identifier	taxonID	type	CVterm	format	language	title	description	additionalInformationURL	UsageTerms	rights	Owner	contributor	creator	bibliographicCitation
7D34D83FFFEB7178FF70B2804A2DCD84.text	7D34D83FFFEB7178FF70B2804A2DCD84.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Historical	<div><p>Historical background</p><p>The history of linyphiid classification is long and convoluted (for details, see Merrett 1963 and Hormiga 2000). With a few exceptions (Bertkau 1878; Gerhardt 1923; Wiehle 1956, 1960), the consensus has been that the Linyphiidae represents a natural group of spiders with highly variable genitalic anatomy (e.g. Hormiga 1994b; Griswold et al. 1998; Miller &amp; Hormiga 2004; Arnedo et al. 2009). Most linyphiid taxonomic classifications resulted from compilations of anatomical information followed by the choice of some particular character or characters deemed important for higher classification, such as the works of Hull (1920), Simon (1926, 1929), Petrunkevitch (1928), and especially Millidge (1977, 1984, 1986, 1988). Most of those studies predate Hennigian phylogenetic systematic theory (Hennig 1957, 1966) and/or lack an explicit phylogenetic approach.Modern studies on linyphiid phylogenetics began in the last decade of the past century (Hormiga 1994a, b; Hormiga 2000; Miller and Hormiga 2004).Although the use of molecular data in spider systematics is well established, the majority of linyphiid phylogenetic studies to date have used only morphological data. The only study to date (Arnedo et al. 2009) that has used a “total evidence” approach to resolve linyphiid familial relationships had a taxon sample too small (49 taxa) to draw detailed conclusions about such a large family. Wang et al. (2015) provide the most comprehensive study up to this point in both data and taxa (211 taxa, five DNA markers, but no morphological characters). However, their taxon sample was primarily tailored to address questions about relationships within the subfamily Erigoninae and, thus, was insufficient to address broad questions about relationships within the family. Recently, Silva-Moreira and Hormiga (2021, 2022) addressed the internal arrangement of linyphiids, but only in the context of establishing the monophyly of the revised genera Jalapyphantes Gertsch &amp; Davis, Selenyphantes Gertsch &amp; Davis, and Diplothyron Millidge. Both studies hypothesized higher level intrafamilial relationships and established new lineages (e.g., the Pocobletus clade, Silva-Moreira &amp; Hormiga 2021) or corroborate and expand previously established groups (e.g., Mounded Posterior Median Eye or MPME clade, a group of linyphiid spiders characterized by having posterior median eyes (PME) on a mound; Silva-Moreira &amp; Hormiga 2015, 2022). Nonetheless, neither study had been designed with the required taxonomic breadth or enough character data to build a robust phylogenetic hypothesis for the family Linyphiidae .</p><p>Our minimal knowledge of most of the spider fauna, especially from tropical areas, is a major obstacle to understanding the diversity of linyphiids. Some estimates suggest that up to 70% of collected spider specimens from those areas already deposited in museum collections are composed of undescribed species (Coddington &amp; Levi 1991; Agnarsson et al. 2013). Furthermore, many of the genera and species described from the Neotropical region have not been included in modern revisionary studies, and many remain known only from the type specimens, and the only information available about them is their original descriptions, some of them dating from the 19 th century. This “taxonomic impediment” (see Evenhuis 2007) has severe consequences for understanding global patterns of diversification within this clade. Good examples in Linyphiidae of how the addition of unknown fauna can dramatically change the understanding of a group can be seen in both the Mynogleninae (Lehtinen 1967; Blest 1979; Frick &amp; Scharff 2014), the Erigoninae (Miller &amp; Hormiga 2004; Miller 2007), the MPME clade; (Silva-Moreira &amp; Hormiga 2015, 2022), and the Pocobletus clade (Silva-Moreira &amp; Hormiga 2021). Some of the lineages whose species are endemic from Afrotropical/Australian and Neotropical regions (sensu Holt et al. 2013), such as Mynoglenines and the MPME clade, have unique features within linyphiids and, so far, are consistently recovered as well-supported clades (Frick &amp; Scharf 2014; Silva-Moreira &amp; Hormiga 2022).</p><p>Even with all those caveats, some hypotheses of linyphiid phylogeny have stood the test of a diversity of analyses and can be considered well corroborated. First, the monophyly of Linyphiidae is well supported by both morphological (Hormiga, 1994b; Griswold et al. 1998; Hormiga 2000; Miller &amp; Hormiga, 2004) and molecular data (Arnedo et al. 2009; Wang et al. 2015; Wheeler et al. 2017; Dimitrov et al. 2017; Hormiga et al., 2021).</p><p>This study aims to produce a new phylogenetic hypothesis of Linyphiidae using both morphological and molecular data. We test the monophyly, composition, and placement of all the seven classic subfamilies of Linyphiidae (as recognized in Tanasevitch 2025): Erigoninae, Dubiaraneinae, Ipainae, Linyphiinae, Micronetinae, Mynogleninae, and Stemonyphantinae) as well as of three recently proposed clades: MPME clade (Silva-Moreira &amp; Hormiga 2015), Porrhommini (Zhao &amp; Li 2017), and Pocobletus clade (Silva-Moreira &amp; Hormiga 2021). This study is the broadest scale phylogenetic matrix for linyphiids assembled to date. Despite the large taxon sample of this study relative to the phylogenetic analyses published so far, we recognize that our sample of linyphiid genera is insufficient for a thorough revision of the classification of this family and thus a detailed revision of such classification falls outside the scope of the study. We hope that the resulting hypotheses will provide a solid phylogenetic background upon which further questions on linyphiid evolution, such as biogeography or comparative biology, can be studied, as well as a foundation for future studies to expand the cladistic classification of this large and diverse family.</p></div>	https://treatment.plazi.org/id/7D34D83FFFEB7178FF70B2804A2DCD84	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.		MagnoliaPress via Plazi	Silva-Moreira, Thiago Da;Kulkarni, Siddharth;Hormiga, Gustavo	Silva-Moreira, Thiago Da, Kulkarni, Siddharth, Hormiga, Gustavo (2025): A total evidence phylogenetic analysis of the spider family Linyphiidae (Araneae, Araneoidea). Zootaxa 5685 (1): 1-78, DOI: 10.11646/zootaxa.5685.1.1, URL: https://doi.org/10.11646/zootaxa.5685.1.1
7D34D83FFFC07155FF70B3504D96CD84.text	7D34D83FFFC07155FF70B3504D96CD84.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Stemonyphantinae Wunderlich 1986	<div><p>Stemonyphantinae (Figs 3, 10)</p><p>Stemonyphantines have been quite consistently hypothesized as the sister group of a clade with all other Linyphiidae (e.g., Wunderlich 1986; Hormiga 1994a, b; Arnedo et al. 2009; Gavish-Regev et al. 2013; Frick and Scharff 2013; Dimitrov et al. 2017). Hormiga et al. (2021) expanded the Stemonyphantinae circumscription by including Weintrauboa and Putaoa (previously placed in Pimoidae) and the genus Pecado . The placement of Stemonyphantinae as the sister lineage to all other Linyphiidae did not change. Silva-Moreira and Hormiga (2022) did retrieve Pecado impudicus (Denis) as a sister group of Stemonyphantes using only phenotypic data. Pecado was erected by Hormiga &amp; Scharff (2005) to place a single species transferred from Labulla Simon, based on a phenotypic cladistic analysis of the latter genus.</p><p>Our preferred hypothesis (Fig. 3) concurs with Hormiga et al. (2021) in both the circumscription and placement of Stemonyphantinae: for the first time, Pecado position in Stemonyphantinae is supported by both molecular and phenotypic evidence. The monophyly of Stemonyphantinae (sensu Hormiga et al., 2021) is strongly supported in all analyses. The monophyly of Stemonyphantinae is supported by six phenotypic synapomorphies: male pedipalp with five or more macrosetae on the tibia (char 5), a dorsal or meso-dorsal oriented cymbium (char 28) with an ecto-marginal process (char 32), paracymbium with an anterobasal apophysis (char 49), the presence of ventral macrosetae on metatarsus I (char 224), and trichobothria on metatarsus IV (char 225).