Triaenonychidae Sorensen 1886
Creators: name: Derkarabetian, Shahan
name: Baker, Caitlin M.
name: Hedin, Marshal
name: Prieto, Carlos E.
name: Giribet, Gonzalo
Publisher: name: Zenodo
Year: 2021
Subjects: Biodiversity
Taxonomy
Animalia
Arthropoda
Arachnida
Opiliones
Triaenonychidae

description: <i>Systematics of Triaenonychidae</i> Here we discuss each of the major lineages within Triaenonychidae, cover relevant taxonomic history and morphological characteristics, provide hypothesised placement of unsampled genera, and highlight potential for future revisionary work. Although there are differences in relationships among major clades across phylogenetic analyses, essentially all major clades and subclades are recovered, with the exception of a few individual taxa showing uncertain placement noted below. We present our discussion based on the IQ-TREE results, noting any uncertainty with the BEAST and SVDQuartets phylogenies in the relevant sections. <i>Clade A</i> Clade A is found in eastern Australia and New Zealand. Morphological phylogenetic studies placed <i>Pyenganella</i> as sister group to <i>Lomanella</i>, sharing several characters including some in the male genitalia (Hunt 1996; Mendes 2009). However, this was a hypothetical placement in Hunt (1996), and not the actual result of analysing his morphological dataset (see his fig. 1 and 2), and was obtained only with particular character weighting schemes in Mendes (2009), who also emphasised caution in this relationship. In the other analyses of Mendes (2009), <i>Pyenganella</i> was recovered as an early diverging core triaenonychid. In our analyses, <i>Pyenganella</i> is always found as sister group to the New Zealand endemic genera <i>Hendea</i> and <i>Hendeola</i> Forster, 1954. A potential synapomorphy for these three genera might be found in the male genitalia: on the ventral plate the single superior seta on each lobe is noted to be stronger and stouter than the 2 <i>–</i> 3 pairs of inferior setae, as noted by Forster (1954) for <i>Hendea</i> and <i>Hendeola</i>. Hickman (1958) noted that the superior setae of <i>Pyenganella</i> are longer than the inferior, but we add that they are also slightly stouter (S. Derkarabetian, pers. obs.) This clade also includes what we call the ‘ <i>Equitius</i> group’ from eastern Australia, with a fairly complicated taxonomic history. The genus <i>Monoxyomma</i> Pocock, 1903 was a superficial group of taxa united by the number of tarsal segments on leg I. The type species was <i>M. spinatum</i> Pocock, 1903, and the genus later included two other species from Queensland, Australia: <i>M. manicatum</i> Roewer, 1920 and <i>M. rotundum</i> Forster, 1955. In a revision of <i>Equitius</i> Simon, 1880, Hunt (1985) synonymised the type species <i>M. spinatum</i> with <i>Equitius</i> (as <i>E. spinatus</i>), but stated that the other two species belong to a ‘new genus together with some undescribed species ranging from northern Queensland to southern New South Wales’. It is unclear whether this undescribed genus has a continuous distribution along eastern Australia or a disjunct distribution; Hunt (1985) states that ‘ <i>Equitius</i> is <i>replaced</i> ’ (emphasis added) in the north and south by species of this new genus, implying allopatry. Across studies, Hunt refers to multiple undescribed genera: the one mentioned above (discussed in Hunt 1985) and ‘new genus C’ and ‘new genus D in Hunt (1996) ’. Both of these new genera were included in the phylogenetic analyses of Hunt (1996) and recovered in a clade with <i>Equitius</i>. (Hunt never explicitly mentioned ‘new genus A’ and ‘new genus B’ implied by the previous statements.) It is likely that one of the new genera in Hunt (1996) is the undescribed genus mentioned in Hunt (1985), and equally likely that both of these genera are represented as sequenced specimens recovered in Clade A. Our sample of MCZ: IZ:23313 collected from southern Queensland may likely be a northern representative of this undescribed