Triaenonychidae Sorensen 1886
Systematics of Triaenonychidae 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 difference...
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Zenodo
2021
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| Online Access: | https://dx.doi.org/10.5281/zenodo.4531733 https://zenodo.org/record/4531733 |
| Summary: | Systematics of Triaenonychidae 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. Clade A Clade A is found in eastern Australia and New Zealand. Morphological phylogenetic studies placed Pyenganella as sister group to Lomanella , 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), Pyenganella was recovered as an early diverging core triaenonychid. In our analyses, Pyenganella is always found as sister group to the New Zealand endemic genera Hendea and Hendeola 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 – 3 pairs of inferior setae, as noted by Forster (1954) for Hendea and Hendeola . Hickman (1958) noted that the superior setae of Pyenganella 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 ‘ Equitius group’ from eastern Australia, with a fairly complicated taxonomic history. The genus Monoxyomma Pocock, 1903 was a superficial group of taxa united by the number of tarsal segments on leg I. The type species was M. spinatum Pocock, 1903, and the genus later included two other species from Queensland, Australia: M. manicatum Roewer, 1920 and M. rotundum Forster, 1955. In a revision of Equitius Simon, 1880, Hunt (1985) synonymised the type species M. spinatum with Equitius (as E. spinatus ), 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 ‘ Equitius is replaced ’ (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 Equitius . (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 ‘ Equitius ’ cf. Monoxyomma (MCZ: IZ:152643 and NHMUC:Aus10b) may be the southern representatives. In the checklist of Triaenonychidae, Kury et al . (2014) included M. manicatum and M. rotundum in Equitius . Given the comments of Hunt (1985), we consider these incertae sedis as the genus level and label them as ‘ Equitius ’. The ‘ Equitius 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. Clade B Clade B is exclusively found in South Africa, with the exception of Ceratomontia argentina 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 ‘ Ceratomontia group’ and the ‘ Roewerania 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 ‘ Ceratomontia group’ mostly corresponds to Clade B, except that in our analyses Paramontia Lawrence, 1934 and Planimontia Kauri, 1961 are included, and Micromontia Lawrence, 1939 is excluded. Taxa from Kauri’ s ‘ Roewerania 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 Lizamontia Kury, 2004 for the three South African species that were included in the otherwise entirely Madagascan genus Acumontia Loman, 1898, to which they are not closely related. These species were originally included as part of Kauri’ s ‘ Roewerania group’, but in our analyses the genus constituted an early-diverging lineage of Clade B, closely related to the ‘ Ceratomontia group’ taxa, a relationship perhaps reflected in its morphology. As detailed in Kury (2004), compared to the ‘ Ceratomontia group’ sensu Kauri, Lizamontia lacks the ventral longitudinal band of small granules, and, unlike most other South African taxa, Lizamontia 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 et al . (2020 b ) recovered a polyphyletic Biacumontia 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 Lispomontia Lawrence, 1937 potentially falls in this clade. Lawrence (1937 b ) discusses its resemblance to Ceratomontia and Biacumontia based on tarsal formula, and its resemblance to ‘ Cryptobunus ’ (now Amatola Lawrence, 1931) and Biacumontia based on cheliceral morphology. As currently defined, Ceratomontia 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 Ceratomontia species were more closely related to the South African genera Monomontia and Austromontia , having several apomorphic characters not shared with the South American Ceratomontia , such as a ventral plate with a narrower base relative to the apex. Our results thus concur with earlier work that Ceratomontia is not monophyletic, and therefore is not a trans-Atlantic genus. The South American Ceratomontia 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 C. capensis Roewer, 1915 from South Africa, necessitating a new genus name for the South American Ceratomontia lineage. However, we refrain from doing so until more South American Ceratomontia can be included in molecular phylogenetic analyses. Clade C Clade C is entirely South American, with the exception of Hedwiga Roewer, 1931 from New Zealand. The monotypic Picunchenops 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 Picunchenops in the subfamily Triaenonychinae (tribe uncertain). Another lineage within Clade C comprises a set of closely related taxa including Triaenonyx Sørensen, 1886, all of which share a fairly similar male genitalic morphology, with an elongate and sometimes uniquely complex stylus (Maury 1987 a , 1987 b , 1988; Pérez-González and Werneck 2018) that is hypertrophied in the case of Araucanobunus Muñoz Cuevas, 1973 (Hunt and Maury 1993). Described as one of two South American genera in Triaenobuninae, the monotypic Araucanobunus was hypothesised to be closely related to Ankaratrix , the only Triaenobunine from Madagascar (Muñoz-Cuevas 1973). Although MuñozCuevas (1973) noted many differences between Araucanobunus and Americobunus 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 Triaenonyx consists of six species, five of which were described before 1916. The inclusion of Hedwiga in this clade was unexpected and no morphological similarities are obvious. Clade D Subclade D1 . This clade is found in South Africa, with the exception of the South American Nuncia verrucosa Maury, 1990 (MCZ: IZ:138122). The species Nuncia verrucosa is representative of a lineage with similar somatic morphology that also includes N. spinulosa Maury, 1990, and potentially N. rostrata Maury, 1990. The type species of Nuncia is