</p><p>The placement of Weintrauboa and Putaoa within Stemonyphantinae required the reinterpretation of some palpal homologies (see discussion in Hormiga et al. 2021). The male palp morphology of Pecado shares some features with that of Stemonyphantes, such as the presence of a secondary apical tegular membrane and a tegular ridge (Gavish-Regev et al. 2013 fig. 3A–B—char 82). Both the conductor (char 81) and the median apophysis (char 82) are widespread throughout Araneoidea, and the loss of both of those sclerites is thought to be a synapomorphy of Linyphiidae (Griswold et al. 1998, 2005; Hormiga 1994b). The analyses of Coddington (1990a) and Griswold et al. (1998) provided the cladistic foundation for studies to address the male palp homologies of araneoids. When a single tegular sclerite is found in araneoids, it is often difficult to determine whether it is a homolog of the conductor or of the median apophysis. This latter problem has led some workers to treat such sclerite as a conductor by default unless there is evidence to the contrary (e.g., Coddington 1990b; Griswold et al. 1998; Griswold et al. 2005). While such a pragmatic approach may provide a consistent system to code the sclerite in a character matrix, it does not necessarily resolve the homology problem per se. The conductor morphology of araneoids is very diverse, varying from a small and membranous (e.g., Pimoa) to a large and heavily sclerotized apophysis (e.g., in many araneids). In Pimoa, Weintrauboa, Putaoa, Stemonyphantes, Pecado, and Labulla, it is possible to see a membranous structure on the tegulum (in Nanoa, the conductor is slightly more sclerotized, Hormiga et al. 2005). In both Weintrauboa and Putaoa, this structure is discrete and interpreted as a membranous conductor as it is located on the apical part of the tegulum. In Labulla and Pecado, this structure is named “ectal tegular flap” or “tegular membrane” (Hormiga and Scharff 2005), and while in Pecado, it is positioned more apically, and in Labulla, it is on the ectal side of the tegulum. The position of the membrane on Stemonyphantes species varies but is usually found closer to the suprategulum (Gavish-Regev et al. 2013). Gavish-Regev et al. (2013 figs 3A, B) and Hormiga et al. (2021) proposed that the tegular ridge is a homolog of the conductor in Stemonyphantes . In Pecado we can observe a tegular structure that the same study interpreted as a conductor homolog. Thus, we have coded all stemonyphantines as having a conductor following Hormiga et al. (2021). We also interpret the median apophysis of Weintrauboa (Hormiga 2003, figs 1G, 5C) and part of the Putaoa pimoid embolic process (Hormiga &amp; Tu, 2008, figs 3B, C) as part of the suprategulum (char 72), the distal suprategular apophysis (char 78). Our phylogenetic hypothesis suggests that the alveolar sclerite (char 24) and the pimoid cymbial sclerite (char 35), both synapomorphic features of pimoids in Tu &amp; Hormiga (2008), are now understood as convergent features (Fig. 10).</p><p>Labulla thoracica (Figs 3, 10)</p><p>Labulla thoracica is the type species and the only representative of the genus Labulla Simon in our dataset. The position of Labulla as one of the earliest branches of Linyphiidae (Figs 1, 3, Supplementary Figs 1–4) is one of the most stable topological results in all analyses that include molecular data, regardless of the optimality criteria used.</p><p>Hormiga &amp; Scharff (2005) suggested that the Labulla clade (with three species) was sister to Tenuiphantes + Bolyphantes (here the Lepthyphantinae, see below) based on the following synapomorphies: ventral position of the ectal suprategular apophysis, the bifid shape of this apophysis, the presence of paracymbial apophyses and presence of a ventral plate epigynal scape (Hormiga and Scharff 2005, fig. 25). While in some of our analyses (with MORPH dataset) we recovered L. thoracica (and Ipa keyserlingi) as sister group to the Lepthyphantinae, our working hypothesis did not recover the same sister-group relationship. This latter result may be due to the lack of molecular data or the absence of other Labulla species in our analyses. Labulla has at least one autapomorphy, the epigynal lateral lobes (char 141), as well as more than 30 other transformations, which include a cymbium much smaller than the bulb (char 25), radix with both a tailpiece (char 100) and a terminal apophysis (char 104) and an epigynum with atrium and spiral grooves (char 143) (Fig. 10). Given the uncertainty we described above, we feel that our study does not have sufficient evidence to place Labulla in any of the major lineages we describe. To this date, Hormiga &amp; Scharff (2005) is still the only study to include more than one representative of the genus Labulla in a phylogenetic analysis. Future studies should include the other two species of this genus to test its placement further and thus, we prefer to leave it as incertae sedis.</p></div>	https://treatment.plazi.org/id/7D34D83FFFC07155FF70B3504D96CD84	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.		MagnoliaPress via Plazi	Silva-Moreira, Thiago Da;Kulkarni, Siddharth;Hormiga, Gustavo	Silva-Moreira, Thiago Da, Kulkarni, Siddharth, Hormiga, Gustavo (2025): A total evidence phylogenetic analysis of the spider family Linyphiidae (Araneae, Araneoidea). Zootaxa 5685 (1): 1-78, DOI: 10.11646/zootaxa.5685.1.1, URL: https://doi.org/10.11646/zootaxa.5685.1.1
7D34D83FFFC37157FF70B0DC4DD9CD08.text	7D34D83FFFC37157FF70B0DC4DD9CD08.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Pocobletinae Silva-Moreira & Kulkarni & Hormiga 2025	<div><p>Pocobletinae new subfamily (Figs 3, 10 — see TAXONOMY section for details)</p><p>Silva-Moreira &amp; Hormiga (2021) recognized the Pocobletus clade, a lineage that includes Jalapyphantes Gertsch &amp; Davies, Selenyphantes Gertsch &amp; Davies, and Pocobletus Simon. These genera were considered part of Linyphiinae by Tanasevich (2025). In this study, we elevate this group to subfamily rank with original composition and diagnosis of Silva-Moreira &amp; Hormiga (2021).</p><p>This clade is supported by 17 synapomorphies, three of which are non-homoplasious (Fig. 10): a concave-coiled tegulum (char 59), an exposed column (char 86), and the presence of a membranous epigynal sac or channel connecting the copulatory openings to the copulatory ducts (char 150). Its monophyly received full support in all analyses and matrices in this study (Fig. 3, Tables 5 and 6) and in the analyses of Silva-Moreira &amp; Hormiga (2021) (q. v. for a more in-depth analysis of this clade).</p></div>	https://treatment.plazi.org/id/7D34D83FFFC37157FF70B0DC4DD9CD08	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.		MagnoliaPress via Plazi	Silva-Moreira, Thiago Da;Kulkarni, Siddharth;Hormiga, Gustavo	Silva-Moreira, Thiago Da, Kulkarni, Siddharth, Hormiga, Gustavo (2025): A total evidence phylogenetic analysis of the spider family Linyphiidae (Araneae, Araneoidea). Zootaxa 5685 (1): 1-78, DOI: 10.11646/zootaxa.5685.1.1, URL: https://doi.org/10.11646/zootaxa.5685.1.1
7D34D83FFFC67153FF70B1CA4E1FCFD8.text	7D34D83FFFC67153FF70B1CA4E1FCFD8.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Linyphiinae Blackwall 1859	<div><p>Linyphiinae (Figs 4 and 5) new circumscription</p><p>Linyphiinae (sensu Tanasevitch 2025) is not monophyletic in any of our analyses. The genera assigned to this subfamily were found scattered through several different lineages across the tree. The polyphyly of Linyphiinae is neither new (e.g., Wang et al. 2015) nor unexpected, given the very loose definition of the subfamily (Millidge, 1993) and the diverse taxa assigned to it. In this study, we propose a new circumscription for the Linyphiinae subfamily that will include most of the traditionally assigned genera to this subfamily, with some exceptions that would be found mainly within Clade A (see TAXONOMY section for details).</p><p>The newly circumscribed Linyphiinae is recovered in all TE analyses with solid support from both ML and Bayesian analyses. Linyphiinae is supported by five synapomorphies (Fig. 11): a compact lamella characteristica (char 113) with a posterior apophysis (char 119), an epigynum with dorsal plate socket (char 131) or a dorsal plate scape (char 132) without socket (char 134), a cymbium dorsal or meso-dorsal oriented (char 113), and an ellipsoid tegulum (char 59). The compact lamella characteristica (LC) and the scape without socket are non-homoplasious synapomorphies of this clade.</p><p>Labullinyphia tersa (Figs 4, 11)</p><p>Van Helsdingen (1985) considered Labullinyphia tersa, from Sri Lanka, closely related to Labulla (hence the new generic name and combination), Microlinyphia Gerhardt, and Frontinellina van Helsdingen (especially to F. locketi van Helsdingen, 1970), based on similarities of the female genitalia and despite the presence of the single dorsal spine on tibia IV. Millidge (1993) illustrated the internal epigynal structures and provided a short description of them. The aforementioned studies on Labullinyphia were based only on the female, as the male remained unknown. Benjamin &amp; Hormiga (2009) provided the first description of the male and based on a phenotypic cladistic analysis, suggested that Labullinyphia nested within erigonines, as sister to Erigone . The erigonine placement of Labullinyphia was supported by synapomorphies such as the absence of epiandrous fusules, retention of araneoid PLS triplet in adult males, absence of the female pedipalpal claw, desmitracheate system, and the male retrolateral tibial apophysis, which are well-known traits of erigonines. Those results were later corroborated by the analysis of Silva-Moreira &amp; Hormiga (2021).</p><p>Molecular analyses from Wheeler et al. (2017) and the present study (Fig. 4) place L. tersa outside erigonines. The results of our MORPH matrix analysis (both equal and implied weights) are congruent with previous studies that used solely phenotypic data (Benjamin &amp; Hormiga 2009; Silva-Moreira &amp; Hormiga 2022) and place Labullinyphia in Erigoninae . Surprisingly, none of the analyses that used molecular data placed this genus within the erigonines (but on TNT, L. tersa was retrieved as sister group to Labulla thoracica). Irfan &amp; Peng (2019) recently described a second species in the genus from China, morphologically similar to the Sri Lanka species and also exhibiting many erigonine synapomorphies. The lack of congruence between phenotypic and molecular data is intriguing, and given the long list of morphological features shared between Labullinyphia and the erigonines, we suspect that the available sequence data for L. tersa is based on a mislabeling error. Future work will require de novo sequencing representatives of this genus to resolve this paradox. We thus consider Labullinyphia as incertae sedis.