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

Revision of the genus Trogulus Latreille: the Trogulus coriziformis species-group of the western Mediterranean (Opiliones : Trogulidae)

Axel L. Schönhofer A B and Jochen Martens A
+ Author Affiliations
- Author Affiliations

A Institut of Zoology, Johannes Gutenberg-University Mainz, Department of Systematic Zoology, Müllerweg 6, D-55099 Mainz, Germany.

B Corresponding author. Email: axel.schoenhofer@uni-mainz.de

Invertebrate Systematics 22(5) 523-554 https://doi.org/10.1071/IS08013
Submitted: 29 March 2008  Accepted: 13 October 2008   Published: 4 December 2008

Abstract

Within the well researched European fauna of harvestmen, the genus Trogulus Latreille exhibits unexpectedly high cryptic diversity. The species’ uniform morphology hinders an exclusively morphological approach to their systematics and taxonomy, and a preliminary molecular study estimated the number of species to be three times higher than currently known. The current study focuses on a clearly defined species-group within Trogulus, combining molecular (~1700 bp 28S rRNA and the cytochrome b gene), distributional, morphometric and morphological data. Relationships are reconstructed using Bayesian inference, maximum parsimony and maximum likelihood and this information is subsequently used to evaluate morphological characters for systematic usability and to identify biogeographical processes leading to speciation events. The Trogulus coriziformis species-group is defined and diagnosed and includes eight species. Three species are redefined: T. coriziformis C. L. Koch, 1839, for which a neotype is designated, and T. aquaticus Simon, 1879 and T. cristatus Simon, 1879 for which lectotypes are designated. Four species are described as new: T. balearicus, sp. nov. from the Balearic Islands, T. huberi, sp. nov. from southern Portugal, T. prietoi, sp. nov. from Andalusia, Spain, and T. pyrenaicus, sp. nov. from the central Pyrenees. Trogulus lusitanicus Giltay, 1931 is used as collective name and probably refers to a composite of species presently difficult to tell apart. Trogulus salfii De Lerma, 1948 is proposed as a synonym for T. coriziformis. Within Trogulus, the molecular genetic data support monophyly and basal placement of the Trogulus coriziformis species-group. The species to differ in external morphology (size, papillation of palps, apophyses of legs, pattern of body papillation, morphometric data), 28S and cytochrome b autapomorphies and to a lesser degree by male genital morphology. The species-group is confined to the western Mediterranean area and its species are allopatrically distributed. Their present distribution corresponds to geological processes in the Miocene and Pliocene indicating that this group of organisms may be of considerable value for further biogeographic studies.


Acknowledgements

We gratefully acknowledge loans of and donations from the following colleagues: Carlos Prieto (Universidad del Pais Vasco, Bilbao), Siegfried Huber (Unteruhldingen, Germany), Joerg Spelda (Museum Mensch und Natur, Munich), Pedro Cardoso (University of Azores, Ponta Delgada), Ambros Hänggi (Naturhistorisches Museum Basel), Paolo Pantini (Museo Civico di Scienze Naturali, Bergamo), Georges Wauthy (Belgian Royal Institute of Natural Sciences, Brussels), Janet Beccaloni (Natural History Museum, London, Great Britain), Jürgen Gruber (Naturhistorisches Museum, Wien, Austria), Jason Dunlop (Naturhistorisches Forschungsinstitut, Museum für Naturkunde, Berlin, Germany), Nikolaj Scharff (Zoological Museum, University of Copenhagen), Marjan Komnenov (Skopje, Republic of Macedonia), Ivo Karaman (Dep. of Biology and Ecology, Novi Sad University, Serbia), Peter Schwendinger (Muséum d’histoire naturelle, Genève, Switzerland), Arturo Muñoz-Cuevas (Muséum National d’Histoire Naturelle, Paris), Christa Deeleman-Reinhold (Ossendrecht), Peter Jäger (Naturmuseum und Forschungsinstitut Senckenberg, Frankfurt am Main, Germany), Roman Ozimec (Zagreb, Croatia). J. Gruber and P. Schwendinger provided helpful information on collection under their care. Marshal Hedin (San Diego State University) sent helpful information on primers for 28S rRNA sequencing. The German Academic Exchange Service (Deutscher Akademischer Austauschdienst, Bonn) supported A. L. S. with a 2.5 month travel grant. The ‘Landesgraduiertenförderung Rheinland-Pfalz’ provided a two year grant to A. L. S. to work on his PhD dissertation. ‘Wagner-Stiftung’ and ‘Feldbausch-Stiftung’, both at Fachbereich Biologie, Mainz University, provided annual travel grants for field work to J. M. To our knowledge all specimens examined were collected according to current regulations.


