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

Enigmatic and extravagant genitalia in the spider genus Mastigusa (Araneae, Cybaeidae) – a taxonomic revision

Filippo Castellucci https://orcid.org/0000-0002-9944-2196 A B * , Andrea Luchetti https://orcid.org/0000-0002-2986-721X A and Nikolaj Scharff B
+ Author Affiliations
- Author Affiliations

A Department of Biological, Geological and Environmental Sciences, University of Bologna, Via Selmi 3, I-40126 Bologna, Italy.

B Zoology Section, Natural History Museum of Denmark, University of Copenhagen, Universitetsparken 15, DK-2100 Copenhagen, Denmark.

* Correspondence to: filippo.castellucci2@unibo.it

Handling Editor: Mark Harvey

Invertebrate Systematics 38, IS23047 https://doi.org/10.1071/IS23047
Submitted: 2 September 2023  Accepted: 21 February 2024  Published: 21 March 2024

© 2024 The Author(s) (or their employer(s)). Published by CSIRO Publishing.

Abstract

Mastigusa is a genus of small palearctic spiders that has recently been moved to the family Cybaeidae after the first inclusion of the genus in a phylogenetic matrix. Three species are currently recognised: M. arietina, M. lucifuga and M. macrophthalma. The status and delimitation, though, has always been problematic due to inconsistency in the characters used to discriminate between these, leading to great confusion in identity and distribution. We present a detailed morphological redescription of the genus and a taxonomic revision of the included species by the combined use of morphological data and molecular species-delimitation techniques based on the mitochondrial COI gene. The status of the three currently described species has been reevaluated and Mastigusa diversa was revalidated based on material from the Iberian Peninsula, North Africa and the United Kingdom. The distribution of Mastigusa species is updated based on novel taxonomic considerations, and comments on the natural history and ecological differences observed in the species are provided.

ZooBank: urn:lsid:zoobank.org:pub:AAD3FAED-440F-4295-B458-455B1D913F81

Keywords: Cybaeidae, DNA barcoding, integrative taxonomy, Mastigusa, morphology, phylogenetics, species delimitation, taxonomic revision.

References

Almquist S (2006) Swedish Araneae, part 2 – families Dictynidae to Salticidae. Insect Systematics & Evolution 63, 285-601.
| Google Scholar |

Azarkina GN, Trilikauskas LA (2012) Spider fauna (Aranei) of the Russian Altai, part I: families Agelenidae, Araneidae, Clubionidae, Corinnidae, Dictynidae and Eresidae. Euroasian Entomological Journal 11, 199-208.
| Google Scholar |

Bidegaray-Batista L, Ferrández MÁ, Arnedo MA (2014) Winter is coming: Miocene and Quaternary climatic shifts shaped the diversification of Western-Mediterranean Harpactocrates (Araneae, Dysderidae) spiders. Cladistics 30, 428-446.
| Crossref | Google Scholar | PubMed |

Bouckaert R, Heled J, Kühnert D, Vaughan T, Wu CH, Xie D, Suchard MA, Rambaut A, Drummond AJ (2014) BEAST 2: a software platform for Bayesian evolutionary analysis. PLoS Computational Biology 10, e1003537.
| Crossref | Google Scholar | PubMed |

Bristowe W (1939) ‘The Comity of Spiders.’ (Ray Society: London, UK)

Castellucci F, Schifani E, Luchetti A, Scharff N (2022) New association between red wood ant species (Formica rufa group) and the myrmecophilic spiders Mastigusa arietina and Thyreosthenius biovatus. Bulletin of Insectology 75, 231-238.
| Google Scholar |

Castellucci F, Scharff N, Luchetti A (2023) Molecular systematics and phylogenetics of the spider genus Mastigusa Menge, 1854 (Araneae, Cybaeidae). Molecular Phylogenetics and Evolution 186, 107833.
| Crossref | Google Scholar | PubMed |

Chyzer C, Kulczynski L (1897) ‘Araneae Hungariae – Zodarioidae, Agelenoidae, Drassoidae, Zoropseoidae, Dysderoidae, Filistatoidae, Calommatoidae, Teraphosoidae.’ (Editio Academiae Scientiarum Hungaricae: Budapest, Hungary, Austro-Hungarian Empire) [In Latin]

Crespo LC, Domènech M, Enguídanos A, Malumbres-Olarte J, Cardoso P, Moya-Laraño J, Frías-López C, Macías-Hernández N, De Mas E, Mazzuca P, Mora E, Opatova V, Planas E, Ribera C, Roca-Cusachs M, Ruiz D, Sousa P, Tonzo V, Arnedo MA (2018) DNA barcode-assisted annotated checklist of the spider (Arachnida, Araneae) communities associated to white oak woodlands in Spanish National Parks. Biodiversity Data Journal 6, e29443.
| Crossref | Google Scholar | PubMed |

Cushing PE (1997) Myrmecomorphy and myrmecophily in spiders: a review. The Florida Entomologist 80, 165-193.
| Crossref | Google Scholar |

Cushing PE (2012) Spider-ant associations: an updated review of myrmecomorphy, myrmecophily, and myrmecophagy in spiders. Psyche 2012, 1-23.
| Crossref | Google Scholar |

Cushing PE, Brückner A, Rogers JW, Horner NV (2022) Trophic specialization of a newly described spider ant symbiont, Myrmecicultor chihuahuensis (Araneae: Myrmecicultoridae). Journal of Arachnology 50, 250-255.
| Crossref | Google Scholar |

Folmer O, Black M, Hoeh W, Lutz R, Vrijenhoek R (1994) DNA primers for amplification of mitochondrial cytochrome c oxidase subunit I from diverse metazoan invertebrates. Molecular Marine Biology and Biotechnology 3, 294-299.
| Google Scholar | PubMed |

Gorneau JA, Crews SC, Cala-Riquelme F, Montana KO, Spagna JC, Ballarin F, Almeida-Silva LM, Esposito LA (2023) Webs of intrigue: museum genomics elucidate relationships of the marronoid spider clade (Araneae). Insect Systematics and Diversity 7, 1-18.
| Crossref | Google Scholar |

Heimer S, Nentwig W (1991) ‘Spinnen Mitteleuropas: Ein Bestimmungsbuch.’ (Paul Parey: Berlin, Germany) [In German]

Jackson AR (1913) On some new and obscure British spiders. Transactions and Annual Report of the “Nottingham Naturalists’ Society” 60, 20-49.
| Google Scholar |

Kalyaanamoorthy S, Minh BQ, Wong TKF, von Haeseler A, Jermiin LS (2017) ModelFinder: fast model selection for accurate phylogenetic estimates. Nature Methods 14, 587-589.
| Crossref | Google Scholar | PubMed |

Kapli P, Lutteropp S, Zhang J, Kobert K, Pavlidis P, Stamatakis A, Flouri T (2017) Multi-rate Poisson tree processes for single-locus species delimitation under maximum likelihood and Markov chain Monte Carlo. Bioinformatics 33(11), 1630-1638.
| Crossref | Google Scholar |

Katoh K, Standley DM (2013) MAFFT multiple sequence alignment software version 7: improvements in performance and usability. Molecular Biology and Evolution 30, 772-780.
| Crossref | Google Scholar | PubMed |

Klausen FE (1974) Spiders new to Norway. Norsk Entomologisk Tidsskrift 21, 191-194.
| Google Scholar |

Kulkarni S, Wood HM, Hormiga G (2023) Advances in the reconstruction of the spider tree of life: a roadmap for spider systematics and comparative studies. Cladistics 39, 479-532.
| Crossref | Google Scholar | PubMed |

Kumar S, Stecher G, Li M, Knyaz C, Tamura K, Battistuzzi FU (2018) MEGA X: molecular evolutionary genetics analysis across computing platforms. Molecular Biology and Evolution 35(6), 1547-1549.
| Crossref | Google Scholar | PubMed |

Lehtinen PT (1967) Classification of the cribellate spiders and some allied families, with notes on the evolution of the suborder Araneomorpha. Annales Zoologici Fennici 4, 199-468.
| Google Scholar |

Lenzini L, Castellucci F, Poso M, Kulczycki A, Simeon E, Greco G, Piccinini A, Legittimo CM (2022) First records of Anagraphis ochracea (Araneae, Gnaphosidae) for continental Italy and Sicily with new observations about its myrmecophilous lifestyle. Arachnologische Mitteilungen 64, 83-92.
| Crossref | Google Scholar |

Locket GH, Millidge AF (1953) ‘British spiders (Vol. 2).’ (Ray Society: London, UK)

Loksa I (1969) Araneae I. In ‘Fauna Hungariae’. Vol. 97, pp. 1–133. (Akadémiai Kiadó: Budapest, Hungary, USSR) [In Hungarian]

Menge A (1854) [Footnotes and taxonomic names.] In ‘Die im Bernstein Befindlichen Organischen Reste der Vorwelt Gesammelt in Verbindung mit Mehreren Bearbeitetet und Herausgegeben’. (Ed. G Berendt) pp. 1–124. (In Commission der Nicolaischen Buchhandlung: Berlin, German Confederation) [In German]

Minh BQ, Nguyen MAT, von Haeseler A (2013) Ultrafast approxim-ation for phylogenetic bootstrap. Molecolar Biology and Evolution 30, 1188-1195.
| Crossref | Google Scholar | PubMed |

Monaghan MT, Wild R, Elliot M, Fujisawa T, Balke M, Inward DJ, Lees DC, Ranaivosolo R, Eggleton P, Barraclough TG, Vogler AP (2009) Accelerated species inventory on Madagascar using coalescent-based models of species delineation. Systematic Biology 58, 298-311.
| Crossref | Google Scholar | PubMed |

Nguyen LT, Schmidt HA, von Haeseler A, Minh BQ (2015) IQ-TREE: a fast and effective stochastic algorithm for estimating maximum-likelihood phylogenies. Molecular Biology and Evolution 32, 268-274.
| Crossref | Google Scholar | PubMed |

Palmgren P (1977) Die Spinnenfauna Finnlands und Ostfennoskandiens. VIII. Argyronetidae, Agelenidae, Hahniidae, Dictynidae, Amaurobiidae, Titanoecidae, Segestriidae, Pholcidae und Sicariidae. Fauna Fennica 30, 1-50 [In German].
| Google Scholar |

Pickard-Cambridge O (1893) On new and rare British spiders. Proceedings of the Dorset Natural History and Antiquarian Field Club 14, 142-164.
| Google Scholar |

Pickard-Cambridge O (1908) On new and rare British Arachnida. Proceedings of the Dorset Natural History and Antiquarian Field Club 28, 121-148.
| Google Scholar |

Pons J, Barraclough TG, Gomez-Zurita J, Cardoso A, Duran DP, Hazell S, Kamoun S, Sumlin WD, Vogler AP (2006) Sequence-based species delimitation for the DNA taxonomy of undescribed insects. Systematic Biology 55(4), 595-609.
| Crossref | Google Scholar |

Puillandre N, Brouillet S, Achaz G (2020) ASAP: assemble species by automatic partitioning. Molecular Ecology Resources 21(2), 609-620.
| Crossref | Google Scholar |

Rambaut A, Drummond AJ, Xie D, Baele G, Suchard MA (2018) Posterior summarization in Bayesian phylogenetics using Tracer 1.7. Systematic Biology 67, 901-904.
| Crossref | Google Scholar | PubMed |

Roberts MJ (1985) ‘The Spiders of Great Britain and Ireland. Vol. 1 – Atypidae to Theridiosomatidae.’ (Harley Books: Colchester, UK).

Roberts MJ (1995) ‘Collins Field Guide – Spiders of Britain and Northern Europe.’ (Collins: London, UK)

Scharff N, Gudik-Sørensen O (2006) Katalog over Danske edderkopper/Catalogue of Danish spiders (Araneae). Entomologiske Meddelelser 74, 3-71 [In Danish].
| Google Scholar |

Simon E (1875) ‘Les arachnides de France.’ Vol. 2, pp. 1–350. (Librarie encyclopédique de Roret: Paris, France) [In French]

Simon E (1898a) Descriptions d’arachnides nouveaux des familles des Agelenidae, Pisauridae, Lycosidae et Oxyopidae. Annales de la Société Entomologique de Belgique 42, 5-34 [In French].
| Google Scholar |

Simon E (1898b) ‘Histoire Naturelle des Araignées. Tome 2.’ pp. 190–380. (Librarie encyclopédique de Roret: Paris, France) [In French].

Simon E (1937) ‘Les arachnides de France. Synopsis générale et catalogue des espèces françaises de l’ordre des Araneae. Tome VI. 5e et derniére partie.’ pp. 979–1298. (Librarie encyclopédique de Roret: Paris, France) [In French]

Stucky BJ (2012) SeqTrace: a graphical tool for rapidly processing DNA sequencing chromatograms. Journal of Biomolecular Techniques 23, 90-93.
| Crossref | Google Scholar | PubMed |

Thorell T (1871) ‘Remarks on synonyms of European spiders. Part II.’ (C. J. Lundström: Uppsala, Sweden)

Tyschchenko VP (1971) ОПРЕДЕЛИТЕЛЬ ПАУКОВ EBPOПEЙСКOЙ ЧАСТП СССР [‘Identification of Spiders of the European part of the USSR.’] (Akademia Nauk SSSR: Leningrad, USSR) [In Russian]

Wasmann E (1894) ‘Kritisches Verzeichniss der myrmecophilin und termitophilen Arthropoden.’ (Felix Dames: Berlin, German Empire). [In German]

Westring N (1861) Araneae svecicae. Göteborgs Kungliga Vetenskaps och Vitterhets Samhälles Handlingar 7, 1-615 [In Latin].
| Google Scholar |

Wheeler WC, Coddington JA, Crowley LM, Dimitrov D, Goloboff PA, Griswold CE, Hormiga G, Prendini L, Ramírez MJ, Sierwald P, Almeida-Silva L, Alvarez-Padilla F, Arnedo MA, Benavides Silva LR, Benjamin SP, Bond JE, Grismado CJ, Hasan E, Hedin M, Izquierdo MA, Labarque FM, Ledford J, Lopardo L, Maddison WP, Miller JA, Piacentini LN, Platnick NI, Polotow D, Silva-Dávila D, Scharff N, Szűts T, Ubick D, Vink CJ, Wood HM, Zhang J (2017) The spider tree of life: phylogeny of Araneae based on target-gene analyses from an extensive taxon sampling. Cladistics 33, 574-616.
| Crossref | Google Scholar | PubMed |

Wunderlich J (1986) ‘Spinnenfauna gestern und heute: Fossile Spinnen in Bernstein und ihre heute lebenden Verwandten.’ (Quelle & Meyer: Weisbaden, West Germany). [In German]

Wunderlich J (1994) Über “Ameisenspinnen” in Mitteleuropa (Arachnida: Araneae). Beiträge zur Araneologie 33, 447-470 [In German].
| Google Scholar |

Wunderlich J (2004) Fossil spiders (Araneae) of the family Dictynidae s. l., including Cryphoecinae and Hahniidae in Baltic and Dominican amber and copal from Madagascar, and on selected extant Holarctic taxa, with new descriptions and diagnoses. Beiträge zur Araneologie 50, 1380-1483.
| Google Scholar |

Zhang J, Madden TL (1997) PowerBLAST: a new network BLAST application for interactive or automated sequence analysis and annotation. Genome Resources 7, 649-656.
| Crossref | Google Scholar | PubMed |

Zhang J, Kapli P, Pavlidis P, Stamatakis A (2013) A general species delimitation method with applications to phylogenetic placements. Bioinformatics 29(22), 2869-2876.
| Crossref | Google Scholar |