</p><p>Linyphia clade (Figs 4, 11)</p><p>The clade that includes Linyphia Latreille is found as one of the lineages of Linyphiinae and comprises, in addition to Linyphia, the genera Microlinyphia, Neriene Blackwall, and Pityohyphantes Simon (Fig. 4). While a lineage with Linyphia, Pityohyphantes, and Microlinyphia is very consistent across analyses, the placement of Neriene in it is much more unstable. Some of our analyses (e.g., M4_RAXML) place all Neriene species together, including three undescribed species, one from the Philippines (Gen. sp. PHY_02) and two from Taiwan (Gen. sp. TW_01 and Gen. sp. TW_02). Ten synapomorphies (all of them homoplasious) support Linyphia clade (Fig. 11), including a pear-shaped male palpal tibia (char 11), a flattened paracymbium (char 52) with a thin medial part (char 51), embolic division with terminal apophysis (char 104) and an epigynum with an atrium (char 142). The implied weighted tree of the MORPH dataset does imply the presence of an atrium with grooves or folds as a non-homoplasious synapomorphy of this clade. This same character in our selected optimization is found as a non-homoplasious synapomorphy of Linyphia + Microlinyphia .</p><p>Frontinella clade</p><p>The Frontinella clade is composed by Frontinella F. O. Pickard-Cambridge, Orsonwelles Hormiga, Neriene japonica (Oi), and one unidentified linyphiid from the Philippines (Gen. sp. PHY_01— Fig. 4). As mentioned above, the placement of Neriene is unstable, and sometimes it is recovered as separate lineage. As we are unable to access the molecular voucher (ICZL 172) of N. japonica we cannot test this hypothesis in more detail. Nevertheless, the Frontinella clade is consistently found in all our analyses, congruent with the results of Arnedo et al. (2009) and Wang et al. (2015), as a natural group supported by 18 synapomorphies, only one of which is non-homoplasious, the presence of a cymbial pocket (char 40). Other synapomorphies include the male palp tibial macrosetae only on the ventral part (char 6), a paracymbium oriented perpendicular to the cymbium (char 47), with the basal section taller than longer (char 50) and medial (char 51)/ apical (char 52) sections absent and a radix with terminal sclerite (char 109) (Fig. 11)</p></div>	https://treatment.plazi.org/id/7D34D83FFFC67153FF70B1CA4E1FCFD8	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.		MagnoliaPress via Plazi	Silva-Moreira, Thiago Da;Kulkarni, Siddharth;Hormiga, Gustavo	Silva-Moreira, Thiago Da, Kulkarni, Siddharth, Hormiga, Gustavo (2025): A total evidence phylogenetic analysis of the spider family Linyphiidae (Araneae, Araneoidea). Zootaxa 5685 (1): 1-78, DOI: 10.11646/zootaxa.5685.1.1, URL: https://doi.org/10.11646/zootaxa.5685.1.1
7D34D83FFFDC7148FF70B7844D3DCD14.text	7D34D83FFFDC7148FF70B7844D3DCD14.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Mynogleninae Lehtinen 1967	<div><p>Mynogleninae and the Australian grade (Figs 6, 12)</p><p>Mynogleninae monophyly has been repeatedly corroborated in a diversity of analyses (e.g., Hormiga 1994, Arnedo et al. 2009, Wang et al. 2015, Silva-Moreira &amp; Hormiga 2022). Frick &amp; Scharff (2013) carried out an extensive phenotypic cladistic analysis on mynoglenines, which included representatives from both Africa and New Zealand and included for the first time Afrotropical mynoglenines. In their hypothesis, the monophyly of Mynogleninae is supported by the following synapomorphies: paracymbium intersegmental membrane covering half of the alveolus, paracymbium with a straight inner margin, a small radix with a proximal part teardrop-shaped almost indistinct from the embolic base, presence of the subocular clypeal sulci, cheliceral setae basis as a distinct bump and at least one ventral distal macroseta on tarsus IV.</p><p>Frick &amp; Scharff (2013) also recognized that a partially sclerotized embolic membrane attached to the tegulum could be an apomorphy of mynoglenines. Our interpretation of the morphology follows Silva-Moreira &amp; Hormiga (2022), where the tegular membrane was not considered homologous to the embolic membrane. In our hypothesis (Fig. 1), Mynogleninae monophyly is supported by 17 synapomorphies, but only one, the presence of subocular clypeal sulci (char 168) in both sexes, is non-homoplasious. Additional homoplasious transformations under our preferred hypothesis include the absence of macrosetae on the male palpal tibia (char 4), the absence of an embolic membrane (char 87), and juxtaposed pedicel sternites on males (char 227) (Fig. 12).</p><p>Mynoglenines are nested within a clade that includes Australolinyphia, Laperousea, two species of Laetesia ( Laetesia raveni and Laetesia sp _NSW), and Palaeohyphantes . To make easier to reference, we have named all the non-mynoglenines in this clade as the “Australian grade.” Palaeohyphantes simplicipalpis is the sister group of Mynogleninae instead of Australolinyphia remota, as in previous studies (Arnedo et al. 2009, Wang et al. 2015). Palaeohyphantes + Mynogleninae is supported by seven synapomorphies, including two or more cymbial macrosetae (char 29), no cymbium retrolateral lobe (char 43), a pear-shaped tegulum (char 59), a curved embolus (char 91), and the absence of dorso-ectal macrosetae on female palp (char 193).</p><p>A third species of Laetesia (“ Laetesia ” sp MAA 2009) is the sister group of the MCP clade (Marginal Cephalothoracic Pits—see below). This species is the terminal labeled as “ Laetesia ” in Arnedo et al. (2009) study. The genus Laetesia Simon currently has 25 species described (WSC, 2025), but with a confusing taxonomy and diagnostic features loosely defined. As a thorough exploration of the taxonomy of Laetesia is far from the scope of this study, we decided to keep Arnedo et al. ’s (2009) tentative generic placement for this specimen in question.</p><p>In our optimization, the Mynogleninae + Australian grade lineage is supported by four synapomorphies: a suprategulum distal in relation to tegulum (char 76), the foramen on the tegulum instead of the suprategulum (char 77), absence of a lamella characteristica (char 111), and a cylindrical or ellipsoid opisthosoma (char 214). All components of the Australian grade are here considered incertae sedis until further better evidence of their placement to be collected and analyzed.</p></div>	https://treatment.plazi.org/id/7D34D83FFFDC7148FF70B7844D3DCD14	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.		MagnoliaPress via Plazi	Silva-Moreira, Thiago Da;Kulkarni, Siddharth;Hormiga, Gustavo	Silva-Moreira, Thiago Da, Kulkarni, Siddharth, Hormiga, Gustavo (2025): A total evidence phylogenetic analysis of the spider family Linyphiidae (Araneae, Araneoidea). Zootaxa 5685 (1): 1-78, DOI: 10.11646/zootaxa.5685.1.1, URL: https://doi.org/10.11646/zootaxa.5685.1.1
7D34D83FFFDE714AFF70B6FC4D2CCD70.text	7D34D83FFFDE714AFF70B6FC4D2CCD70.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Micronetinae Hull 1920	<div><p>“ Micronetinae ”</p><p>Micronetinae Hull, 1920 was circumscribed by Saaristo &amp;Tanasevitch (1996) and diagnosed based on their complex embolic division (which they described in detail) and the presence of the outward pointing suprategular apophysis just below the tip of the cymbium in the unexpanded palp. Among their genitalic features, micronetines have an enlarged U-shaped paracymbium, with pockets and an enlargement of the seminal duct prior to the entrance on the embolus proper (Fickert’s gland). Although these diagnostic characters were carefully described and illustrated, Saaristo &amp; Tanasevitch (1996) did not provide a phylogenetic justification to their concept of Micronetinae .</p><p>None of our analyses support the monophyly of Micronetinae sensu Saaristo and Tanasevitch (1996) (see T able 5). In that respect, our results are consistent with previous studies (e.g., Arnedo et al. 2009; Wang et al. 2015; Zhao and Li 2017). No phylogenetic analysis with a sufficient taxon sample has ever supported the monophyly of micronetines. The “micronetine” representatives in our analyses are placed across four different lineages: Centromerus clade, Lepthyphantinae, Agyneta clade, and Microneta clade, in addition to the isolated Allomengea lineage. All those clades, except for the Microneta clade, were recovered with full support in all our large and small datasets analyses. The Centromerus clade (clade F in Wang et al. 2015) and the Lepthyphantinae (Clade A in Arnedo et al. 2009, and Clade A in Wang et al. 2015) are the first of the “micronetine branches” of Clade B (Fig. 7).</p><p>Centromerus clade (Fig. 7)</p><p>The Centromerus clade is a small lineage composed of only three genera: Centromerus Dahl; Saaristoa Millidge; and Doenitzius Oi. Both Saaristoa and Doenitzius are small genera (five and two species, respectively) and (with the exception of S. sammamish (Levi &amp; Levi)) restricted to the Palearctic and Sino-Japanese regions (sensu Holt et al. 2013). After the original descriptive work, few studies have addressed these genera. While there is no formal taxonomic revision of Centromerus, there have been occasional species transfers and nomenclatural acts concerning the species of the genus. Millidge (1977) placed Centromerus within his Lepthyphantes group based on the conformation of the seminal ducts.</p></div>	https://treatment.plazi.org/id/7D34D83FFFDE714AFF70B6FC4D2CCD70	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.		MagnoliaPress via Plazi	Silva-Moreira, Thiago Da;Kulkarni, Siddharth;Hormiga, Gustavo	Silva-Moreira, Thiago Da, Kulkarni, Siddharth, Hormiga, Gustavo (2025): A total evidence phylogenetic analysis of the spider family Linyphiidae (Araneae, Araneoidea). Zootaxa 5685 (1): 1-78, DOI: 10.11646/zootaxa.5685.1.1, URL: https://doi.org/10.11646/zootaxa.5685.1.1