References


Avram S. (1971) Quelques espéces nouvelles ou connues du genre Trogulus Latr. (Opiliones). Travaux de l’institut de Spéologie “Émile Racovitza” 10, 245–272. open url image1

Boyer S. L., Clouse R., Benavides L. R., Sharma P., Schwendinger P. J., Kuranarathna I., Giribet G. (2007) Biogeography of the world: a case study from cyphophthalmid Opiliones, a globally distributed group of arachnids. Journal of Biogeography 34, 2070–2085.
Crossref | GoogleScholarGoogle Scholar | open url image1

Boyer S. L., Karaman I., Giribet G. (2005) The genus Cyphophthalmus (Arachnida, Opiliones, Cyphophthalmi) in Europe: a phylogenetic approach to Balkan Peninsula biogeography. Molecular Phylogenetics and Evolution 36, 554–567.
Crossref | GoogleScholarGoogle Scholar | PubMed | PubMed | open url image1

Brignoli P. M. (1968) Note su Sironidae, Phalangodidae e Trogulidae italiani, cavernicoli ed endogei (Opiliones). Fragmenta Entomologica 5(3), 259–293. open url image1

Brignoli P. M., Rafaelli E. (1978) Nuovi dati e problemi aperti su alcuni Opilioni Italiani (Arachnida, Opiliones). Bollettino della Societa Entomologica Italiana 110(4–6), 86–99. open url image1

Canestrini G. (1875) Intorno ai Chernetidi ed Opilionidi della Calabria. Atti della Accademia Scientifica Veneto-Trentino-Istriana 4, 1–12. open url image1

Cardoso P., Scharff N., Gaspar C., Henriques S. S., Carvalho R. , et al. () Rapid biodiversity assessment of spiders (Araneae) using semi-quantitative sampling: a case study in a Mediterranean forest. Insect Conservation and Diversity In press , open url image1

Chemini C. (1983) Trogulus martensi n. sp. from northern Italy (Arachnida, Opiliones). Bollettino della Societa Entomologica Italiana 115(8–10), 125–129. open url image1

Chemini C., Martens J. (1988) Trogulus cisalpinus n. sp. from the Italian Alps (Arachnida: Opiliones: Trogulidae). Mitteilungen aus dem Zoologischen Museum in Berlin 64(1), 71–81. open url image1

Cunningham C. W. (1997) Is congruence between data partitions a reliable predictor of phylogenetic accuracy? Empirically testing an iterative procedure for choosing among phylogenetic methods. Systematic Biology 46, 464–478.
CAS | CAS | Crossref | Crossref | PubMed | PubMed |
open url image1

Dahl F. (1903). Eine eigenartige Metamorphose der Troguliden, eine Verwandlung von Amopaum in Dicranolasma und von Metopoctea in Trogulus. Sitzungsbericht der Gesellschaft naturforschender Freunde Berlin, 278–292.

De Jong H. (1998) In search of historical biogeographic patterns in the western Mediterranean terrestrial fauna. Biological Journal of the Linnean Society 65, 99–164.
Crossref | Crossref |
open url image1

De Lerma B. (1948) Opilionidi cavernicoli della Campania. Studi Speleologici e Faunistici sull Italia Meridionale. Supplemento al Bollettino della Società dei Naturaisti in Napoli 57(9), 160–164. open url image1

Di Caporiacco L. (1949) Un nuova opilione e qualche altro Arachnide delle Alpi Orobie. Atti del museo civico di Storia naturale Trieste 17, 120–121. open url image1

Farris J. S., Kallersjo M., Kluge A. G., Bult C. (1994) Testing significance of congruence. Cladistics 10, 315–320.
Crossref | GoogleScholarGoogle Scholar | open url image1

Felsenstein J. (1985) Confidence limits on phylogenies: an approach using the bootstrap. Evolution 39, 783–791.
Crossref | GoogleScholarGoogle Scholar | open url image1

Giltay L. (1931) Note sur une espèce nouvelle de Trogulus du Portugal. Bulletin du Musee Royal d’Historie naturelle de Belgique 7(27), 1–2. open url image1

Gozo A. (1908) Gli Aracnidi di caverne italiane. Bollettino della Società entomologica italiana, Genova 38, 109–139. open url image1

Gruber J. (1968) Ergebnisse zoologischer Sammelreisen in der Türkei: Calathocratus beieri, ein neuer Trogulide aus Anatolien (Opiliones, Arachnida). Annalen des Naturhistorischen Museums in Wien 72, 435–441. open url image1

Gruber J. (1969) Ergebnisse des österreichisch-türkischen Anatolien Expeditionen 9. – Weberknechte der Familien Sironidae und Trogulidae aus der Türkei (Opiliones, Arachnida). Review of the Faculty of Science University of Istanbul 34(1/2), 75–88. open url image1

Huelsenbeck J. P., Ronquist F. R. (2001) MrBayes: Bayesian inference of phylogenetic trees. Bioinformatics 17, 754–755.
Crossref | GoogleScholarGoogle Scholar | CAS | CAS | PubMed | PubMed | open url image1

Karaman I. M. ( ) The taxonomical status and diversity of Balkan Sironids (Opiliones, Cyphophthalmi) with description of twelve new species. Zoological Journal of the Linnean Society, London In press , open url image1

Koch C. L. (1839). Die Arachniden. Getreu nach der Natur abgebildet und beschrieben. In: C. W. Hahn and C. L. Koch, Vol. 5, C. H. Zeh’sche Buchhandlung, Nürnberg.