N. obesa (Simon, 1899), found in the diverse New Zealand Nuncia 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 Nuncia were sampled, but their position among South African species was also found in the Sanger-based phylogeny of Baker et al . (2020 b ). Speleomontia 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: Sorensenella Pocock, 1902 and Karamea Forster, 1954. The South African ‘ Adaeum group’ includes several genera that share very similar somatic morphology to that of Adaeum 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 Cryptadaeum Lawrence, 1931, Micradaeum Lawrence, 1931, Montadaeum Lawrence, 1931, Paradaeum Lawrence, 1931, and Heteradaeum Lawrence, 1963 fall into the ‘ Adaeum 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 Dingupa 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 Mensamontia in Clade C. Subclade D2 . This subclade includes the three South African genera Roewerania Lawrence, 1934, Graemontia Staręga, 1992, and Micromontia . The male genitalia of Roewerania and Graemontia both show a deeply cleft dorsal plate (Kauri 1961; Kury 2006). This lineage likely includes the unsampled genera Austronuncia Lawrence, 1931, the monotypic Gunvoria Kauri, 1961, and the monotypic Yulella Lawrence, 1939. Gunvoria also shows a deeply cleft dorsal plate (Kauri 1961). Yulella natalensis (Lawrence, 1937) was originally described as a species of Roewerania Lawrence (1963) noted morphological similarities between Austronuncia and Yulella , and A. spinipalpus Lawrence, 1931 has extremely (and unusually) elongate pedipalps and chelicerae, much like Roewerania lignicola 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. Subclade D3 . This clade includes all taxa from Madagascar, excluding Flavonuncia and Ankaratrix , 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. Acumontia ): 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 Hovanuncia 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, Staręga (1992) states a synonymy for Flavonuncia and Hovanuncia 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 Ivohibea Lawrence, 1959, Millomontia Lawrence, 1959, and Millotonyx Lawrence, 1959 will almost certainly fall in this clade: all three genera share the same general body plan as Acumontia 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. Subclade D4 . This clade includes Ankaratrix from Madagascar as the sister group to three genera that are either endemic to Western Australia ( Breviacantha Kauri, 1954 and Dingupa ) or South Australia ( Yatala Roewer, 1942). Recently, Porto and Pérez-González (2020) explored soil crypsis in Ankaratrix while also describing four new species. The unsampled genus Perthacantha Roewer, 1931 is also likely included in this clade, as Hunt (1996) stated ‘ Perthacantha Roewer 1931 (= Dingupa 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 Dingupa show that the male genitalia are similar to the drawings for Breviacantha gisleni Kauri, 1954 (Kauri 1954). In the SVDQuartets analysis, subclade D4 is not monophyletic with Ankaratrix grouped with subclade D2, although with weak support. Subclade D5 . 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 Calliuncus 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 Calliuncus instead). The Tasmanian genera Notonuncia Hickman, 1958, Bryonuncia Hickman, 1958, and Ankylonuncia Hickman, 1958 were recovered as a closely related group. As shown in Hunt (1996) Bryonuncia and Ankylonuncia 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. Ankylonuncia barrowensis Hickman, 1958), are homologous to the ‘plates enveloping the stylus’. The monotypic genus Tasmanonyx 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 Leionuncia Hickman, 1958 most likely falls in this clade, close to Notonuncia , as recovered in Hunt (1996). The ‘ Nunciella group’ includes two deeply divergent clades. In our results, the genus Nunciella Roewer, 1929 is paraphyletic across three different lineages: (1) a clade including the Nunciella samples from Western Australia (MCZ: IZ:35908 and MCZ: IZ:132908) and N. cheliplus Roewer 1931 from Victoria; (2) N. kangarooensis Hunt, 1971, a species endemic to Kangaroo Island, South Australia; and (3) N. tasmaniensis 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 Nunciella contains ‘a miscellany of species of doubtful affinity’. The type species of Nunciella is N. aspera (Pocock, 1902), a species known from Western Australia, although Pocock’ s original description states the location for this species (then called Triaenonyx aspera ) as just ‘Australia’. Two new genus names will be needed for ‘ Nunciella ’. However, we do not name new genera here as detailed analyses of Nunciella are required. In this group, we also identify a new genus, called ‘new genus B Australia NHMUC:Aus025b’ (Fig. 1 J ), 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 Nuncia unifalculata is recorded from the extremely isolated Crozet Islands, which have never had a continental connection. This species was first described as Promecostethus unifalculatus 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 Nuncia . Later, Hickman (1939) redescribed this species based on a large set of adult specimens and formally transferred it to Nuncia . Forster (1954), when describing the New Zealand Nuncia (i.e. true Nuncia ) and considering Hickman’ s description, concluded that N. unifalculata does no : Published as part of Derkarabetian, Shahan, Baker, Caitlin M., Hedin, Marshal, Prieto, Carlos E. & 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) on pages 148-153, DOI: 10.1071/is20047, http://zenodo.org/record/4531309 : {"references": ["Hunt, G. S. (1996). A preliminary phylogenetic analysis of Australian Triaenonychidae (Arachnida: Opiliones). Revue Suisse de Zoologie hors serie I, 295 - 308.", "Mendes, A. C. (2009). Avaliacao do status sistematico dos taxons supragenericos da infra-ordem Insidiatores Loman, 1902 (Arachnida, Opiliones, Laniatores). Ph. D. thesis, Universidade Federal do Rio de Janeiro, Museu Nacional.", "Forster, R. R. (1954). 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