7D34D83FFFDE714BFF70B2704EFFC998.text	7D34D83FFFDE714BFF70B2704EFFC998.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Lepthyphantinae Simon 1929	<div><p>Lepthyphantinae new circumscription and Fickert’s gland homology (Figs 7, 12)</p><p>Lepthyphantinae new circumscription is a very speciose lineage with more than 400 species (WSC 2025), concentrating about 70% of the diversity of the polyphyletic micronetines (sensu Saaristo &amp; Tanasevitch 1996).</p><p>Saaristo’s (1975) Lepthyphantinae [partim] and Millidge’s (1977) Lepthyphantes group were later referred to as Micronetinae (Saaristo &amp; Tanasevitch 1996), based on the presence of the Fickert’s gland in the male palp. In our analysis, the Lepthyphantinae (new circumscription) is supported by ten synapomorphies. The presence of a special seta on the patella of the male palp (char 126) is recovered as non-homoplasious, while the remaining changes, such as the presence of an ecto-basal cymbial process (char 33), a paracymbium of about the same size as the rest of the cymbium (char 46), paracymbium with an apophysis or process as a distal branch of its apical part (char 52), a sperm duct switchback (char 71), and an epigynum with a ventral plate scape (char 136) optimized as homoplastic.</p><p>The Fickert’s gland (char 103) was not recovered as a synapomorphy for any of the “micronetinae” clades, similar to the results of Arnedo et al. (2009). Our ancestral reconstruction (Fig. 14A) analysis suggests at least two independent origins of Fickert’s gland: one in the Lepthyphantinae, found in Lepthyphantes Menge Tenuiphantes Saaristo &amp; Tanasevich Bolyphantes Koch, and Drapetisca Menge with a loss in Floronia Simon and a second origin within the Agyneta clade ( A. rurestris + A. ramosa Jackson). The position of Floronia (together with Tapinopa Westring) as sister group to the remaining Lepthyphante s group receives full support across all our analyses. Both Floronia and Tapinopa lack Fickert’s gland, and for this reason, Millidge (1977) placed them in the Oreonetides group.</p><p>Based on Merret’s (1963) illustrations, Saaristo &amp; Tanasevitch (1996) noticed that the position of Fickert’s gland in Agyneta was different from that in the rest of Lepthyphantinae (sensu Saaristo, 1999), appearing in the former as a “small secondary dilatation within the embolus [proper]” instead of a larger one within the radix in the latter (Figs 14B, C). While we made no distinction between the two morphologies of the Fickert’s gland when coding them in our matrices, our results suggest that those structures might have separate origins and, thus, not be homologous. A detailed study of the structure of Fickert’s gland in both groups is needed to understand the evolution of this structure better.</p></div>	https://treatment.plazi.org/id/7D34D83FFFDE714BFF70B2704EFFC998	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.		MagnoliaPress via Plazi	Silva-Moreira, Thiago Da;Kulkarni, Siddharth;Hormiga, Gustavo	Silva-Moreira, Thiago Da, Kulkarni, Siddharth, Hormiga, Gustavo (2025): A total evidence phylogenetic analysis of the spider family Linyphiidae (Araneae, Araneoidea). Zootaxa 5685 (1): 1-78, DOI: 10.11646/zootaxa.5685.1.1, URL: https://doi.org/10.11646/zootaxa.5685.1.1
7D34D83FFFD07144FF70B4644EB4CFD8.text	7D34D83FFFD07144FF70B4644EB4CFD8.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Ipainae	<div><p>Ipainae</p><p>Saaristo (2007) erected the subfamily Ipainae to group the genera Ipa Saaristo Epibellowia Tanasevitch, Metaleptyphantes Locket, Solenysa Simon, Uralophantes Esyunin, Wubanoides Eskov, and Epigytholus Tanasevitch. Saaristo (2007) enumerated a long list of diagnostic features for the subfamily, including the radical lamella, “the shape of the paracymbium” [sic], a filiform embolus, and a movable epigynum “whereas the first and last are possible autapomorphies of the entire family” (Saaristo 2007:38). In the same study, the author also hypothesized that Ipainae would be the sister group to Micronetinae as they share the presence of Fickert’s gland in the male palp (Saaristo 2007). However, as pointed by Arnedo et al. (2009), none of those assertions were tested as characters in phylogenetic analysis, and thus, the monophyly of Ipainae was regarded by them with some skepticism. The phenotypic analysis of Tu &amp; Hormiga (2011) placed Solenysa within the “Distal erigonines” clade. A subsequent study, based entirely on molecular data (Wang et al. 2015), included for the first time more than one ipaine genus ( Solenysa and Wubanoides) and did not support the monophyly of Ipainae . Silva-Moreira &amp; Hormiga (2022) coded Ipa keyserlingi (Ausserer) together with some of the species of Solenysa used in Tu &amp; Hormiga (2011) and tested the monophyly of Ipainae for the first time in a cladistic framework, with results similar to those of Wang et al. (2015), rejecting again Ipainae monophyly but this time with a phenotypic dataset.</p><p>We have tested the monophyly of Ipainae in a total evidence framework. We could not obtain Ipa specimens suitable for sequencing, so our hypothesis of placement for this genus is based on phenotypic data. Our hypothesis is congruent with those of Wang et al. (2015) and Silva-Moreira &amp; Hormiga (2022). None of our analyses recovered Ipainae as a clade (see Table 5). Ipa position varied widely depending on the dataset and optimality criteria (always with very poor nodal support—see Supplementary files) but never as sister to Solenysa or Wubanoides, being the latter consistently recovered as sister to Parawubanoides within the Lepthyphantinae (Fig. 7).</p><p>Finally, the position of Solenysa was strongly influenced by the inclusion of molecular data. The MORPH dataset places Solenysa in Erigoninae, sister to Gonatium rubens (Fig. 12), while both DNA and TE datasets place Solenysa genus in Clade B, sister to a clade comprising Allomengea + Agyneta clade + Microneta clade + Erigoninae (Fig. 8) a result similar to those of Wang et al. (2015). In both cases, Solenysa is strongly supported as a natural group. The genus is supported by five non-homoplasious synapomorphies: lamella characteristica posterior apophysis flattened (char 120), epigynum attached to opisthosoma trough a solenoid (char 128), clypeus with pits (char 172), prosomal pits (char 198) and prosomal petiole (char 199).</p><p>While the placement of the different genera of Ipainae is not fully resolved, the data strongly suggest that this subfamily is not a natural group. The distinctive features pointed out by Saaristo (2007) are very likely convergent features, as suggested by our MORPH dataset. The Ipainae subfamily should not be used as a taxonomic rank as it refers to a non-natural group.</p></div>	https://treatment.plazi.org/id/7D34D83FFFD07144FF70B4644EB4CFD8	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.		MagnoliaPress via Plazi	Silva-Moreira, Thiago Da;Kulkarni, Siddharth;Hormiga, Gustavo	Silva-Moreira, Thiago Da, Kulkarni, Siddharth, Hormiga, Gustavo (2025): A total evidence phylogenetic analysis of the spider family Linyphiidae (Araneae, Araneoidea). Zootaxa 5685 (1): 1-78, DOI: 10.11646/zootaxa.5685.1.1, URL: https://doi.org/10.11646/zootaxa.5685.1.1
7D34D83FFFD07145FF70B3184DFFCED4.text	7D34D83FFFD07145FF70B3184DFFCED4.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Erigoninae Emerton 1882	<div><p>Erigoninae (Figs 9, 12)</p><p>Erigonines, commonly known as “dwarf spiders” or “money spiders,” is one of the oldest recognized groups within linyphiids. The name Erigoninae goes back to Simon’s (1884: 181) “erigonine,” and this subfamily is the most diverse linyphiid group. While most linyphiids have relatively uniform somatic morphology, many male erigonines exhibit a great diversity of cephalic modifications (Hormiga, 2000). This diversity may have contributed at some point to their rank as a family— Erigonidae (Gerhardt 1923) or Micryphantidae (Bertkau 1878; Wiehle 1956, 1960). Some of the largest analyses of linyphiid phylogenetic relationships have the erigonines as the main focus (Hormiga 2000; Miller &amp; Hormiga 2004; Frick et al. 2010; Wang et al. 2015; Arnedo &amp; Hormiga 2021; Lin et al. 2022). Erigonines are also the most diverse, species-rich group of linyphiids. Erigonines correspond to “Clade B” (partim) of Arnedo et al. (2009) and Wang et al. (2015). The monophyly of erigonines has been corroborated by phenotypic data (e.g., Hormiga 2000, Miller &amp; Hormiga 2004 Tu &amp; Hormiga 2011, Silva-Moreira &amp; Hormiga 2022, Lin et al. 2022), but with the inclusion of molecular data (Arnedo et al. 2009, Wang et al. 2015), classic erigonines form an unresolved clade with Microneta, Agyneta, Helophora (among other genera, see Wang et al. (2015)) nested within it. Arnedo et al. (2009, Fig. 9) do report that one of the trees that was used in forming the consensus of the hypothesis tree finds Erigoninae monophyletic, but all other analyses were congruent with Wang et al. (2015), recovering an “Erigonines + Micronetines” clade.</p><p>In our preferred hypothesis, erigonines are recovered as a natural group (Fig. 9), and split in two major lineages . The first, the “ Neomaso clade,” contains Macrargus Dahl, Gongylidiellum Simon, Hilaira Simon, Millidgella Kammerer, Ostearius Hull, Laminacauda Millidge, and Neomaso Forster. The second is the “distal erigonines” clade (sensu Wang et al. 2015), the largest clade within Erigoninae in which Erigone Audouin is nested. Even considering the small taxon overlap between this study and Hormiga’s (2000) and Miller &amp; Hormiga’s (2004), our results are congruent with the phenotypic analyses of those latter two studies. Erigoninae monophyly received, for the first time, the support of a large-scale total evidence analysis. The Neomaso clade is supported by seven synapomorphies, including the presence of a retrolateral groove in the cymbium (char 42), a paracymbium with a blunt edge (char 52), an embolic membrane without papillae (char 88), median (char 159) and lateral (char 160) eyes clustered and tracheolar taenidia (char 237). The Distal Erigonines lineage is supported by eight synapomorphies, including the branched medial tracheal trunks (char 234), much wider than the lateral ones (char 235), and invading the prosoma through the pedicel (char 236).