identifier	taxonID	type	CVterm	format	language	title	description	additionalInformationURL	UsageTerms	rights	Owner	contributor	creator	bibliographicCitation
03A2D5375E55FFFDD1B1F924FBC7BDB3.text	03A2D5375E55FFFDD1B1F924FBC7BDB3.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Laniatores	<html xmlns:mods="http://www.loc.gov/mods/v3">
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            <p> Laniatores</p>
            <p> All investigated species of  Laniatores , in 28 families, possess three distinct sensilla on the distal-third end of the distalmost tarsomere of leg pairs I and II, roughly aligned along the sagittal axis (Fig. 2). The only exception is  Sandokan truncatus Thorell, 1891 (  Sandokanidae ), in which only the distal sensillum was clearly identified in the corresponding position. The distal sensillum has been termed the ‘hooded sensillum’ (Gainett et al., 2017b) and the mid and proximal sensilla have been termed ‘sensilla basiconica’ (after Willemart et al., 2007, 2009). Below, we describe the variation observed in the morphology and topology of these three sensilla on the distalmost tarsomeres I and II. </p>
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	https://treatment.plazi.org/id/03A2D5375E55FFFDD1B1F924FBC7BDB3	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Gainett, Guilherme;Sharma, Prashant P;Fernandes, Nathália;Pinto-Da-Rocha, Ricardo;Giribet, Gonzalo;Willemart, Rodrigo Hirata	Gainett, Guilherme, Sharma, Prashant P, Fernandes, Nathália, Pinto-Da-Rocha, Ricardo, Giribet, Gonzalo, Willemart, Rodrigo Hirata (2019): Evolution of a sensory cluster on the legs of Opiliones (Arachnida) informs multi-level phylogenetic relationships. Zoological Journal of the Linnean Society 187 (1): 143-165, DOI: 10.1093/zoolinnean/zlz016, URL: https://academic.oup.com/zoolinnean/article/187/1/143/5437571
03A2D5375E53FFF9D229F9E4FD3FB81F.text	03A2D5375E53FFF9D229F9E4FD3FB81F.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Dyspnoi	<html xmlns:mods="http://www.loc.gov/mods/v3">
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            <p> Dyspnoi</p>
            <p> The three species of  Dyspnoi studied show three sensilla aligned along the sagittal axis in a corresponding position as  Laniatores , on the distalmost tarsomeres of legs I and II. The three sensilla differ from the surrounding sensilla chaetica and trichomes by having a flat articulation membrane (Figs 7, 8). The proximal two sensilla are short and conical, being also termed sensilla basiconica. The distal sensillum has a very similar morphology to the hooded sensillum in  Laniatores and, therefore, we refer to it using the same name. </p>
            <p>The hooded sensillum is longer than both sensilla basiconica (Figs 7A, C, 8A, D). The shaft has longitudinal ridges and gradually tapers into a fine tip (Figs 7C, 8A). The apical portion has two pore-like structures in a region without ridges, but shows no subterminal swelling (see Ps in Figs 7D, 8D). The distal pore-like structure is smaller than the proximal (Fig. 7D). The side opposite to the pore-like structures is similar to the rest of the shaft, with ridges.</p>
            <p> The sensilla basiconica are thin pegs, being the shortest sensilla in the tarsomere (Figs 7A, B, 8A–C). The proximal sensillum in  Anelasmocephalus sp. (  Trogulidae ) is shorter than the mid sensillum, both being shorter than the distal-hooded sensillum (Fig. 8A), but in the other species we could not access their relative length. The shaft is bent forward in its mid portion, in an angle of almost 90º (Figs 7B, 8A– C). Apically, it bears a subterminal pore with a short slit, which faces down in  Nemastoma bimaculatum Fabricius, 1775 (  Nemastomatidae ) (Fig. 7A, B) or up in  Anelasmocephalus sp. (Fig. 8A–C). The slit apparently does not divide the shaft into two flaps as in  Laniatores . We observed no articulation membrane fusion between the three sensilla and three shafts are more interspaced than in  Laniatores (Figs 7A, 8A). </p>
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	https://treatment.plazi.org/id/03A2D5375E53FFF9D229F9E4FD3FB81F	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Gainett, Guilherme;Sharma, Prashant P;Fernandes, Nathália;Pinto-Da-Rocha, Ricardo;Giribet, Gonzalo;Willemart, Rodrigo Hirata	Gainett, Guilherme, Sharma, Prashant P, Fernandes, Nathália, Pinto-Da-Rocha, Ricardo, Giribet, Gonzalo, Willemart, Rodrigo Hirata (2019): Evolution of a sensory cluster on the legs of Opiliones (Arachnida) informs multi-level phylogenetic relationships. Zoological Journal of the Linnean Society 187 (1): 143-165, DOI: 10.1093/zoolinnean/zlz016, URL: https://academic.oup.com/zoolinnean/article/187/1/143/5437571
03A2D5375E51FFF5D229FB52FDB6BB37.text	03A2D5375E51FFF5D229FB52FDB6BB37.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Cyphophthalmi	<html xmlns:mods="http://www.loc.gov/mods/v3">
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            <p> Cyphophthalmi</p>
            <p> The distal third of the last tarsomeres of legs I and II of the cyphophthalmid species studied show a structure termed the subapical process (Fig. 11). Juberthie (1979, 1988, 2000) first described the subapical process, which occurs isolated on the distal third of tarsomeres I and II only, of males and females of some  Cyphophthalmi species. Willemart &amp; Giribet (2010) later found this structure to be widespread in  Cyphophthalmi . Our investigation of two  Cyphophthalmi species, belonging to one family each (  Troglosironidae and  Pettalidae ), confirms the occurrence of this structure in the corresponding position of the hooded sensillum and sensilla basiconica found in the other suborders. In the species here investigated, this single sensillum occurs among other types of sensilla (solenidia, sensilla chaetica), but no short and conic sensilla, such as sensillum basiconicum, is present in the tarsomere (Fig. 11A). The shaft is wide at the base (~6.5 µm) and tapers to a fine rounded tip (Fig. 11C), possessing a high density of longitudinal ridges (Fig. 11B, inset). The apical portion has no subterminal swelling and no pore-like structures </p>
            <p>CHARACTERS AND CODING</p>
            <p> In the previous section, we described the comparative incidence of sensilla with external morphological similarities in the shaft and articulation membrane, with a restricted distribution to the distal third of the distalmost tarsomeres of legs I and II of representatives of all suborders of  Opiliones . In order to understand the variation in Phalangida (  Eupnoi +  Dyspnoi +  Laniatores ), we established morphological characters and hypothesized, based on position and morphological similarity, that the subapical process of  Cyphophthalmi is homologous to the hooded sensillum of Phalangida. </p>
            <p> This conserved sensory field shows variation in the association of the articulation membrane between the shafts, composition of sensillar types and shaft morphology. In order to trace how this sensory field has been modified across  Opiliones evolution, we propose the following characters and character states: </p>
            <p>Character 1: Sensory cluster, number of sensilla. States: (1a) one, (1b) three.</p>
            <p>Character 2: Sensory cluster, articulation membrane configuration. States: (2a) distal, mid and proximal sensilla not fused (notation: ‘1 2 3’); (2b) distal sensillum isolated, mid and proximal sensilla fused (notation: ‘1 (2 3) ’); (2c) distal, mid and proximal sensilla fused (notation: ‘ (1 2 3) ’).</p>
            <p>Character 3: Hooded sensilla, hood morphology. States: (3a) terminal swelling (spoon-shaped), (3b) sub-terminal swelling (‘death’s hood’), (3c) no swelling (regular).</p>
            <p>Character 4: Mid and proximal sensilla, apical opening: (4a) pore-like, (4b) complete slit.</p>
            <p> Supporting Information, File S2 contains the l i s t o f f e a t u r e s o r g a n i z e d b y c h a r a c t e r s a n d character states for all species studied. States listed are representative of the morphology of leg pairs I and II, unless otherwise stated in the few cases of serial polymorphisms observed. In  Avima octomaculata Agoristenidae (  Laniatores ),  Poassa limbata (  Laniatores ,  Nomoclastidae ) and  Astrobunus grallator (  Eupnoi ,  Sclerosomatidae ) legs I and II have different character states (for character 2). Apart from that, species in which we could access the character in both legs always showed the same character state and morphology (27 species). In order to ensure comparability, we chose to code leg pair I for our analysis. For eight species in which legs I could not be accessed properly for character 2, the sensilla on leg pair II was used as proxy for coding leg pair I (species marked with asterisk in Supporting Information, File S2). </p>
            <p>ANCESTRAL STATE RECONSTRUCTION</p>
            <p> Variation in the number of sensilla composing the sensory field (distal-third regions of the distalmost tarsomeres of legs I and II) (character 1) has two equally parsimonious optimizations (cost = 2; PTP &lt;0.05; mean: 2.97; median: 3): either having one sensillum (1a, white) is the ancestral state of  Opiliones , which has increased to three sensilla (1b, black) in Phalangida (  Eupnoi +  Dyspnoi +  Laniatores ); or having three sensilla is the ancestral state of  Opiliones (1 b, black), which has become secondarily reduced to one sensillum (1a, white) in  Cyphophthalmi (Fig. 12). Reversion from three (1b, black) to one sensillum (1a, white) is unambiguously recovered in the laniatorean family  Sandokanidae (  Sandokan truncatus ) (Fig. 12). </p>
            <p> Association between the articulation membranes of the three sensilla (character 2) is inapplicable for  Cyphophthalmi species and ambiguous for the most recent common ancestor of Phalangida and Palpatores (Fig. 13). The ancestral state in  Laniatores is recovered as ‘1 (2 3) ’. Several transformations occur in the equally parsimonious reconstructions (cost = 9; PTP &lt;0.05; mean: 18.69; median: 19), with ambiguous ancestral states in most cases. The character state, all sensilla fused (2c, black), occurs independently is some families of  Laniatores (  Podoctidae ,  Stygnommatidae ,  Biantidae and  Agoristenidae ) and is recovered as the ancestral character state for the  Gonyleptoidea clade that excludes  Stygnopsidae ,  Agoristenidae and  Stygnidae (Fig. 13). Three separated sensilla (2a, white) were acquired independently in  Dyspnoi and a clade inside the laniatorean superfamily  Gonyleptoidea (  Metasarcidae +Cometidae+  Gonyleptidae ) (Fig. 13). </p>
            <p> Character 3 concerns the tip of the distal sensillum with hood morphology, which is a condition that is not directly applicable in  Cyphophthalmi and  Eupnoi , since the distalmost sensillum in  Cyphophthalmi apparently has no pore and in  Eupnoi the pore is not subterminal. Therefore, coding encompasses the comparable hood morphology observed in  Dyspnoi and  Laniatores only. Character 3 has undergone several changes in the phylogeny and is ambiguously optimized (cost = 8; PTP &lt;0.05; mean: 11.91; median: 12). In the clade Grassatores, five transformation events are unambiguous (Fig. 14). Terminal swelling (3a, white) has independently evolved from subterminal swelling (3b, green) in the laniatorean families  Podoctidae ,  Biantidae and  Guasiniidae . No swelling (3c, black) has independently evolved two times: in the family  Stygnopsidae (Gonyleptoidea) and in a clade inside  Gonyleptoidea including the families  Metasarcidae ,  Cosmetidae and two  Gonyleptidae (Fig. 14). </p>
            <p> Finally, the type of apical opening of the mid and proximal sensilla (character 4, inapplicable for  Cyphophthalmi ) is ambiguously reconstructed in Phalangida (cost = 1; PTP &lt;0.05; mean: 4.91; median: 5) (Fig. 15), but having a pore-like opening (4a, white) is clearly ancestral for Palpatores (  Eupnoi +  Dyspnoi ), whereas having a complete slit (4b) is unambiguously ancestral for  Laniatores . </p>
            <p>Ancestral state reconstructions under maximum likelihood using a one-parameter Markov model yielded virtually identical ancestral states for all nodes (Supporting Information, File S4).</p>
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	https://treatment.plazi.org/id/03A2D5375E51FFF5D229FB52FDB6BB37	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Gainett, Guilherme;Sharma, Prashant P;Fernandes, Nathália;Pinto-Da-Rocha, Ricardo;Giribet, Gonzalo;Willemart, Rodrigo Hirata	Gainett, Guilherme, Sharma, Prashant P, Fernandes, Nathália, Pinto-Da-Rocha, Ricardo, Giribet, Gonzalo, Willemart, Rodrigo Hirata (2019): Evolution of a sensory cluster on the legs of Opiliones (Arachnida) informs multi-level phylogenetic relationships. Zoological Journal of the Linnean Society 187 (1): 143-165, DOI: 10.1093/zoolinnean/zlz016, URL: https://academic.oup.com/zoolinnean/article/187/1/143/5437571