Komposch C. (2000) Trogulus falcipenis, spec. nov., ein neuer Brettkanker aus den Alpen und dem Dinarischen Gebirge. Spixiana 23(1), 1–14. open url image1

Kraus O. (1959) Weberknechte aus Spanien (Arachn., Opiliones). Mitteilungen aus dem Zoologischen Museum in Berlin 35(2), 293–304. open url image1

Kraus O. (1961) Die Weberknechte der Iberischen Halbinsel (Arach., Opiliones). Senckenbergiana Biologica 42(4), 331–363. open url image1

Mallatt J., Giribet G. (2006) Further use of nearly complete 28S and 18S rRNA genes to classify Ecdysozoa: 37 more arthropods and a kinorhynch. Molecular Phylogenetics and Evolution 40, 772–794.
Crossref | GoogleScholarGoogle Scholar | CAS | CAS | PubMed | PubMed | open url image1

Mallatt J., Sullivan J. (1998) 28S and 18S rDNA sequences support the monophyly of lampreys and hagfishes. Molecular Biology and Evolution 15, 1706–1718.
CAS | CAS | PubMed | PubMed |
open url image1

Mallatt J. M., Garey J. R., Shultz J. W. (2004) Ecdysozoan phylogeny and Bayesian inference: first use of nearly complete 29S and 18S rRNA gene sequences to classify the arthropods and their kin. Molecular Phylogenetics and Evolution 31, 178–191.
Crossref | GoogleScholarGoogle Scholar | CAS | PubMed | PubMed | open url image1

Marcellino I. (1965) Su alcuni Trogulidae (Arachnida, Oipliones) della Sicilia e dell Appenino centrale. Bollettino delle sedute della Accademia Gioenia di Scienze Naturali in Catania 4(8/5), 323–340. open url image1

Marcellino I. (1970) Opilioni delle Alpi Apuane. Lavori della Società Italiana di Biogeografia 1, 363–389. open url image1

Marcellino I. (1971) Opilioni dell’ Appennino Centrale. Lavori della Società Italiana di Biogeografia 2, 401–422. open url image1

Marcellino I. (1975) Opilioni (Arachnida) delle alpi occidentali. Bollettino del Museo Civico di Storia Naturale di Verona 2, 119–144. open url image1

Marcellino I. (1982) Opilioni cavernicoli italiani. Lavori della Società Italiana di Biogeografia 7, 33–52. open url image1

Martens J. (1978). ‘Spinnentiere, Arachnida: Weberknechte, Opiliones.’ (Die Tierwelt Deutschlands 64. G. Fischer, Jena.)

Martens J., Chemini C. (1988) Die Gattung Anelasmocephalus Simon, 1879. Biogeographie, Artgrenzen und Biospezies-Konzept (Opiliones: Trogulidae). Zoologische Jahrbucher. Abteilung fur Systematik, Geographie und Biologie der Tiere 115, 1–48. open url image1

Mello-Leitão C. (1936) Les Opilions de Catalogne. Treballs del Museu de Ciències Naturals de Barcelona (sér. entom.) 11(9), 1–18. open url image1

Novak T. (2005) An overview of harvestmen (Arachnida: Opiliones) in Bosnia and Herzegovina. Natura Croatica 14(4), 301–350. open url image1

Passamaneck Y., Halanych K. M. (2006) Lophotrochozoan phylogeny assessed with LSU and SSU data: evidence of lophophorate polyphyly. Molecular Phylogenetics and Evolution 40(1), 20–28.
Crossref | GoogleScholarGoogle Scholar | CAS | PubMed | PubMed | open url image1

Posada D., Buckley T. (2004) Model selection and model averaging in phylogenetics: advantages of Akaike information criterion and Bayesian approaches over likelihood ratio tests. Systematic Biology 53, 793–808.
Crossref | GoogleScholarGoogle Scholar | PubMed | PubMed | open url image1

Posada D., Crandall K. A. (1998) Modeltest: Testing the model of DNA substitution. Bioinformatics (Oxford, England) 14, 817–818.
Crossref | GoogleScholarGoogle Scholar | CAS | PubMed | PubMed | open url image1