</p><p>Some of our analyses recovered results similar to those of Arnedo et al. (2009) and Wang et al. (2015), where the Microneta clade is nested within the Neomaso clade or with some particular genera (notably Gongylidiellum) recovered within the Microneta clade. All those topologies result from datasets that include the 28S sequences. As we discussed earlier, the inclusion of the 28S data results in topologies that contradict highly corroborated lineages. Given that we have considered those results artifactual and the finding of a monophyletic Erigoninae with strong phenotypic support, we regard the paraphyly of Erigonines in some analyses as another artifact caused by the inclusion of a marker with extreme size disparity. In our optimizations, Erigoninae is supported by 11 synapomorphies (all homoplastic), including the absence of macrosetae on the male palp tibia (char 6), male palp with a prolateral tibial apophysis (char 15), distal suprategular apophysis extending ventrally from the suprategulum (char 79) and females with a single clypeal seta below the AME (char 173).</p><p>None of the classic phenotypic diagnostic features of erigonines, such as the clawless female pedipalp (char 189) or the presence of a retrolateral tibial apophysis (char 17) in the male pedipalp, optimized as synapomorphies for this clade. This is due to the fact that those characters are being tested for the first time in a wider taxonomic context. Taxa previously placed in the erigonines based (in part) on those two characters, such as Labullinyphia tersa (Benjamin &amp; Hormiga, 2009) or Solenysa (Tu &amp; Hormiga 2011; contra Saaristo 2007), fall somewhere else in the tree once the molecular data are included.</p></div>	https://treatment.plazi.org/id/7D34D83FFFD07145FF70B3184DFFCED4	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.		MagnoliaPress via Plazi	Silva-Moreira, Thiago Da;Kulkarni, Siddharth;Hormiga, Gustavo	Silva-Moreira, Thiago Da, Kulkarni, Siddharth, Hormiga, Gustavo (2025): A total evidence phylogenetic analysis of the spider family Linyphiidae (Araneae, Araneoidea). Zootaxa 5685 (1): 1-78, DOI: 10.11646/zootaxa.5685.1.1, URL: https://doi.org/10.11646/zootaxa.5685.1.1
7D34D83FFFD27146FF70B2934D56CDC7.text	7D34D83FFFD27146FF70B2934D56CDC7.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Linyphiidae Blackwall 1859	<div><p>Family Linyphiidae Blackwall, 1859</p><p>Linyphiidae Blackwall, 1859: 261 .</p><p>Type genus: Linyphia Latreille, 1804 .</p></div>	https://treatment.plazi.org/id/7D34D83FFFD27146FF70B2934D56CDC7	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.		MagnoliaPress via Plazi	Silva-Moreira, Thiago Da;Kulkarni, Siddharth;Hormiga, Gustavo	Silva-Moreira, Thiago Da, Kulkarni, Siddharth, Hormiga, Gustavo (2025): A total evidence phylogenetic analysis of the spider family Linyphiidae (Araneae, Araneoidea). Zootaxa 5685 (1): 1-78, DOI: 10.11646/zootaxa.5685.1.1, URL: https://doi.org/10.11646/zootaxa.5685.1.1
7D34D83FFFD37147FF70B59C4A32C90C.text	7D34D83FFFD37147FF70B59C4A32C90C.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Erigoninae Emerton 1882	<div><p>Subfamily Erigoninae Emerton, 1882</p><p>Erigoninae Emerton, 1882, Locket &amp; Millidge 1953, Millidge 1984, 1993,</p><p>Gonatini + Lophocarenini + Walckenaerini, Simon,1884.</p><p>Erigoninae + Gonatinae Petrunkevitch, 1928</p><p>Donacocharae [ad partem] + Erigoninae + Walckenaerinae Wiehle 1956</p><p>Erigoninae + Walckenaerinae Merrett 1963, 1965</p><p>Type genus: Erigone Audouin, 1826 .</p><p>Diagnosis (after Miller 2007): Members of the Erigoninae are distinguished from other linyphiids by the absence of epiandrous gland spigots, by the absence of the tarsal claw in the female pedipalp, the absence of prolateral macrosetae on the femur and a reduced number of setae on the clypeus (Miller 2007, fig 7, 18).</p><p>Composition: In our study, Erigoninae includes, in addition to the type genus the following genera: Asperthorax Oi, Ceratinella Emerton, Collinsia Pickard-Cambridge, Concavocephalus Eskov, Dicymbium Menge, Diplocentria Hull, Gnathonarium Karsch, Gonatium Menge, Gongylidiellum Simon, Gongylidioides Oi, Hilaira Simon, Hylyphantes Simon, Hypselistes Simon, Laminacauda Millidge, Lasiargus Kulczyński, Maso Simon, Micrargus Dahl, Millidgella Kammerer, Moebelia Dahl, Nematogmus Simon, Neomaso Forster, Oedothorax Bertkau, Oia Wunderlich, Oreoneta Kulczyński, Ostearius Hull, Paikiniana Eskov, Rhabdogyna Millidge, Savignia Blackwall, Scotinotylus Simon, Sisicottus Bishop &amp; Crosby, Sphecozone Pickard-Cambridge, Styloctetor Simon, Ummeliata Strand, and Walckenaeria Blackwall. For additional members of this subfamily please refer to Miller (2007).</p></div>	https://treatment.plazi.org/id/7D34D83FFFD37147FF70B59C4A32C90C	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.		MagnoliaPress via Plazi	Silva-Moreira, Thiago Da;Kulkarni, Siddharth;Hormiga, Gustavo	Silva-Moreira, Thiago Da, Kulkarni, Siddharth, Hormiga, Gustavo (2025): A total evidence phylogenetic analysis of the spider family Linyphiidae (Araneae, Araneoidea). Zootaxa 5685 (1): 1-78, DOI: 10.11646/zootaxa.5685.1.1, URL: https://doi.org/10.11646/zootaxa.5685.1.1
7D34D83FFFD37147FF70B67E4AF7CDC6.text	7D34D83FFFD37147FF70B67E4AF7CDC6.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Lepthyphantinae Simon 1929	<div><p>Subfamily Lepthyphantinae Simon, 1929 new circumscription</p><p>Linyphiini [ad partem] Simon, 1984,</p><p>Lepthyphantes [ad partem] Petrunkevitch 1928,</p><p>Linyphiinae [ad partem] Locket &amp; Millidge 1953,</p><p>Leptyphantae Wiehle 1956,</p><p>Syedrulae Merrett 1963, 1965 [ad partem],</p><p>Lepthyphantinae Millidge 1984, I983,</p><p>Type genus: Lepthyphantes Menge, 1866 .</p><p>Diagnosis: Male members of the subfamily Lepthyphantinae are diagnosed by the presence of a special seta on the patella of the palp (Roberts 1987, figs 72B, 75A–B, 77C; Hormiga 1994b, fig. 12A), a paracymbium of about the same size as the rest of the cymbium, with a distal apophysis or process (Hormiga 1994b, figs 12A, 13A), and a sperm duct switchback (Hormiga 1994b, fig. 12C), while females have an epigynum with a ventral plate scape (Roberts 1987, figs 75A–D).</p><p>N.B. The taxonomic history and composition of Lepthyphantinae are intermingled with that of Micronetinae, most obviously expressed in the re-delimitation of the latter subfamily by the detailed studies of Saaristo &amp; Tanasevitch (1996), which included Lepthyphantes and allied genera in their circumscription of Micronetinae . Our phylogenetic analyses conclude that Lepthyphantes and Microneta belong to different clades, and thus Micronetinae sensu Saaristo &amp; Tanasevitch (1996) is not monophyletic. Resolving monophyletic groups to which the Linnaean labels ‘Lepthyphantinae’ and ‘Micronetinae’ can be applied entails significant changes in their circumscription. As noted, the classification proposal presented here is limited by the taxa included in our analyses. Additional work is required to address all the genera that had been formerly classified under Micronetinae .</p><p>Composition: As re-circumscribed here, the composition of this subfamily is similar to Saaristo’s (1975) Lepthyphantinae, Millidge’s (1977) Lepthyphantes group + Oreonetide s group [partim], Micronetinae (Saaristo &amp; Tanasevitch, 1996) [partim], Clade A in Arnedo et al. (20009), and Clade A in Wang et al. (2015). In addition to the type genus, Lepthyphantinae is represented in our study by the following genera: Acanthoneta Eskov &amp; Marusik, Anguliphantes Saaristo &amp; Tanasevitch, Arcuphantes Chamberlin &amp; Ivie, Bifurcia Saaristo, Tu &amp; Li, Bolyphantes Koch, Crispiphantes Tanasevitch, Denisiphantes Tu, Li &amp; Rollard, Drapetisca Menge, Eldonnia Tanasevitch, Floronia Simon, Fusciphantes Oi, Himalaphantes Tanasevitch, Incestophantes Tanasevitch, Ipa Saaristo, Lidia Saaristo &amp; Marusik, Macrargus Dahl, Mughiphantes Saaristo &amp; Tanasevitch, Nippononeta Eskov, Oryphantes Hull, Parawubanoides Eskov &amp; Marusik, Poeciloneta Kulczyński, Syedra Simon, Tapinopa Westring, Tenuiphantes Saaristo &amp; Tanasevitch, Ternatus Sun, Li &amp; Tu, Vagiphantes Saaristo &amp; Tanasevitch, and Wubanoides Eskov (Table 8).</p></div>	https://treatment.plazi.org/id/7D34D83FFFD37147FF70B67E4AF7CDC6	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.		MagnoliaPress via Plazi	Silva-Moreira, Thiago Da;Kulkarni, Siddharth;Hormiga, Gustavo	Silva-Moreira, Thiago Da, Kulkarni, Siddharth, Hormiga, Gustavo (2025): A total evidence phylogenetic analysis of the spider family Linyphiidae (Araneae, Araneoidea). Zootaxa 5685 (1): 1-78, DOI: 10.11646/zootaxa.5685.1.1, URL: https://doi.org/10.11646/zootaxa.5685.1.1
