Kury, Adriano B., 2015, Opiliones are no longer the same—on suprafamilial groups in harvestmen (Arthropoda: Arachnida), Zootaxa 3925 (3), pp. 301-340: 316-323

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Phylogeny of Laniatores  

The armored harvestmen ( Laniatores   ) are here presented in more detail, separated from the rest of the Opiliones   . In Figs. 12 View FIGURE 12 to 19, the relevant hypotheses found in the literature concerning the branching pattern of the Laniatores   are presented. The six following genera are here used to represent the major groups of Laniatores   as currently understood: Erebomaster   (for the Travunioidea s.s.), Equitius   (for the Triaenonychoidea), Synthetonychia   (for the Synthetonychiidae   ), Gnomulus   (for the Sandokanidae   / Oncopodidae   , consistently deemed to hold a special position), Phalangodes   and Gonyleptes   . See Table 4 for details.

The monophyly of this suborder has not been challenged since Perty (1833), who ranked the Cosmetidae   together with the Eupnoi.

Hypothesis L 1 ( Fig. 12 View FIGURE 12 ) was prevalent by the end of 19 th century. The young Triaenonychidae   and Oncopodidae   did not have yet a separate placement, and the Cladonychiidae   were not yet recognized, its species being merged along with Oncopodidae   in the Phalangodidae   .

Hypothesis L 2 ( Fig. 13 View FIGURE 13 ) represented a major step forward, when Loman (1901; 1902) separated the Triaenonychidae   as “sister” (a coordinate group) to the other Laniatores   , calling it suborder Insidiatores. An expanded usage of the Insidiatores was later recovered by Kury (2003), also including Erebomaster   , although formally this is not Loman’s original concept. Pocock (1902), in spite of his critics, followed this arrangement. Loman (1901) also separated Gnomulus   from the Phalangodidae   .

Hypothesis L 3 ( Fig. 14 View FIGURE 14 ) was a further refinement by Loman (1903), who created the names Sterrhonoti (for the Sandokanidae   , then called Oncopodidae   ) and Camptonoti (for the rest of his Laniatores   ). Setting the Sandokanidae   apart proved to be very popular, persisting in the literature for many decades. In the 1920 s and early 1930 s, a subdivision of the Laniatores   became out of fashion. The families were simply listed as coordinate categories (e.g., Roewer 1923).

Hypothesis L 4 ( Fig. 15 View FIGURE 15 ): a major breakthrough happened in the mid- 1930 s, when Hadži (1935) recognized the close relationship between the southern hemisphere Triaenonychidae   and the European Cladonychiinae. This group was later resurrected as Travunioidea ( Erebomaster   + Equitius   + Synthetonychia   ) by Kratochvíl (1958) and as an “expanded” Insidiatores by Kury (2003), but it is currently widely regarded as a paraphylum. Kratochvíl (1958) put all remaining Laniatores   as a monophylum (a hitherto unchallenged hypothesis, which he called “Oncopodoidea” and was later named Grassatores by Kury). Kratochvíl also combined Hadži’s group with Mello- Leitão’s special placement for Oncopodidae   as sister group of the other Grassatores. Martens (1980) conserved exactly this arrangement, which also appeared in the combined molecular + morphological analyses by Giribet et al. (1999; 2002). This hypothesis was in the second half of the 20 th century and at the turn of 21 st century.

Hypothesis L 5 ( Fig. 16 View FIGURE 16 ): Mello-Leitão (1944) led to an extreme version of Loman’s (1903) attempt to segregate the Oncopodidae   and put it at the base of his tree of “oculariate” Laniatores   . Hadži’s component Erebomaster   + Equitius   is ever present, although nested a node above. Šilhavý (1961), who seemed to hold Mello- Leitão’s views in high regard (see, for instance, Šilhavý 1973), followed this scheme, naming the Sandokanidae   (then Oncopodidae   ) as Oncopodomorphi and all non-oncopodid Laniatores   as Gonyleptomorphi. Bristowe (1976) did the same, separating the Sandokanidae   from the rest, establishing a monofamilial suborder Oncopodines vs. the Laniatores   s.s. .

Hypothesis L 6 ( Fig. 17 View FIGURE 17 ): Shultz elaborated morphological ( Shultz 1998) and molecular phylogenetic analyses ( Shultz & Regier 2001) that agreed with the notion ( Dumitrescu 1976 and Kury 1993) that the “expanded” Insidiatores were diphyletic. However, the branching pattern obtained was different from Kury-Dumitrescu scheme.

Hypothesis L 7 ( Fig. 18 View FIGURE 18 ): supported by Kury (2002), Giribet & Kury (2007) and Mendes (2009), also has the Travunioidea and Triaenonychoidea as two clades, but with the sister group sequence inverted in relation to hypothesis O 8. The special position for Sandokanidae   is also refuted by this hypothesis; this taxon appears nested within the Grassatores.

Hypothesis L 8 ( Fig. 19 View FIGURE 19 ): finally Giribet and collaborators (2010) and Sharma & Giribet (2011) in a molecular analysis also did not recover the “expanded” Insidiatores in full. They also found Sandokanidae   nested within Grassatores, although in a different position from hypothesis L 7. The need is clear to do a total evidence analysis to reconcile the molecular with the morphological view.