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Figure 10 in The evolution of pedipalps and glandular hairs as predatory devices in harvestmen (Arachnida, Opiliones)

Figure 10. Use of pedipalps during feeding. All macro photographs, except (E) and (F), taken in the field; (E) and (F) taken in captivity. A, B, Rilaena triangularis (Phalangiidae), the prey is held between the pedipalps, glandular setae partly in contact and contaminated with prey setae: A, juvenile feeding on a captured springtail; B, adult feeding on a mosquito. C, juvenile Opilio canestrinii (Phalangiidae) feeding on a psocopteran insect. D, unidentified Gagrellinae feeding on fungi, which is held both with pedipalpal claws and chelicerae. E, Ischyropsalis kollari (Ischyropsalididae) feeding on snail; the pedipalps are only used as feelers. F, Trogulus martensi (Trogulidae) feeding on snail; the pedipalps are highly reduced and enclosed in tergal processus, thus not visible. G, Sabacon cavicolens feeding on captured springtail. Photos (A) by Jan van Duinen; (B), (C), (E), and (F) by Jorg Pageler; (D) by Melvyn Yeo, with kind permission; (G) by Axel Schonhofer.

Figure 5 in The evolution of pedipalps and glandular hairs as predatory devices in harvestmen (Arachnida, Opiliones)

Figure 5. Ontogenetic dimorphism of pedipalps. A, B, Cynortella quadrimaculata (Cosmetidae): A, juvenile exhibiting elongated cylindrical pedipalps with modified claw; B, adult female, exhibiting shortened pedipalps with flattened tibia and raptorial claw. C, D, Dicranolasma pauper (Dicranolasmatidae): C, juvenile, exhibiting spines on femur and clavate setae on patella, tibia, and tarsus; D, adult female, exhibiting a relatively shorter pedipalp, lacking spines and clavate setae. E, F, Dicranopalpus spp. (Phalangiidae), E, juvenile of Dicranopalpus cf. pyrenaeus, with spine-like elongated plumose setae on femur and tibia (black colour); F, adult female of Dicranopalpus ramosus, lacking the spine-like plumose setae. Scale bars: A, B = 1 mm; C, D = 0.2 mm; E, F = 0.5 mm.

Figure 4 in The evolution of pedipalps and glandular hairs as predatory devices in harvestmen (Arachnida, Opiliones)

Figure 4. Negative allometry of pedipalpal length. Plot of the relative pedipalp length in relation to the body length in Palpatores for species lacking glandular setae ('none' in the image), with plumose setae, and with clavate setae. Thin lines between symbols mark different stages of the same species.

Figure 3 in The evolution of pedipalps and glandular hairs as predatory devices in harvestmen (Arachnida, Opiliones)

Figure 3. Joint and muscle modifications in the eupnoid clamp morphotype. A, B, 3D reconstruction from microcomputed tomography images, with different segments differentially coloured. Joints in arachnid appendages are usually only operated by flexor muscles and extension is achieved by elastic transarticular sclerites or internal haemolymph pressure. The pivot of the patella–tibia joint is tilted, such that it articulates laterally. This joint is controlled by two pairs of opposing muscles. C, patellar joints in Metagagrella cf. formosa (Sclerosomatidae). cl-de, claw depressor muscle; cl-le, claw levator muscle; co, condyle; fe, femur; fe-ap, femoral apophysis; pa, patella; pa-ap, patellar apophysis; pa-fm, patella flexor muscle; ta, tarsus; ta-fm, tarsus flexor muscle; ti, tibia; ti-ab, tibia abductor muscle; ti-ad, tibia adductor muscle; tr, trochanter.

Figure 1 in The evolution of pedipalps and glandular hairs as predatory devices in harvestmen (Arachnida, Opiliones)

Figure 1. Morphological variation of harvestmen pedipalps. All macro photographs were taken in the field. A, Siro acaroides (Sironidae) of the basal-most suborder Cyphopthalmi, exhibiting small, nonmodified (leg-like) pedipalps. B, Holoscotolemon querilhaci (Cladonychiidae), a more basal lineage of the suborder Laniatores, exhibiting massive raptorial pedipalps. C, an unidentified species of Epedanidae, a more derived lineage of Laniatores, exhibiting extreme elongation of the femur and patella, shifting the raptorial segments distally. D, Caddo agilis (Caddidae) the most basal lineage in the suborder Eupnoi, exhibiting thick femoral spines and a dense field of glandular setae on the prolateral sides of pedipalps (secretion droplets appear like whitish dew-drops between both pedipalps). E, Platybunus sp. (Phalangiidae) with thick femoral spines and both glandular seta bearing patellar and tibial apophyses. F, male Phalangium opilio (Phalangiidae), exhibiting pedipalps with highly increased length, and all spines, apophyses, and glandular setae reduced. G, Juvenile Gagrellinae harvestman (Sclerosomatidae) with well-developed patellar and tibial apophyses, bearing glandular setae. H, Sabacon viscayanus (Sabaconidae), belonging to the suborder Dyspnoi, showing a highly modified pedipalp with dense coverage of glandular setae and hyperflexible patellar–tibia and tibia–tarsus joints. I, juvenile Mitostoma chrysomelas (Nemastomatidae), exhibiting the typical dyspnoid 'tentacle' pedipalp, densely covered in glandular setae. Photos (C) and (G) by Melvyn Yeo and (I) by Jorg Pageler, with kind permission, all others by Axel Schonhofer.

Figure 2 in The evolution of pedipalps and glandular hairs as predatory devices in harvestmen (Arachnida, Opiliones)

Figure 2. Scanning electron micrographs showing characters of harvestmen pedipalps. Cyphophthalmi: A, Meghalaya sp. (Stylocellidae). Laniatores: B, Galibrotus cf. riedeli (Biantidae). Eupnoi: C, Gagrella cf. disticta (Sclerosomatidae). D, Megabunus rhinoceros (Phalangiidae); E, Dicranopalpus cf. pyrenaeus (Phalangiidae), juvenile; F, Ballarra longipalpis (Neopilionidae). Dyspnoi: G, Acropsopilio neozealandiae (Acropsopilionidae); H, Dendrolasma mirabile (Nemastomatidae); I, Mediostoma stussineri (Nemastomatidae). Condition of tarsal tip and pretarsus (claw): J, Meghalaya sp., claw reduced, knob-like tip; K, Dibunus similis (Dibunidae), enlarged raptorial claw (here flexed condition); L, Cynortellana quadrimaculata (Cosmetidae), juvenile, modified pretarsus, inset shows one of numerous pores at the bulbous region; M, Phalangium opilio (Phalangiidae), smooth claw; N, Amilenus aurantiacus (Phalangiidae), pectinate claw (arrowhead); O, Ballarra longipalpis, claw totally reduced, tarsal tip rounded; P, Mediostoma stussineri, claw totally reduced, tarsal tip tapered; Q, Acropsopilio neozealandiae, claw highly but not totally reduced (arrowhead). ap, apophysis; cs, clavate seta; cx, coxa; fe, femur; pa, patella; ps, plumose seta; pt, pretarsus (claw); sc, sensilla chaetica; ta, tarsus; ti, tibia; tr, trochanter. Scale bars: A–I, K, M = 100 µm, J, N–P = 30 µm, L = 50 µm (inset 1 µm), Q = 10 µm.

FIGURE 23. Hypothesis C4 in Opiliones are no longer the same—on suprafamilial groups in harvestmen (Arthropoda: Arachnida)

FIGURE 23. Hypothesis C4 of phylogeny of the Cyphophthalmi (Giribet et al. 2011). Finally material of Ogoveidae was obtained for molecular study. The result was this asymmetrical tree, with Pettalidae as sister group of the rest as in Clouse et al. (2010). They recovered one of Shear's clades, the Ogoveoidea. Three new infraordinal names were proposed.

FIGURE 20. Hypothesis C1 in Opiliones are no longer the same—on suprafamilial groups in harvestmen (Arthropoda: Arachnida)

FIGURE 20. Hypothesis C1 of phylogeny of the Cyphophthalmi (Shear 1980/1993, morph). Shear (1980) was the first attempt ever of a cladistic analysis of the Cyphophthalmi. The analysis resulted in a symmetrical arrangement. Shear proposed the two infraorders Tropicophthalmi and Temperophthalmi. A sixth family was added in 1993. This graph is a combination of both works by Shear.

FIGURE 17. Hypothesis L6 in Opiliones are no longer the same—on suprafamilial groups in harvestmen (Arthropoda: Arachnida)

FIGURE 17. Hypothesis L6 of phylogeny of the Laniatores (Shultz 1998). Shultz morphological analysis (1998) and Shultz & Regier molecular analysis (2001) made a scarce sample of the Laniatores. The Insidiatores were dismantled, but the position of Synthetonychia was not made explicit, by not having been used in the analyses. The proximity of the Travuniidae with the Grassatores resembled Loman's view of the Laniatores.

FIGURE 14. Hypothesis L3 in Opiliones are no longer the same—on suprafamilial groups in harvestmen (Arthropoda: Arachnida)

FIGURE 14. Hypothesis L3 of phylogeny of the Laniatores (Loman 1903). Only two years after creating the Insidiatores, Loman (1903) separated the Oncopodidae (which he called Sterrhonoti, here represented by Gnomulus) against the other Laniatores sensu stricto, called Camptonoti. This hypothesis lost support with the removal of the Cladonychiidae (here represented by Erebomaster) from the Phalangodidae by Briggs (1969).

FIGURE 19. Hypothesis L8 in Opiliones are no longer the same—on suprafamilial groups in harvestmen (Arthropoda: Arachnida)

FIGURE 19. Hypothesis L8 of phylogeny of the Laniatores (Giribet et al. 2010, mol). Grassatores remained firm, and the Insidiatores were dismantled, but in a novel way: the Synthetonychiidae were for the first time considered sister group to the other Laniatores. This result has been repeated by Sharma & Giribet (2011, mol). The new name Eulaniatores is proposed here to identify this clade, frontally conflicting with the Tricospilata concept illustrated in Fig. 18.

FIGURE 13. Hypothesis L2 in Opiliones are no longer the same—on suprafamilial groups in harvestmen (Arthropoda: Arachnida)

FIGURE 13. Hypothesis L2 of phylogeny of the Laniatores (Loman 1901; 1902). This author proposed a sharp separation between the Triaenonychidae (which he called Insidiatores, here represented by Equitius) against all other Laniatores (which he called simply Laniatores [sensu stricto] and here called Lomaniatores). Pocock (1902) followed this hypothesis of phylogeny. The term Insidiatores was later reused in a broader sense.

FIGURE 11. Hypothesis O11 in Opiliones are no longer the same—on suprafamilial groups in harvestmen (Arthropoda: Arachnida)

FIGURE 11. Hypothesis O11 of phylogeny of the Opiliones (Giribet et al. 1999, comb; 2002, comb). Following his 1997 thesis, Giribet recovered the early branching off for the Cyphophthalmi and resurrected the name Phalangida for its sister group. A new clade was also firstly proposed and named, causing the diphyly of the Palpatores—the Dyspnolaniatores. Both this hypothesis and J are the modern contrasting views of the deep relationships among Opiliones groups.

FIGURE 16. Hypothesis L5 in Opiliones are no longer the same—on suprafamilial groups in harvestmen (Arthropoda: Arachnida)

FIGURE 16. Hypothesis L5 of phylogeny of the Laniatores Šilhavý (1961). Šilhavý recognized the Travunioidea of Kratochvíl (here marked as "Insidiatores", in blue) and made a change contraposing Oncopodidae against all other Laniatores, which has been anticipated by Mello-Leitão (1944), who had produced a surprisingly similar phylogeny (the famous cactus). This arrangement was conserved by Bristowe (1976).

FIGURE 18. Hypothesis L7 in Opiliones are no longer the same—on suprafamilial groups in harvestmen (Arthropoda: Arachnida)

FIGURE 18. Hypothesis L7 of phylogeny of the Laniatores (Kury 2002). A series of morphological studies—Kury (2002, morph), Giribet & Kury (2007, morph) and Mendes (2009, morph)—also favored the dismantlement of the Insidiatores, but joining the Triaenonychidae with the Grassatores. The new name Tricospilata is here proposed for this clade.

FIGURE 10. Hypothesis O10 in Opiliones are no longer the same—on suprafamilial groups in harvestmen (Arthropoda: Arachnida)

FIGURE 10. Hypothesis O10 of phylogeny of the Opiliones (Shultz 1998, morph). This analysis recovered the old Sundevall's concept (Cyphophthalmi versus Phalangida), and Simon's Plagiostethi (by then already universally called "Palpatores"). Within Palpatores, he recognized the Dyspnoi, burying the Apagosterni. Surprisingly, a molecular analysis by Shultz & Regier (2001, mol) recovered the same configuration. After some analyses which did not recover the Palpatores (see hypothesis K in Fig. 11 below), Giribet et al. (2010, mol) obtained the same pattern.

FIGURE 9. Hypothesis O9 in Opiliones are no longer the same—on suprafamilial groups in harvestmen (Arthropoda: Arachnida)

FIGURE 9. Hypothesis O9 of phylogeny of the Opiliones (Martens 1980, morphological, non-numerical). By the end of the 20th century, the original concept of Palpatores was lost, so that a hypothesis recovering this clade received the new name Cyphopalpatores Martens. This was the first cladistic analysis of the Opiliones.

FIGURE 7. Hypothesis O7 in Opiliones are no longer the same—on suprafamilial groups in harvestmen (Arthropoda: Arachnida)

FIGURE 7. Hypothesis O7 of phylogeny of the Opiliones (Mello-Leitão 1944). This author combined some older hypotheses such as the dichotomy Laniatores versus Palpatores (still using the original Thorell's concept) and reintroduced the Apagosterni.

FIGURE 5. Hypothesis O5 in Opiliones are no longer the same—on suprafamilial groups in harvestmen (Arthropoda: Arachnida)

FIGURE 5. Hypothesis O5 of phylogeny of the Opiliones (Pocock 1902). Pocock suggested a group formed by today's Phalangioidea + Ischyropsalidoidea, thus introducing the name and the concept of Apagosterni. Loman (1903) was the first to accept this clade, which alternately lost and regained favor, but endured for a century.

FIGURE 15. Hypothesis L4 in Opiliones are no longer the same—on suprafamilial groups in harvestmen (Arthropoda: Arachnida)

FIGURE 15. Hypothesis L4 of phylogeny of the Laniatores Kratochvíl (1958). After the work of Forster (1954), who created the Synthetonychiidae, this family went straight into the vicinity of the Triaenonychidae in the views of all most authors. Kratochvíl proposed a symmetrical arrangement of the groups we now know as Insidiatores versus Grassatores (then respectively called Travunioidea and Oncopodoidea). The morphological cladistic analysis by Martens (1980) recovered this arrangement, as well as both combined analyses by Giribet et al. (1999; 2002). Kury in his catalogue (2003) resurrected and expanded the original concept of Loman's Insidiatores, using this name for the same clade recognized by Kratochvíl as Travunioidea.