7D34D83FFFD47140FF70B59C4E98C903.text	7D34D83FFFD47140FF70B59C4E98C903.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Linyphiinae Blackwall 1859	<div><p>Subfamily Linyphiinae Blackwall, 1859 new circumscription</p><p>Linyphiinae Blackwall, 1859 (ad partem),</p><p>Linyphieae [ad partem] Merrett 1963, 1965</p><p>Linyphiinae Millidge, 1977 [ad partem]</p><p>Linyphiinae Millidge 1984, I983</p><p>Type genus: Linyphia Latreille, 1804 .</p><p>Dubiaraneinae Millidge, 1993 (new synonymy). Type Genus: Dubiaranea Mello-Leitão, 1943 .</p><p>Diagnosis: All members of Linyphiinae are diagnosed by a compact characteristic lamella with an apophysis on the male palp (Silva-Moreira &amp; Hormiga 2022, figs 6f, 14f) and an epigynum with a dorsal plate scape (Millidge 1991, figs 42, 51, 58) or socket (Millidge 1991, figs 13, 21).</p><p>Composition: Linyphiinae new circumscription includes a fraction of the subfamilies Linyphiinae and the entirety of Dubiaraneinae (both sensu Millidge 1993): “clade D” of Wang et al. (2015) and the MPME clade (sensu Silva-Moreira &amp; Hormiga 2021). In this study, they are represented by the type genus, plus representatives of the following genera: Diplothyron Millidge, Dubiaranea Mello-Leitao, Eurymorion Millidge, Frontinella, F.O. Pickard-Cambridge, Lomaita Bryant, Microlinyphia Gerhardt, Neriene Blackwall, Notiohyphantes Millidge, Novafrontina Millidge, Orsonwelles Hormiga, Pityohyphantes Simon, and Plectembolus Millidge &amp; Russell-Smith, as well as six undescribed genera (Table 8). The exact composition of Linyphiinae requires a more extensive study which is beyond the scope of this paper.</p></div>	https://treatment.plazi.org/id/7D34D83FFFD47140FF70B59C4E98C903	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.		MagnoliaPress via Plazi	Silva-Moreira, Thiago Da;Kulkarni, Siddharth;Hormiga, Gustavo	Silva-Moreira, Thiago Da, Kulkarni, Siddharth, Hormiga, Gustavo (2025): A total evidence phylogenetic analysis of the spider family Linyphiidae (Araneae, Araneoidea). Zootaxa 5685 (1): 1-78, DOI: 10.11646/zootaxa.5685.1.1, URL: https://doi.org/10.11646/zootaxa.5685.1.1
7D34D83FFFD47140FF70B6844BFECEE0.text	7D34D83FFFD47140FF70B6844BFECEE0.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Mynogleninae Lehtinen 1967	<div><p>Subfamily Mynogleninae Lehtinen, 1967</p><p>Mynogleninae Lehtinen, 1967, Millidge 1894, 1993, Tanasevich, 2025.</p><p>Type genus: Haplinis Simon.</p><p>Diagnosis: Mynoglenines are diagnosed by the presence of clypeal sulci (or pits) with cuticular pores served by gland, present in both sexes as well as juveniles (Blest, 1979; Hormiga, 1994; Arnedo et al. 2009, Frick &amp; Scharff 2014).)</p><p>Composition: In this study, Mynoglenines are represented by the genera Haplinis, Novafroneta, Parafroneta, and Pseudafroneta . For additional members of this clade, refer to Frick and Scharff (2014).</p></div>	https://treatment.plazi.org/id/7D34D83FFFD47140FF70B6844BFECEE0	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.		MagnoliaPress via Plazi	Silva-Moreira, Thiago Da;Kulkarni, Siddharth;Hormiga, Gustavo	Silva-Moreira, Thiago Da, Kulkarni, Siddharth, Hormiga, Gustavo (2025): A total evidence phylogenetic analysis of the spider family Linyphiidae (Araneae, Araneoidea). Zootaxa 5685 (1): 1-78, DOI: 10.11646/zootaxa.5685.1.1, URL: https://doi.org/10.11646/zootaxa.5685.1.1
7D34D83FFFD47140FF70B0244CD2CD49.text	7D34D83FFFD47140FF70B0244CD2CD49.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Pocobletinae Silva-Moreira & Kulkarni & Hormiga 2025	<div><p>Subfamily Pocobletinae Silva-Moreira &amp; Hormiga, new subfamily</p><p>Erigoninae [ad partem] + Linyphiinae [ad partem] Petrunkevitch 1928</p><p>Stemonyphantes group (Millidge, 1977, 1984) [ad partem]</p><p>Pocobletus clade. Silva-Moreira &amp; Hormiga 2021</p><p>Type genus: Pocobletus Simon, 1894</p><p>Diagnosis: Pocobletines are diagnosed by a concave-coiled tegulum (versus a diversity of linyphiid tegular morphologies, such as globular and flattened tegulum; e.g., Hormiga 2000: fig. 11), an exposed column in the male palp (Silva-Moreira and Hormiga 2021, figs 14; 32; 33; 43) (versus a hidden column in the unexpanded palp; e.g., see Hormiga 2000: figs 1, 13, 21) and, in females, a membranous sac or channel connecting the epigynal copulatory opening to the copulatory duct (Silva-Moreira and Hormiga 2021, figs 17A–E; 32; 37; 44) (versus the widespread absence of such sac; e.g., Hormiga &amp; Scharff 2005: figs 7B, 10D, 13B, C).</p><p>Composition: The subfamily Pocobletinae (Fig. 3), referred to as the Pocobletus clade in Silva-Moreira &amp; Hormiga (2021),comprises in addition to the type genus the genera Jalapyphantes Gertsch &amp; Davis and Selenyphantes Gertsch &amp; Davis (Table 8).</p><p>Distribution: Southern North America, Central and South America.</p></div>	https://treatment.plazi.org/id/7D34D83FFFD47140FF70B0244CD2CD49	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.		MagnoliaPress via Plazi	Silva-Moreira, Thiago Da;Kulkarni, Siddharth;Hormiga, Gustavo	Silva-Moreira, Thiago Da, Kulkarni, Siddharth, Hormiga, Gustavo (2025): A total evidence phylogenetic analysis of the spider family Linyphiidae (Araneae, Araneoidea). Zootaxa 5685 (1): 1-78, DOI: 10.11646/zootaxa.5685.1.1, URL: https://doi.org/10.11646/zootaxa.5685.1.1
7D34D83FFFD57141FF70B1434E40CFD3.text	7D34D83FFFD57141FF70B1434E40CFD3.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Linyphiidae Blackwall 1859	<div><p>Linyphiidae genera included in this study considered incertae sedis</p><p>Allomengea Strand, Australolinyphia Wunderlich, Labulla Simon, Labulinyphia van Helsdingen, Laetesia Simon, Laperousea Dalmas, Palaeohyphantes Millidge, Solenysa Simon, Turinyphia van Helsdingen.</p><p>Supplementary Files</p><p>All supplementary files can be found at Zenodo (https://doi.org/10.5281/zenodo.13134490). The package includes a printed version of the phenotypic matrix, a list of characters with descriptions and comments (both as PDF files), the five gene alignments (as FASTA files), all matrices used (as NEXUS files), and all trees generated in this study (as.tre files).</p></div>	https://treatment.plazi.org/id/7D34D83FFFD57141FF70B1434E40CFD3	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.		MagnoliaPress via Plazi	Silva-Moreira, Thiago Da;Kulkarni, Siddharth;Hormiga, Gustavo	Silva-Moreira, Thiago Da, Kulkarni, Siddharth, Hormiga, Gustavo (2025): A total evidence phylogenetic analysis of the spider family Linyphiidae (Araneae, Araneoidea). Zootaxa 5685 (1): 1-78, DOI: 10.11646/zootaxa.5685.1.1, URL: https://doi.org/10.11646/zootaxa.5685.1.1
7D34D83FFFD57141FF70B59C4B19C9CA.text	7D34D83FFFD57141FF70B59C4B19C9CA.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Stemonyphantinae Wunderlich 1986	<div><p>Subfamily Stemonyphantinae Wunderlich, 1986</p><p>Linyphiinae Locket &amp; Millidge 1953 [ad partem],</p><p>Linyphieae Merrett 1963, 1965 [ad partem],</p><p>Stemonyphantes group (Millidge, 1977, 1984) [ad partem],</p><p>Stemonyphantinae Wunderlich, 1986, Hormiga et al. 2021,</p><p>Type genus Stemonyphantes Menge, 1866 .</p><p>Diagnosis (updated from Hormiga et al. 2021): Male stemonyphantines are distinguished from other linyphiids by the presence on the tegulum of a conductor (absent in all other linyphiids), and sometimes a median apophysis (also absent in all other linyphiids), and an integral or partially integral paracymbium. The apical region of the cymbium of most stemonyphantines is either narrow and elongated ( Stemonyphantes) or conical ( Weintrauboa, Pecado, Putaoa) (e.g., see figures in Blauvelt (1936), Merrett (1963), van Helsdingen (1968), and Gavish-Regev et al. (2013)). Composition: Stemonyphantinae includes, in addition to the type genus, the genera Pecado, Putaoa, and Weintrauboa .</p><p>Unranked linyphiid clades</p><p>ConoSur clade— Notholepthyphantes Millidge, Patagoneta Millidge, Juanfernandezia Koçak &amp; Kemal.</p><p>MCP clade— Bathyphantes Menge, Diplostyla Emerton, Kaestneria Wiehle, Porrhomma Simon, Vesicapalpus Millidge.</p><p>Centromerus clade— Centromerus Dahl, Doenitzius Oi, Saaristoa Millidge.</p><p>Agyneta clade— Agyneta Hull, Helophora Menge, Nippononeta Eskov, Parameioneta Locket.</p><p>Microneta clade— Asthenargus Simon &amp; Fage, Microneta Menge, Ryojius Saito &amp; Ono.</p></div>	https://treatment.plazi.org/id/7D34D83FFFD57141FF70B59C4B19C9CA	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.		MagnoliaPress via Plazi	Silva-Moreira, Thiago Da;Kulkarni, Siddharth;Hormiga, Gustavo	Silva-Moreira, Thiago Da, Kulkarni, Siddharth, Hormiga, Gustavo (2025): A total evidence phylogenetic analysis of the spider family Linyphiidae (Araneae, Araneoidea). Zootaxa 5685 (1): 1-78, DOI: 10.11646/zootaxa.5685.1.1, URL: https://doi.org/10.11646/zootaxa.5685.1.1
7D34D83FFFA07134FF70B7844DD0C928.text	7D34D83FFFA07134FF70B7844DD0C928.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Linyphiidae Blackwall 1859	<div><p>Linyphiidae subdivisions sensu Simon (1884)</p><p>(N.B. Simon considered linyphiids as one of the six subdivisions of the family Theridionidae, which is roughly equivalent to modern Theridiidae + Linyphiidae)</p></div>	https://treatment.plazi.org/id/7D34D83FFFA07134FF70B7844DD0C928	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.		MagnoliaPress via Plazi	Silva-Moreira, Thiago Da;Kulkarni, Siddharth;Hormiga, Gustavo	Silva-Moreira, Thiago Da, Kulkarni, Siddharth, Hormiga, Gustavo (2025): A total evidence phylogenetic analysis of the spider family Linyphiidae (Araneae, Araneoidea). Zootaxa 5685 (1): 1-78, DOI: 10.11646/zootaxa.5685.1.1, URL: https://doi.org/10.11646/zootaxa.5685.1.1
