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Phylogenetic Analysis of Phalangida (Arachnida, Opiliones) Using Two Nuclear Protein-Encoding Genes Supports Monophyly of Palpatores

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Phylogenetic Analysis of Phalangida (Arachnida, Opiliones) Using Two Nuclear Protein-Encoding Genes Supports Monophyly of Palpatores 2001. The Journal of Arachnology 29:189±200 PHYLOGENETIC ANALYSIS OF PHALANGIDA (ARACHNIDA, OPILIONES) USING TWO NUCLEAR PROTEIN-ENCODING GENES SUPPORTS MONOPHYLY OF PALPATORES Jeffrey W. Shultz: Department of Entomology, University of Maryland, College Park, Maryland 20742 USA Jerome C. Regier: Center for Agricultural Biotechnology, University of Maryland Biotechnology Institute, College Park, Maryland 20742 USA ABSTRACT. Recent phylogenetic studies of Opiliones have shown that Cyphophthalmi and Phalangida (5 Palpatores 1 Laniatores) are sister groups, but higher relationships within Phalangida remain contro- versial. Current debate focuses on whether Palpatores (5 Caddoidea 1 Phalangioidea 1 Ischyropsalidoidea 1 Troguloidea) is monophyletic or paraphyletic, with Ischyropsalidoidea 1 Troguloidea (5 Dyspnoi) being more closely related to Laniatores. The latter hypothesis was favored in recent combined studies of ri- bosomal DNA and morphology. Here higher relationships within Phalangida are examined using two nuclear protein-encoding genes, elongation factor-1a (EF-1a) and RNA polymerase II (Pol II), from 27 opilion species representing seven superfamilies. Cyphophthalmi was used as the outgroup. Nucleotide and inferred amino acid sequences were analyzed using maximum-parsimony and maximum-likelihood methods. All analyses recovered Palpatores as the monophyletic sister group to Laniatores with moderate to strong empirical support. Most palpatorean superfamilies were also recovered, but relationships among them were ambiguous or weakly supported. A monophyletic Palpatores was also obtained when EF-1a and Pol II sequences were analyzed together with 18S and 28S rDNA sequences. Keywords: Molecular systematics, elongation factor-1a, RNA polymerase II, Opiliones Members of the arachnid order Opiliones The goal of the present study is to reconstruct (harvestmen, shepherd spiders, daddy long- phylogenetic relationships among major line- legs, etc.) are abundant and often highly vis- ages within Opiliones using nucleotide and in- ible members of many terrestrial ecosystems. ferred amino-acid sequences from two nuclear The group is estimated to encompass about protein-encoding genes, elongation factor-1a 5000 species (Shear 1982). The basic biology (EF-1a) and the largest subunit of RNA poly- of most opilions is unexplored; but the order merase II (Pol II). is known to encompass substantial behavioral The monophyly of Opiliones is well estab- and morphological diversity, with much recent lished (Weygoldt & Paulus 1979; Shultz work focusing on the structure and evolution 1990), and three principal opilion groups are of mating systems (e.g., MacõÂas-OrdoÂnÄez widely recognized, namely, Cyphophthalmi, 1997; Martens 1993; Mora 1987, 1990, 1991; Palpatores and Laniatores (Hansen & Sùren- Ramires & Giaretta 1994) and the evolution- sen 1904; Shear 1982). Cyphophthalmi (5 Si- ary morphology of genitalia (e.g., Martens ronoidea) is a group of tiny (, 5 mm), hard- 1976, 1980, 1986; Martens, Hoheisel & GoÈtze bodied and somewhat mite-like opilions 1981; Hoheisel 1980; Shultz 1998). An un- characterized by many unique traits, such as derstanding of the phylogenetic relationships a short male genital organ (spermatopositor), within Opiliones, and between Opiliones and elevated ozopores (ozophores), a specialized other arachnid orders, is pivotal to further pro- spine on leg IV of males (adenostyle) and ab- gress in these and other ®elds of arachnolog- sence of a genital operculum (Shear 1982; ical research. However, many relationships are Shultz 1998). Laniatores is a species-rich unclear and the focus of ongoing investigation group of heavily sclerotized opilions that have and debate (e.g., Giribet et al. 1995, 1999; Gi- radiated extensively in the neotropics and ribet & Wheeler 1999; Shultz 1998, 2000). southeastern Asia (Shear 1982). Members of 189 190 THE JOURNAL OF ARACHNOLOGY this undisputedly monophyletic group are gida). Phalangida has also been supported by characterized by large, often raptorial palps, sperm ultrastructure (Juberthie & Manier unsegmented ovipositors and other unique 1978), arrangement of extrinsic pharyngeal features. Palpatores is a morphologically di- muscles (Shultz 2000) and, especially, molec- verse assemblage of four superfamilies (Cad- ular sequence data. Giribet et al. (1999) using doidea, Ischyropsalidoidea, Phalangioidea, 18S and 28S ribosomal DNA and Shultz & Troguloidea) united by few morphological Regier (unpublished) using EF-1a and Pol II synapomorphies. Caddoidea and Phalangioi- amino acid sequences have recovered both dea include the typical ``daddy longlegs'' fa- Cyphophthalmi and Phalangida as monophy- miliar to inhabitants of northern temperate re- letic sister clades. gions and are characterized, in part, by Recent morphological and molecular anal- well-developed coxal lobes (coxapophyses) yses appear to have established Phalangida as and segmented ovipositors. Ischyropsalidoi- a monophyletic group, but there is debate con- dea and Troguloidea are less familiar groups cerning basal relationships within this group. but encompass a substantial range of morpho- These opilions are traditionally divided into logical diversity. They are characterized, in the Palpatores and Laniatores, with Palpatores part, by reduced or absent coxapophyses, di- often divided into Dyspnoi (5 Ischyropsali- aphanous cheliceral teeth and unsegmented doidea 1 Troguloidea) and Eupnoi (5 Cad- ovipositors. doidea 1 Phalangioidea) (Hansen & Sùrensen Opilion systematists have long debated the 1904; SÏilhavy 1961; Juberthie & Manier phylogenetic relationships of the three prin- 1978; Shultz 1998). However, Giribet et al. cipal opilion lineages. SÏilhavy (1961) placed (1999) conducted a phylogenetic analysis of Cyphophthalmi and Palpatores together as the Opiliones using 18S and 28S ribosomal DNA sister group to Laniatores on the basis of the sequences and recovered two topologies, one arrangement of tarsal claws; that is, claws of favoring a monophyletic Palpatores and one all legs are similar in Cyphophthalmi and Pal- favoring a paraphyletic Palpatores, with Dy- patores, but those of legs I and II differ from spnoi being the sister group to Laniatores. The those of legs III and IV in Laniatores. A sim- molecular characters alone did not strongly fa- ilar but more well-defended system was pro- vor one hypothesis over the other, but a mor- posed by Martens and co-workers (Martens phology-based analysis and the combined mo- 1976, 1980, 1986; Hoheisel 1980; Martens et lecular-morphological study tended to support al. 1981). They also united Cyphophthalmi the Dyspnoi 1 Laniatores hypothesis. In a and Palpatores (5 Cyphopalpatores) but ar- subsequent study, Giribet & Wheeler (1999) gued that the palpatorean superfamily Trogu- explored the signi®cance of indels on the phy- loidea was the sister group to a clade com- logenetic analysis of Opiliones. Using 18S ri- prising Cyphophthalmi and the remaining bosomal DNA sequences and morphological Palpatores, a system that rendered Palpatores characters from Giribet et al. (1999), Giribet a paraphyletic group. Their Cyphopalpatores & Wheeler recovered Dyspnoi 1 Laniatores. hypothesis was based largely on original mor- However, given their exclusion of the 28S phological studies of the ovipositor and penis, rDNA and use of a problematic morphology character systems that are uniquely derived in matrix (see Methods for details), it is probably Opiliones. Consequently, the root and branch- best to regard Giribet & Wheeler's contribu- ing pattern of the tree proposed by Martens et tion as a demonstration of a new analytical al. was based on speculative character trans- method rather than a study of opilion relation- formation series rather than inferences derived ships. from outgroups. Shultz (1998) assessed the The present study examines higher relation- Cyphopalpatores concept through a parsimo- ships within Phalangida using nucleotide and ny-based morphological analysis of genitalic amino-acid sequences from EF-1a and Pol II and non-genitalic characters using a generic with the speci®c aim of testing the Dyspnoi taxon sample similar to that of Martens et al. 1 Laniatores hypothesis. Sequences were ob- and outgroups (i.e., scorpions and xiphosu- tained from 27 opilion taxa representing the rans) to polarize non-genitalic characters. The major opilion superfamilies (i.e., Sironoidea, results supported Cyphophthalmi as the sister Travunioidea, Gonyleptoidea, Phalangioidea, group to Laniatores 1 Palpatores (5 Phalan- Caddoidea, Ischyropsalidoidea and Troguloi- SHULTZ & REGIERÐMOLECULAR SYSTEMATICS OF OPILIONES 191 dea). Maximum-parsimony and maximum- parentheses: Sironoidea: Parasiro coiffaiti likelihood analyses strongly and consistently Juberthie 1956 (Sironidae) (AF240852; recovered Palpatores and Laniatores as mono- AF241025 - AF241027), Siro acaroides (Ew- phyletic sister groups under all character par- ing 1923) (Sironidae) (AF240855; AF241034 titions, weighting schemes and analytical - AF241036). Travunioidea: Equitius doriae methods. This strong support for the mono- Simon 1880 (Triaenonychidae) (AF240867; phyly of Palpatores contrasts with the ambig- AF241068 - AF241070), Sclerobunus robus- uous or weakly supported conclusions derived tus (Packard 1877) (Triaenonychidae) from studies of 18S and 28S ribosomal DNA (AF240858; AF241042 - AF241044). Gony- and morphology (Giribet et al. 1999; Giribet leptoidea: Gonyleptes fragilis Mello-LeitaÄo & Wheeler 1999) or morphology alone 1922 (Gonyleptidae:
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