genus, whereas the two we have identified as ‘ <i>Equitius</i> ’ cf. <i>Monoxyomma</i> (MCZ: IZ:152643 and NHMUC:Aus10b) may be the southern representatives. In the checklist of Triaenonychidae, Kury <i>et al</i>. (2014) included <i>M. manicatum</i> and <i>M. rotundum</i> in <i>Equitius</i>. Given the comments of Hunt (1985), we consider these <i>incertae sedis</i> as the genus level and label them as ‘ <i>Equitius</i> ’. The ‘ <i>Equitius</i> group’ including the potential (and yet another) undescribed genus from the Otway Ranges of Victoria (MCZ: IZ:152660 and SDSU _TAC: OP2779) will require more detailed taxonomic attention. <i>Clade B</i> Clade B is exclusively found in South Africa, with the exception of <i>Ceratomontia argentina</i> from South America (MCZ: IZ 152640, discussed below). Species in this clade are typically smaller in size relative to the other South African taxa (Clade D1). Kauri (1961) proposed two main groups, the ‘ <i>Ceratomontia</i> group’ and the ‘ <i>Roewerania</i> group’, divided according to the respective presence or absence of a longitudinal band of small granules ventrally on the pedipalp femur. Both groups are paraphyletic in our analyses. Kauri’ s ‘ <i>Ceratomontia</i> group’ mostly corresponds to Clade B, except that in our analyses <i>Paramontia</i> Lawrence, 1934 and <i>Planimontia</i> Kauri, 1961 are included, and <i>Micromontia</i> Lawrence, 1939 is excluded. Taxa from Kauri’ s ‘ <i>Roewerania</i> group’ are predominantly found in subclades D1 and D2, those subclades containing all the other South African taxa. More recently, Kury (2004) described the genus <i>Lizamontia</i> Kury, 2004 for the three South African species that were included in the otherwise entirely Madagascan genus <i>Acumontia</i> Loman, 1898, to which they are not closely related. These species were originally included as part of Kauri’ s ‘ <i>Roewerania</i> group’, but in our analyses the genus constituted an early-diverging lineage of Clade B, closely related to the ‘ <i>Ceratomontia</i> group’ taxa, a relationship perhaps reflected in its morphology. As detailed in Kury (2004), compared to the ‘ <i>Ceratomontia</i> group’ <i>sensu</i> Kauri, <i>Lizamontia</i> lacks the ventral longitudinal band of small granules, and, unlike most other South African taxa, <i>Lizamontia</i> lacks sexual dimorphism in the chelicerae and stout spines on the pedipalp femur. A potential synapomorphy for Clade B, found in the male genitalia, is fused parastyli that wrap around the stylus like a coat (e.g. Kauri 1961; Kury 2004), but all presumptive genera of this clade have not been examined. It is likely that a specieslevel revision of this lineage will find that some South African genera are paraphyletic; indeed, Baker <i>et al</i>. (2020 <i>b</i>) recovered a polyphyletic <i>Biacumontia</i> Lawrence, 1931. Many taxa are differentiated by leg tarsus segmentation, and species assignment and identification can be ambiguous based on available descriptions, the majority of which were done by R.F. Lawrence in the 1930s before the standardised use of genitalic morphology in Opiliones systematics. The unsampled monotypic genus <i>Lispomontia</i> Lawrence, 1937 potentially falls in this clade. Lawrence (1937 <i>b</i>) discusses its resemblance to <i>Ceratomontia</i> and <i>Biacumontia</i> based on tarsal formula, and its resemblance to ‘ <i>Cryptobunus</i> ’ (now <i>Amatola</i> Lawrence, 1931) and <i>Biacumontia</i> based on cheliceral morphology. As currently defined, <i>Ceratomontia</i> is one of two transcontinental triaenonychid genera, with four species in South America largely distributed east of the Andes in Argentina, Uruguay, and south-eastern Brazil, and 18 species found in South Africa. Each regional group is represented in our dataset by a single sample and our analyses support paraphyly, a result also found in the morphological analyses of Mendes and Kury (2008). In their study, the South African <i>Ceratomontia</i> species were more closely related to the South African genera <i>Monomontia</i> and <i>Austromontia</i>, having several apomorphic characters not shared with the South American <i>Ceratomontia</i>, such as a ventral plate with a narrower base relative to the apex. Our results thus concur with earlier work that <i>Ceratomontia</i> is not monophyletic, and therefore is not a trans-Atlantic genus. The South American <i>Ceratomontia</i> have a penis with an undivided ventral plate and five setae on each side of the stylus, whereas the South African group has a cleft ventral plate and four setae on each side. The type species of the genus is <i>C. capensis</i> Roewer, 1915 from South Africa, necessitating a new genus name for the South American <i>Ceratomontia</i> lineage. However, we refrain from doing so until more South American <i>Ceratomontia</i> can be included in molecular phylogenetic analyses. <i>Clade C</i> Clade C is entirely South American, with the exception of <i>Hedwiga</i> Roewer, 1931 from New Zealand. The monotypic <i>Picunchenops</i> is a single-site cave-obligate species from Argentina that shows extreme troglomorphy, including complete loss of eyes. When it was described, Maury (1988) placed it into Triaenonychidae based on male genitalia, but could not confidently assign it to any of the triaenonychid subfamilies. Later, Kury (2003) placed <i>Picunchenops</i> in the subfamily Triaenonychinae (tribe uncertain). Another lineage within Clade C comprises a set of closely related taxa including <i>Triaenonyx</i> Sørensen, 1886, all of which share a fairly similar male genitalic morphology, with an elongate and sometimes uniquely complex stylus (Maury 1987 <i>a</i>, 1987 <i>b</i>, 1988; Pérez-González and Werneck 2018) that is hypertrophied in the case of <i>Araucanobunus</i> Muñoz Cuevas, 1973 (Hunt and Maury 1993). Described as one of two South American genera in Triaenobuninae, the monotypic <i>Araucanobunus</i> was hypothesised to be closely related to <i>Ankaratrix</i>, the only Triaenobunine from Madagascar (Muñoz-Cuevas 1973). Although MuñozCuevas (1973) noted many differences between <i>Araucanobunus</i> and <i>Americobunus</i> Muñoz Cuevas, 1972, comparisons with other South American taxa were lacking, largely because few South American triaenonychids were known at this time. The Triaenonychidae type genus <i>Triaenonyx</i> consists of six species, five of which were described before 1916. The inclusion of <i>Hedwiga</i> in this clade was unexpected and no morphological similarities are obvious. <i>Clade D</i> <i>Subclade D1</i>. This clade is found in South Africa, with the exception of the South American <i>Nuncia verrucosa</i> Maury, 1990 (MCZ: IZ:138122). The species <i>Nuncia verrucosa</i> is representative of a lineage with similar somatic morphology that also includes <i>N. spinulosa</i> Maury, 1990, and potentially <i>N. rostrata</i> Maury, 1990. The type species of <i>Nuncia</i> is <i>N. obesa</i> (Simon, 1899), found in the diverse New Zealand <i>Nuncia</i> E2 subclade. As such, this South American lineage will need a new genus name. We refrain from naming this clade here as not all South American <i>Nuncia</i> were sampled, but their position among South African species was also found in the Sanger-based phylogeny of Baker <i>et al</i>. (2020 <i>b</i>). <i>Speleomontia</i> Lawrence, 1931 was one of two South African genera included in the subfamily Sorensenellinae, and based on our results are completely unrelated to the Sorensenellinae from New Zealand: <i>Sorensenella</i> Pocock, 1902 and <i>Karamea</i> Forster, 1954. The South African ‘ <i>Adaeum</i> group’ includes several genera that share very similar somatic morphology to that of <i>Adaeum</i> Karsch, 1880, typically large-bodied with a highly denticulate dorsal scutum, often found dirt-encrusted. Based on this morphology, it is almost certain that the unsampled genera <i>Cryptadaeum</i> Lawrence, 1931, <i>Micradaeum</i> Lawrence, 1931, <i>Montadaeum</i> Lawrence, 1931, <i>Paradaeum</i> Lawrence, 1931, and <i>Heteradaeum</i> Lawrence, 1963 fall into the ‘ <i>Adaeum</i> group’ as well (Lawrence 1931, 1963). These taxa are largely differentiated by the structure of the sternum and tarsal claw segmentation. This lineage essentially corresponds to the subfamily Adaeinae as currently defined, excluding <i>Dingupa</i> Forster, 1952 from Australia. In the BEAST and SVDQuartets analyses D1 is sister group to Clade E, rendering Clade D paraphyletic. Similarly, both analyses placed <i>Mensamontia</i> in Clade C. <i>Subclade D2</i>. This subclade includes the three South African genera <i>Roewerania</i> Lawrence, 1934, <i>Graemontia</i> Star&amp;eogon;ga, 1992, and <i>Micromontia</i>. The male genitalia of <i>Roewerania</i> and <i>Graemontia</i> both show a deeply cleft dorsal plate (Kauri 1961; Kury 2006). This lineage likely includes the unsampled genera <i>Austronuncia</i> Lawrence, 1931, the monotypic <i>Gunvoria</i> Kauri, 1961, and the monotypic <i>Yulella</i> Lawrence, 1939. <i>Gunvoria</i> also shows a deeply cleft dorsal plate (Kauri 1961). <i>Yulella natalensis</i> (Lawrence, 1937) was originally described as a species of <i>Roewerania</i>; Lawrence (1963) noted morphological similarities between <i>Austronuncia</i> and <i>Yulella</i>, and <i>A. spinipalpus</i> Lawrence, 1931 has extremely (and unusually) elongate pedipalps and chelicerae, much like <i>Roewerania lignicola</i> Lawrence, 1934 (Lawrence 1931, 1934). Although the BEAST analyses recover a paraphyletic D2 with respect to D3, the unsampled taxa would still be associated with D2 genera in a D2 + D3 clade. <i>Subclade D3</i>. This clade includes all taxa from Madagascar, excluding <i>Flavonuncia</i> and <i>Ankaratrix</i>, most of which were described by Lawrence (1959) and have remained largely unstudied since then. Many of these genera show extremely similar somatic morphology (e.g. <i>Acumontia</i>): large bodied, most with large dorsal spines on the scutum, an elongate spine on the eye mound, and leg I with 4+ tarsal segments. The only genus not fitting this general pattern is <i>Hovanuncia</i> Lawrence, 1959, which lacks spines on the eye mound and scutum and only has three tarsal segments on leg I. In a checklist of Afrotropical Opiliones, Star&amp;eogon;ga (1992) states a synonymy for <i>Flavonuncia</i> and <i>Hovanuncia</i>; however, this synonymy only appeared in the abstract and was never followed, not even in the checklist included in the same paper. Obviously, these genera are distinct. The unsampled Madagascan genera <i>Ivohibea</i> Lawrence, 1959, <i>Millomontia</i> Lawrence, 1959, and <i>Millotonyx</i> Lawrence, 1959 will almost certainly fall in this clade: all three genera share the same general body plan as <i>Acumontia</i> and similar genitalic morphology. A sequenced specimen which could not be assigned to an existing genus based on morphology (MCZ: IZ:52828) potentially represents a new genus. <i>Subclade D4</i>. This clade includes <i>Ankaratrix</i> from Madagascar as the sister group to three genera that are either endemic to Western Australia (<i>Breviacantha</i> Kauri, 1954 and <i>Dingupa</i>) or South Australia (<i>Yatala</i> Roewer, 1942). Recently, Porto and Pérez-González (2020) explored soil crypsis in <i>Ankaratrix</i> while also describing four new species. The unsampled genus <i>Perthacantha</i> Roewer, 1931 is also likely included in this clade, as Hunt (1996) stated ‘ <i>Perthacantha</i> Roewer 1931 (= <i>Dingupa</i> Forster 1952)’, though without any explanation or justification. We are keeping these two genera distinct until a more direct comparison can be made. Our examinations of <i>Dingupa</i> show that the male genitalia are similar to the drawings for <i>Breviacantha gisleni</i> Kauri, 1954 (Kauri 1954). In the SVDQuartets analysis, subclade D4 is not monophyletic with <i>Ankaratrix</i> grouped with subclade D2, although with weak support. <i>Subclade D5</i>. This is a widespread Australian lineage that somewhat corresponds to a group recovered in Hunt’ s (1996) morphological phylogenetic analyses, though it does not include <i>Calliuncus</i> Roewer, 1931. This group had an apomorphy of a penis stylus completely enveloped by plates, a character shared with taxa in Clade E1 (where we find <i>Calliuncus</i> instead). The Tasmanian genera <i>Notonuncia</i> Hickman, 1958, <i>Bryonuncia</i> Hickman, 1958, and <i>Ankylonuncia</i> Hickman, 1958 were recovered as a closely related group. As shown in Hunt (1996) <i>Bryonuncia</i> and <i>Ankylonuncia</i> were not recovered in the above-mentioned group, and hence do not share this apomorphy. However, they were found to be each other’ s closest relatives united by a penis with a ‘mid-dorsal large spiny lobe’. It is possible that re-examination and reinterpretation of genitalia may find that the ‘large spiny lobes’, which do surround the stylus (e.g. <i>Ankylonuncia barrowensis</i> Hickman, 1958), are homologous to the ‘plates enveloping the stylus’. The monotypic genus <i>Tasmanonyx</i> is unique in this lineage in having hind claws with the lateral branches longer than median prong, a characteristic shared with the nonmonophyletic Sorensenellinae. The unsampled genus <i>Leionuncia</i> Hickman, 1958 most likely falls in this clade, close to <i>Notonuncia</i>, as recovered in Hunt (1996). The ‘ <i>Nunciella</i> group’ includes two deeply divergent clades. In our results, the genus <i>Nunciella</i> Roewer, 1929 is paraphyletic across three different lineages: (1) a clade including the <i>Nunciella</i> samples from Western Australia (MCZ: IZ:35908 and MCZ: IZ:132908) and <i>N. cheliplus</i> Roewer 1931 from Victoria; (2) <i>N. kangarooensis</i> Hunt, 1971, a species endemic to Kangaroo Island, South Australia; and (3) <i>N. tasmaniensis</i> Hickman, 1958 from Tasmania. This non-monophyly is not surprising given the fact this genus is diagnosed by the number of tarsal segments of the first leg and Hunt’ s (1985) statement that <i>Nunciella</i> contains ‘a miscellany of species of doubtful affinity’. The type species of <i>Nunciella</i> is <i>N. aspera</i> (Pocock, 1902), a species known from Western Australia, although Pocock’ s original description states the location for this species (then called <i>Triaenonyx aspera</i>) as just ‘Australia’. Two new genus names will be needed for ‘ <i>Nunciella</i> ’. However, we do not name new genera here as detailed analyses of <i>Nunciella</i> are required. In this group, we also identify a new genus, called ‘new genus B Australia NHMUC:Aus025b’ (Fig. 1 <i>J</i>), which differs from related taxa in both somatic and male genitalic morphology. The two specimens sequenced are both from the Otway Ranges, Victoria, but we refrain from describing this genus here until more specimens can be obtained for detailed anatomical study. A species currently known as <i>Nuncia unifalculata</i> is recorded from the extremely isolated Crozet Islands, which have never had a continental connection. This species was first described as <i>Promecostethus unifalculatus</i> by Enderlein (1909) based on four specimens, three immatures and one adult. However, upon examination, Roewer (1923) concluded that all specimens were immature, and postulated that it may belong to the transcontinental genus <i>Nuncia</i>. Later, Hickman (1939) redescribed this species based on a large set of adult specimens and formally transferred it to <i>Nuncia</i>. Forster (1954), when describing the New Zealand <i>Nuncia</i> (i.e. true <i>Nuncia</i>) and considering Hickman’ s description, concluded that <i>N. unifalculata</i>
description: Published as part of <i>Derkarabetian, Shahan, Baker, Caitlin M., Hedin, Marshal, Prieto, Carlos E. &amp; Giribet, Gonzalo, 2021, Phylogenomic re-evaluation of Triaenonychoidea (Opiliones: Laniatores), and systematics of Triaenonychidae, including new families, genera and species, pp. 277-288 in Invertebrate Systematics 167 (1)</i> on pages 148-153, DOI: 10.1071/is20047, http://zenodo.org/record/4531309
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