7D34D83FFFA07136FF70B3CC4DBDCE64.text	7D34D83FFFA07136FF70B3CC4DBDCE64.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Linyphiidae Blackwall 1859	<div><p>Linyphiidae subdivisions sensu Petrunkevitch (1928)</p><p>N.B. Elaphopus, as a replacement name for Elaphidion, should be also a synonym of Saloca Simon. However, both Roewer and the WSC overlooked the fact that Elaphopus is an available name and, as such, competes for synonymy, being senior to Saloca Simon by four decades. Prószyński &amp; Staręga (1971) avoided the collision by considering both genera separated “The genus Elaphopus Menge includes only the discussed species [ Elaphopus flagellifer] and is probably closely related to the genus Saloca Simon Simon ”, which might be interpreted as the undoing of the synonymy between those two genera. Since a deeper investigation on the status of those names is needed and for the sake of nomenclatural stability, we consider Elaphopus as a nomen dubium .</p><p>Elaphopus Menge, 1879, nomen dubium .</p><p>Elaphidion Menge 1869: 224. Type-species: Elaphidion flagelliferum Menge, 1879, by monotypy. Junior homonym of Elaphidion Audinet-Serville, 1834 ( Coleoptera). Junior subjective synonym of Saloca Simon Simon, 1926 by Roewer (1942: 701); synonymy rejected by Prószyński &amp; Staręga (1971: 150).</p><p>Elaphopus Menge 1879: 1 (in index). Original correct spelling as per first reviser, Prószyński &amp; Staręga (1971: 150). Replacement name for Elaphidion Menge, 1869. Type-species: automatically Elaphidion flagelliferum Menge, 1879.</p><p>Elaphopus — Prószyński &amp; Staręga 1971: 150 (taxonomy).</p><p>Elaphipus: Menge 1879: 8 (in index). Original incorrect spelling.</p><p>Eleaphipus: Petrunkevitch, 1928: 122. Subsequent incorrect spelling.</p><p>Saloca Simon Simon, 1926</p><p>Saloca Simon Simon 1926: 501 . Type-species: Walckenaeria Blackwall diceros O. Pickard-Cambridge, 1871, by monotypy.</p><p>1— Erigoninae: Acartauchenius Simon, Alioranus Simon, Anomalaria Dahl (a junior synonym of Agyneta Hull), Araeoncus Simon (including the junior synonym Scleroschema Hull), Arctilaira Chamberlin (a junior synonym of Hilaira Simon), Aulacocyba Simon (a junior synonym of Microctenonyx Dahl), Auletta O. Pickard-Cambridge (a junior synonym of Macrargus Dahl), Baryphyma Simon, Blaniargus Simon (a junior synonym of Micrargus Dahl), Brattia Simon, Caledonia O. Pickard-Cambridge, Caracladus Simon, Ceratinopsis Emerton, Cineta Simon (currently a nomen dubium), Cnephalocotes Simon, Collinsia O. Pickard-Cambridge (including the junior synonym Coryphaeolana Strand, and Parerigone Crosby &amp; Bishop), Dactylopisthes Simon, Diastanillus Simon, Diplocephalus Bertkau (including the junior synonym Plaesiocraerus Simon, Streptosphaenus Simon), Eboria Falconer (a junior synonym of Semljicola Strand), Eordea Simon, Erigone Audouin, Erigonides Strand (a junior synonym of Hylyphantes Simon), Erigonopsis Hewitt (a junior synonym of Erigonops Scharff), Evansia O. Pickard-Cambridge, Frontella Kulczyński, Glyphesis Simon, Grammonota Emerton (including the junior synonym Itys Strand), Graphomoa Chamberlin (a junior synonym of Pocobletus Simon), Hillhousia F.O. Pickard-Cambridge (a junior synonym of Erigone Audouin), Hybocoptus Simon, Hypselistes Simon, Lophomma Menge, Lucrinus O. Pickard-Cambridge, Lygarina Simon, Mecopisthes Simon, Mermessus O. Pickard-Cambridge, Metopobactrus Simon, Minyriolus Simon, Mioxena Simon, Nematogmus Simon, Nothocyba Simon (a junior synonym of Micrargus Dahl), Notioscopus Simon, Panamomops Simon, Peponocranium Simon, Phanetta Keyserling, Plaesianillus Simon, Priperia Simon, Saloca Simon (including the junior synonym Elaphophus), Scotoneta Simon (a junior synonym of Lessertia Smith), Scotynotylus Simon, Silometopus Simon, Sphecozone O. Pickard-Cambridge (including the junior synonym Clitolyna Simon and Hypselistoides Tullgren), Stajus (a junior synonym of Entelecara Simon), Styloctetor Simon (including the junior synonym Anacotyle Simon), Tapinocyba Simon (including the junior synonym Colobocyba Simon), Thapsagus Simon, Thaumatoncus Simon, Thyreosthenius Simon, Tiso Simon, Trachelocamptus Simon (a junior synonym of Erigonoplus Simon), Trichoncus Simon, Troxochrota Kulczyński, Troxochrus Simon, Tutaibo Chamberlin, Typhistes Simon, Typhochrestus Simon, Utopiellum Strand (a junior synonym of Hilaria Simon), Wubana Chamberlin, Zilephus Simon.</p><p>2—Gonatiinae: Abacoproeces Simon, Aigola Chamberlin (a junior synonym of Oreonetides Strand), Anitsia Chamberlin (a junior synonym of Collinsia O. Pickard-Cambridge), Anthrobia Tellkampf, Atypena Simon, Catabrithorax Chamberlin (a junior synonym of Collinsia O. Pickard-Cambridge), Centromerus Dahl, Dicymbium Menge, Dismodicus Simon, Entelecara Simon, Gnathonarium Karsch, Gonatium Menge, Gongylidiellum Simon, Gongylidium Menge, Halorates Hull, Hilaira Simon, Hylyphantes Simon, Hypomma Dahl, Lasiargus Kulczyński, Lessertia Smith, Maro O. Pickard-Cambridge, Montilaira O. Pickard-Cambridge, Oedothorax Bertkau, Ostearius Hull, Piesocalus Simon, Pocadicnemis Simon, Pseudogonatium Strand (a junior synonym of Zornella Jackson), Rhaebothorax Simon (a junior synonym of Mecynargus Kulczyński), Savignia Blackwall (including the junior synonym Delorrhipis Simon), Scotargus Simon, Spirembolus Chamberlin, Tmeticides Strand (currently a nomen dubium), Tmeticus Menge, Tortembolus Crosby (a junior synonym of Spirembolus Chamberlin), Trematocephalus Dahl Dahl, Walckenaeria Blackwall (including the junior synonym Cornicularia Menge, Prosopotheca Simon, Tigellinus Simon, Trachynotus Dahl, and Wideria Simon).</p><p>Linyphiinae — Agyneta Hull (including the junior synonym Eupolis O. Pickard-Cambridge), Allomengea Strand, Anibontes Chamberlin, Asthenargus Simon &amp; Fage, Bathyphantes Menge, Bolyphantes Strand, Cryptocleptes Simon, Diplocentria Hull, Donacochara Simon, Drapetisca Menge, Emenista Simon, Eryciniola Urquhart (currently Nanometa Simon, in the family Tetragnathidae), Florinda O. Pickard-Cambridge, Floronia Simon, Formicina Canestrini (a junior synonym of Cresmatoneta Simon), Haplinis Simon, Labulla Simon, Labullula Strand, Laetesia Simon, Laperousea Dalmas, Lepthyphantes Menge, Leptorhoptrum Kulczyński, Linyphia Latreille, Linyphioides Strand (currently a nomen dubium), Macrargus Dahl, Microcentria Schenkel (a junior synonym of Diplocentria Hull), Microneta Menge, Micronetata Dahl (a junior synonym of Maro O. Pickard-Cambridge), Micryphantes (currently a nomen dubium), Obrima Simon (a junior synonym of Obrimona Strand), Oreonetides Strand, Parasintula Simon (a junior synonym of Centromerus Dahl), Phaulothrix Bertkau (a junior synonym of Leptothrix Menge), Pocobletus Simon, Porrhomma Simon (including the junior synonym Opistoxys Simon), Rhabdoria (a junior synonym of Centromerus Dahl), Semljicola Strand, Sintula Simon, Smermisia Simon, Solenysa Simon, Sthelota Simon, Syedra Simon, Tapinasta Simon (a junior synonym of Centromerus Dahl), Tapinopa Westring, Taranucnus Simon, Tarsiphantes Strand, Troglohyphantes Joseph (including its junior synonym Typhloneta Kulczyński).</p><p>N.B. The genus Typhloneta was described by Kulczyński (1914) to encompass three species ( T. affinis, T. fugax, and T. salax), without designing a type species. Fage (1919) transferred all three species of Typhloneta to Troglohyphantes Joseph, indirectly synonymizing the two genera but without designating a type species for the former genus. As per the ICZN provision in Article 24.2, we here establish Typhloneta fugax Kulczyński, 1914 the type species of the genus Typhloneta and formalize Typhloneta as a junior synonym of Troglohyphantes Joseph.</p><p>3—Lophocareninae: Ceraticelus Simon, Ceratinella Emerton, Comaroma Bertkau, Erigonoplus Simon (including the junior synonym Cotyora Simon), Exechophysis Simon (a junior synonym of Pelecopsis Simon), Floricomus Crosby &amp; Bishop, Lophocarenum Menge (a junior synonym of Pelecopsis Simon), Thyreobaeus Simon, Trichopterna Kulczyński.</p><p>4—Masoninae: Maso Simon, Minicia Thorell, Notiomaso Banks, Satilatlas Keyserling.</p><p>5—Tennesseellinae: Tennesseellum Petrunkevitch.</p></div>	https://treatment.plazi.org/id/7D34D83FFFA07136FF70B3CC4DBDCE64	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.		MagnoliaPress via Plazi	Silva-Moreira, Thiago Da;Kulkarni, Siddharth;Hormiga, Gustavo	Silva-Moreira, Thiago Da, Kulkarni, Siddharth, Hormiga, Gustavo (2025): A total evidence phylogenetic analysis of the spider family Linyphiidae (Araneae, Araneoidea). Zootaxa 5685 (1): 1-78, DOI: 10.11646/zootaxa.5685.1.1, URL: https://doi.org/10.11646/zootaxa.5685.1.1
7D34D83FFFA27136FF70B1F94BFBCCDC.text	7D34D83FFFA27136FF70B1F94BFBCCDC.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Linyphiidae Blackwall 1859	<div><p>Family Linyphiidae</p><p>1—Centromereae: Centromerita Dahl, Centromerus Dahl, Macrargus Dahl, Oreonetides Strand.</p><p>2—Donacocharae: Drepanotylus Holm, Donacochara Simon, Halorates Hull, Hilaira Simon, Hylyphantes Simon, Leptothrix Menge, Leptorhoptrum Kulczyński, Ostearius Hull.</p><p>3—Leptyphantae: Bolyphantes Strand, Drapetisca Menge, Floronia Simon, Lepthyphantes Menge, Poeciloneta Kulczyński, Tapinopa Westring.</p><p>4—Linyphieae: Bathyphantes Menge, Helophora Menge, Labulla Simon, Linyphia Latreille (including the junior synonym Prolinyphia Homann), Kaestneria Wiehle, Mengea F. O. Pickard-Cambridge (a junior synonym of Allomengea Strand), Microlinyphia Gerhardt, Neriene Blackwall, Pityohyphantes Simon, Stemonyphantes Menge, Stylophora Menge (a junior synonym of Diplostyla Emerton).</p><p>5—Micronetae: Agyneta Hull (including the junior synonym Aprolagus Simon and Meioneta Hull), Maro O. Pickard-Cambridge, Microneta Menge, Syedrula Simon, Syedra Simon, Theonina Simon.</p><p>6—Porrhommae: Hilhousia (a junior synonym of Erigone Audouin), Porrhomma Simon.</p></div>	https://treatment.plazi.org/id/7D34D83FFFA27136FF70B1F94BFBCCDC	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.		MagnoliaPress via Plazi	Silva-Moreira, Thiago Da;Kulkarni, Siddharth;Hormiga, Gustavo	Silva-Moreira, Thiago Da, Kulkarni, Siddharth, Hormiga, Gustavo (2025): A total evidence phylogenetic analysis of the spider family Linyphiidae (Araneae, Araneoidea). Zootaxa 5685 (1): 1-78, DOI: 10.11646/zootaxa.5685.1.1, URL: https://doi.org/10.11646/zootaxa.5685.1.1
7D34D83FFFA27137FF70B2394A6DC895.text	7D34D83FFFA27137FF70B2394A6DC895.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Micryphantidae	<div><p>Family Micryphantidae</p><p>7—Masoninae: Maso Simon, Perimones Jackson, Minicia Thorell</p><p>8—Pelecopsinae: Abacoproeces Simon, Ceratinella Emerton, Comaroma Bertkau, Hypselistes Simon, Mecopisthes Simon, Pelecopsis Simon (including the junior synonym Trichopterna Kulczyński).</p><p>9—Walckenaerinae: Araeoncus Simon, Baryphyma Simon, Cineta Simon, Cnephalocotes Simon, Dicymbium Menge, Dismodicus Simon, Enidia Smith, Entelecara Simon, Evansia O. Pickard-Cambridge, Gnathonarium Karsch, Gonatium Menge, Lasiargus Kulczyński, Lessertia Smith, Metopobactrus Simon, Micrargus Dahl, (including the junior synonym Nothocyba Simon), Minyrioloides Schenkel (a junior synonym of Baryphyma Simon), Minyriolus Simon, Mioxena Simon, Moebelia Dahl, Nematogmus Simon, Panamomops Simon, Peponocranium Simon, Pocadicnemis Simon, Saloca Simon, Silometopus Simon, Styloctetor Simon (including the junior synonym Anacotyle Simon), Tapinocyboides Wiehle, Tiso Simon, Trichoncus Simon, Trematocephalus Dahl, Typhochrestus Simon, Walckenaeria Blackwall (including the junior synonym Cornicularia Menge, Prosopotheca Simon, Tigellinus Simon, Trachynella Braendegaard, Wideria Simon).</p><p>9— Erigoninae: Acartauchenius Simon, Asthenargus Simon &amp; Fage, Aulacocyba Simon, Collinsia O. Pickard-Cambridge, Diplocentria Hull, Diplocephalus Bertkau, Erigone Audouin, Erigonella Dahl, Erigonidium Smith, Erigonopterna Miller, Glyphesis Simon, Gongylidiellum Simon, Gongylidium Menge, Hylyphantes Simon, Lophomma Menge, Microcentria Schenkel (a junior synonym of Diplocentria Hull), Monocephalus Smith, Notioscopus Simon, Oedothorax Bertkau, Rhaebothorax Simon (a junior synonym of Mecynargus Kulczyński), Savignia Blackwall, Tapinocyba Simon, Tmeticus Menge, Troxochrus Simon, Thyreosthenius Simon.</p></div>	https://treatment.plazi.org/id/7D34D83FFFA27137FF70B2394A6DC895	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.		MagnoliaPress via Plazi	Silva-Moreira, Thiago Da;Kulkarni, Siddharth;Hormiga, Gustavo	Silva-Moreira, Thiago Da, Kulkarni, Siddharth, Hormiga, Gustavo (2025): A total evidence phylogenetic analysis of the spider family Linyphiidae (Araneae, Araneoidea). Zootaxa 5685 (1): 1-78, DOI: 10.11646/zootaxa.5685.1.1, URL: https://doi.org/10.11646/zootaxa.5685.1.1
7D34D83FFFA37137FF70B7F14CAACC81.text	7D34D83FFFA37137FF70B7F14CAACC81.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Linyphiidae Subfamilial	<div><p>Linyphiidae subfamilial classification sensu Locket &amp; Millidge (1953)</p><p>1—Subfamily Erigoninae: Acartauchenius Simon, Araeoncus Simon, Asthenargus Simon &amp; Fage, Aulacocyba Simon, Baryphyma Simon, Caledonia O. Pickard-Cambridge, Ceratinella Emerton, Cnephalocotes Simon, Collinsia O. Pickard-Cambridge, Dicymbium Menge, Dismodicus Simon, Diplocentria Hull, Diplocephalus Bertkau, Eboria Falconer (a junior synonym of Semljicola Strand), Entelecara Simon, Erigone Audouin, Erigonidium Smith, Erigonella Dahl, Evansia O. Pickard-Cambridge, Glyphesis Simon, Gnathonarium Karsch, Gonatium Menge, Gongylidiellum Simon, Gongylidium Menge, Hypomma Dahl (a junior synonym of Enidia Smith), Hypselistes Simon, Hybocoptus Simon, Jacksonella Millidge, Lessertia Smith, Lophomma Menge, Maso Simon, Mecopisthes Simon, Metopobactrus Simon, Micrargus Dahl, Minyriolus Simon, Minyrioloides Schenkel (a junior synonym of Baryphyma Simon), Mioxena Simon, Moebelia Dahl, Monocephalus Smith, Notioscopus Simon, Oedothorax Bertkau, Panamomops Simon, Peponocranium Simon, Pelecopsis Simon (including the junior synonym Lophocarenum Menge, Trichoncus Simon, and Trichopterna Kulczyński), Perimones Jackson, Pocadicnemis Simon, Praestigia Millidge, Rhaebothorax Simon (a junior synonym of Mecynargus Kulczyński), Saloca Simon, Savignia Blackwall, Scotargus Simon, Silometopus Simon, Styloctetor Simon (including the junior synonym Anacotyle Simon), Tapinocyba Simon, Tapinocyboides Wiehle, Tiso Simon, Thyreosthenius Simon, Tmeticus Menge, Trematocephalus Dahl, Troxochrus Simon, Typhochrestus Simon, Walckenaeria Blackwall (including the junior synonym Cornicularia Menge, Prosopotheca Simon, Tigellinus Simon, Trachynella Braendegaard, and Wideria Simon).</p><p>2—Subfamily Linyphiinae: Agyneta Hull, Bathyphantes Menge, Bolyphantes Strand, Centromerus Dahl, Centromerita Dahl, Diplostyla Emerton, Donacochara Simon, Drapetisca Menge, Drepanotylus Holm, Floronia Simon, Halorates Hull, Helophora Menge, Hilaira Simon, Hillhousia F.O. Pickard-Cambridge (a junior synonym of Erigone Audouin), Kaestneria Wiehle, Labulla Simon, Lepthyphantes Menge, Leptorhoptrum Kulczyński, Linyphia Latreille, Macrargus Dahl, Maro O. Pickard-Cambridge, Meioneta Hull (a junior synonym of Agyneta Hull), Mengea F. O. Pickard-Cambridge (a junior synonym of Allomengea Strand), Microneta Menge, Oreonetides Strand, Ostearius Hull, Phaulothrix Bertkau, Poeciloneta Kulczyński, Porrhomma Simon, Stemonyphantes Menge, Syedrula Simon, Syedra Simon, Tapinopa Westring, Taranucnus Simon.</p></div>	https://treatment.plazi.org/id/7D34D83FFFA37137FF70B7F14CAACC81	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.		MagnoliaPress via Plazi	Silva-Moreira, Thiago Da;Kulkarni, Siddharth;Hormiga, Gustavo	Silva-Moreira, Thiago Da, Kulkarni, Siddharth, Hormiga, Gustavo (2025): A total evidence phylogenetic analysis of the spider family Linyphiidae (Araneae, Araneoidea). Zootaxa 5685 (1): 1-78, DOI: 10.11646/zootaxa.5685.1.1, URL: https://doi.org/10.11646/zootaxa.5685.1.1
7D34D83FFFA37130FF70B2054C1BCB20.text	7D34D83FFFA37130FF70B2054C1BCB20.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Linyphiidae Subfamilial	<div><p>Linyphiidae Subfamilial Division sensu Merrett (1963, 1965)</p><p>1—Subfamily Linyphiinae</p><p>1a Syedruleae: Agyneta Hull (including the junior synonym Meioneta Hull), Bolyphantes, Strand, Centromerita Dahl, Centromerus Dahl, Drapetisca Menge, Floronia Simon, Helophora Menge, Kaestneria Wiehle, Lepthyphantes Menge, Macrargus Dahl, Maro O. Pickard-Cambridge, Microneta Menge, Oreonetides Strand, Poeciloneta Kulczyński, Syedra Simon, Syedrula Simon, Tapinopa Westring.</p><p>1b Linyphieae: Allomengea Strand, Bathyphantes Menge, Diplostyla Emerton, Labulla Simon, Linyphia Latreille, Porrhomma Simon, Stemonyphantes Menge, Taranucnus Simon.</p></div>	https://treatment.plazi.org/id/7D34D83FFFA37130FF70B2054C1BCB20	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.		MagnoliaPress via Plazi	Silva-Moreira, Thiago Da;Kulkarni, Siddharth;Hormiga, Gustavo	Silva-Moreira, Thiago Da, Kulkarni, Siddharth, Hormiga, Gustavo (2025): A total evidence phylogenetic analysis of the spider family Linyphiidae (Araneae, Araneoidea). Zootaxa 5685 (1): 1-78, DOI: 10.11646/zootaxa.5685.1.1, URL: https://doi.org/10.11646/zootaxa.5685.1.1
7D34D83FFFA47131FF70B1114CFDCDE1.text	7D34D83FFFA47131FF70B1114CFDCDE1.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Linyphiidae Subfamilial	<div><p>Linyphiidae Subfamilial Division sensu Millidge (1984, 1993)</p><p>1— Dubiaraneinae: Dubiaranea Mello-Leitão, Notiohyphantes Millidge.</p><p>2— Erigoninae: Abacoproeces Simon, Acartauchenius Simon, Alioranus Simon, Araeoncus Simon, Aulacocyba Simon, Baryphyma Simon, Caledonia O. Pickard-Cambridge, Caracladus Simon, Ceraticelus Simon, Ceratinella Emerton, Ceratinopsis Emerton, Cnephalocotes Simon, Collinsia O. Pickard-Cambridge, Dicymbium Menge, Diplocephalus Bertkau, Dismodicus Simon, Donacochara Simon, Drepanotylus Holm, Eboria Falconer (a junior synonym of Semljicola Strand), Erigonella Dahl, Emenista Simon, Entelecara Simon, Eridantes Crosby &amp; Bishop, Erigone Audouin, Erigonella Dahl, Erigonidium Smith, Erigonoplus Simon, Evansia O. Pickard-Cambridge, Exechophysis Simon, Glyphesis Simon, Gnathonargus Bishop &amp; Crosby, Gnathonarium Karsch, Gonatium Menge, Gongylidiellum Simon, Gongylidium Menge, Grammonota Emerton, Halorates Hull, Hilaira Simon, Hybocoptus Simon, Hypomma Dahl, Hypselistes Simon, Islandiana Braendegaard, Lessertia Smith, Lophomma Menge, Maso Simon, Mecopisthes Simon, Metopobactrus Simon, Microcyba Holm, Milleriana Denis (a junior synonym of Collinsia O. Pickard-Cambridge), Minicia Thorell, Minyrioloides Schenkel (a junior synonym of Baryphyma Simon), Minyriolus Simon, Moebelia Dahl, Monocephalus Smith, Notioscopus Simon, Oedothorax Bertkau, Origanates Crosby &amp; Bishop, Panamomops Simon, Pelecopsis Simon, Peponocranium Simon, Perimones Jackson, Phanetta Keyserling, Phaulothrix Bertkau, Pocadicnemis Simon, Praestigia Millidge, Rhaebothorax Simon (a junior synonym of Mecynargus Kulczyński), Saloca Simon, Savignia Blackwall, Sciastes Bishop &amp; Crosby, Scotinotylus Simon, Scylaceus Bishop &amp; Crosby, Scytiella Simon, Silometopus Simon, Sisicottus Bishop &amp; Crosby, Souessoula Crosby &amp; Bishop, Spirembolus Chamberlin, Stajus Simon (a junior synonym of Entelecara Simon), Strongyliceps Fage, Tachygyna Chamberlin &amp; Ivie, Tapinocyba Simon, Tapinocyboides Wiehle, Thyreosthenius Simon, Tiso Simon, Tmeticus Menge, Trematocephalus Dahl, Trichoncoides Denis, Trichoncus Simon, Trichopterna Kulczyński, Troxochrus Simon, Tunagyna Miller, Typhochrestus Simon, Walckenaeria Blackwall, Zornella Jackson.</p><p>3— Lepthyphantinae: Bolyphantes Strand, Centromerita Dahl, Centromerus Dahl, Centrophantes Miller &amp; Polenec, Floronia Simon, Gnathantes Hull, Labullula Strand, Lepthyphantes Menge, Locketidium Jocqué, Macrargus Dahl, Metaleptyphantes Locket, Oreonetides Strand, Poeciloneta Kulczyński, Syedra Simon, Tapinopa Westring, Troglohyphantes Joseph.</p><p>4— Linyphiinae: Bathyphantes Menge, Cresmatoneta Simon, Cryptolinyphia Millidge, Diplostyla Emerton, Frontinellina van Helsdingen, Kaestneria Wiehle, Laetesia Simon, Laperousea Dalmas, