Accepted by T. Wesener: 20 Mar. 2018; published: 17 May 2018
ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN 1175-5334 (online edition)
Copyright © 2018 Magnolia Press
Zootaxa 4420 (3): 415–429
http://www.mapress.com/j/zt/
Article
415
https://doi.org/10.11646/zootaxa.4420.3.7
http://zoobank.org/urn:lsid:zoobank.org:pub:5F0337E2-CBFE-413E-BDFF-3D6131BE1609
Out of the dark void: Ommatoiulus longicornis n. sp., a new julid from Spain
(Diplopoda, Julida) with notes on some troglobiomorphic traits in millipedes
NESRINE AKKARI
1
, JOSÉ D. GILGADO
2,3
, VICENTE M. ORTUÑO
3
& HENRIK ENGHOFF
4
1
Naturhistorisches Museum Wien, Burgring 7, 1010 Wien, Austria. E-mail: nesrine.akkari@nhm-wien.ac.at
2
Section of Conservation Biology, Department of Environmental Sciences, University of Basel. St. Johanns-Vorstadt 10, CH-4056,
Basel, Switzerland. E-mail: josedomingo.gilgadohormaechea@unibas.ch
3
Grupo de Investigación de Biología del Suelo y de los Ecosistemas Subterráneos. Departamento de Ciencias de la Vida. Facultad de
Biología, Ciencias Ambientales y Química. Universidad de Alcalá. A.P. 20 Campus Universitario. E-28805. – Alcalá de Henares,
Madrid, Spain. E-mail: vicente.ortuno@uah.es
4
Natural History Museum of Denmark, University of Copenhagen, Universitetsparken 15, DK-2100 København Ø, Denmark.
E-mail: henghoff@snm.ku.dk
Abstract
A new millipede species, Ommatoiulus longicornis n. sp. is described from the subterranean habitat in eastern Spain, and
the elongation of its antennae is discussed in relation to the habitat where it dwells. We take the occasion to discuss the
elongation of appendages as a troglobiomorphic character in millipedes and shed the light on the various, sometimes con-
troversial, hypotheses that could have led to these modifications in millipedes and other subterranean arthropods.
Key words: Schizophyllini, Spain, taxonomy, new species, troglomorphism
Introduction
The fauna of the subterranean environment is classified, according to its degree of dependence, as troglobionts
(true hypogean), troglophiles (facultative hypogean) and trogloxenes (accidentally found in the hypogean
environment). A detailed explanation of these terms can be found in Sket (2008). Here, we take the occasion to
discuss the concept of 'troglomorphism' or more correctly 'troglobiomorphism', which refers to convergent
morphological traits observed in many (but not all) troglobionts as well as in some troglophiles (Christiansen 1962;
Juberthie & Decu 1994; Desutter-Grandcolas 1997; Hernando et al. 1999; Dethier & Hubart 2005; Pipan & Culver
2012; Culver & Pipan 2015).
The term troglobiomorphism refers to the fact that “cave” animals have similar traits (troglobiomorphies) due
to convergent selective pressures in the hypogean environment (Culver & Pipan 2015). However, this is not as
simple, as not all troglobionts are troglobiomorphic and some troglobiomorphies are only observable in certain taxa
(see list of troglobiomorphies in Christiansen 2012). Several troglobiomorphies might also be expressions of
previous adaptations to other environments, i.e. exaptations to the hypogean environment (Hernando et al. 1999;
Heads 2010; Gilgado et al. 2014).
It was recently demonstrated that the subterranean realm is composed of different habitats, which leads to a
divergent evolution in some characters (Pipan & Culver 2012, Trontelj et al. 2012; Culver & Pipan 2015), leading
to the existence of specialized troglobionts with different sizes and degrees of appendage elongation. According to
Culver & Pipan (2015), this leads to a contradiction within the troglobiomorphism hypothesis and makes it
necessary to refer to these morphological traits as convergences, or divergences, rather than uncritically classifying
them as troglobiomorphies.
There are several types of subterranean environments with various characteristics (Culver & Pipan 2009, 2014;
Ortuño et al. 2014). The deep hypogean environment is traditionally understood as constituted by bigger caves and
the fissure network of the bedrock. The latter was reported by Racovitza (1907) as the true habitat of many