Prieto C. (2003) Primera Actualización de la check-list de los Opiliones de la Península Ibérica e Islas Baleares. Revista Iberica de Arachnologia 8, 125–141. open url image1

Rambla M. (1960) Contributión al estudio de los Opiliones de la Fauna Ibérica. 1a. Nota sobre Opiliones de Andalucía. Publicaciones del Instituto de Biologia Aplicada Barcelona 31, 5–16. open url image1

Rambla M. (1967) Opiliones de Portugal. Revista de Biologia, Lisboa 6(1/2), 1–34. open url image1

Rambla M. (1972). Opiliones (Arachn.) de las Baleares. Rapport du congrès de la CIESM – Commission Internationale pour l’Exploration Scientifique de la Mer Méditerranée 21(3), 89–92.

Roewer C. F. (1923). ‘Die Weberknechte der Erde. Systematische Bearbeitung der bisher bekannten Opiliones.’ (Gustav Fischer, Jena, Germany.)

Roewer C. F. (1925) Opilioniden aus dem nördlichen und östlichen Spanien, gesammelt von Dr. F. Haas in den Jahren 1914–19. Senckenbergiana 7(5), 177–179. open url image1

Roewer C. F. (1950) Über Ischyropsalididae und Trogulidae. Weitere Weberknechte XV. Senckenbergiana 31(1/2), 11–56. open url image1

Rosenbaum G., Lister G. S., Duboz C. (2002) Reconstruction of the tectonic evolution of the western Mediterranean since the Oligocene. Journal of the Virtual Explorer 8, 107–130. open url image1

Saitou N., Nei M. (1987) The neighbor-joining method: A new method for reconstructing phylogenetic trees. Molecular Biology and Evolution 4, 406–425.
CAS | PubMed | PubMed |
open url image1

Schmidt H. A., Strimmer K., Vingron M., von Haeseler A. (2002) Tree-Puzzle: maximum likelihood phylogenetic analysis using quartets and parallel computing. Bioinformatics 18, 502–504.
Crossref | GoogleScholarGoogle Scholar | CAS | PubMed | PubMed | open url image1

Schönhofer A. L. , and Martens J. (in press). Revision of the genus Trogulus Latreille: the Trogulus hirtus species-group (Opiliones: Trogulidae). In ‘Towards a Natural History of Arthropods and Other Organisms – in memoriam Konrad Thaler’. (Eds C. Kropf and P. Horak.) Contributions to Natural History 12, Bern.

Sharma P., Karunarathna I., Giribet G. ( ) On the endemic Sri Lankan genus Pettalus (Opiliones, Cyphophthalmi, Pettalidae) with a description of a new species and a discussion of its diversity. Journal of Arachnology , open url image1

Šilhavý V. (1967) Anarthrotarsus martensi, ein neuer Weberknecht aus Griechenland (Arach., Opiliones). Senckenbergiana Biologica 48(3), 175–178. open url image1

Šilhavý V. (1968) Trogulus rossicus sp. nov., ein neuer Weberknecht aus der UdSSR (Arachnida, Opiliones). Reichenbachia 10(5), 25–29. open url image1

Šilhavý V. (1969) Gli Opilionidi dell’Italia settentrionale e centrale. Atti dell’Istituto Veneto di Scienze, Lettere ed Arti. Classe di Scienze Fisiche. Matematiche e Naturali 127, 495–498. open url image1

Simon E. (1879). ‘Les Arachnides de France. Contenant les ordres des Chernetes, Scorpiones et Opiliones.’ (Librairie Encyclopédique de Roret 7, Paris.)

Simon E. (1882) Aracnidi raccolti da G. Cavanna al Vulture, al Pollino ed in altri luoghi dell’Italia meridionale e centrale, nei mesi di luglio ed agosto 1880. Bullettino della Società entomologica italiana 14, 31–48. open url image1

Swofford D. L. (2001). PAUP*: Phylogenetic analysis using parsimony and other methods, Version 4.0b8a. Sinauer Associates, Sunderland/Massachusetts.

Trossarelli F. (1934) Elenco degli Opilionidi italiani del Museo di Milano. Atti della società italiana di scienze naturali di Milano 73, 19–24. open url image1

Winchell C. J., Sullivan J., Cameron C. B., Swalla B. J., Mallatt J. (2002) Evaluating hypotheses of deuterostome phylogeny and chordate evolution with new LSU and SSU ribosomal DNA data. Molecular Biology and Evolution 19, 762.
CAS | PubMed | PubMed |
open url image1

Zangheri P. (1966) Repertorio sistematico e topografico della flora e fauna vivente e fossile della Romagna. Memorie Fuori, Serie No. 1. Museo Civico di Storia Naturale di Verona 2, 485–854. open url image1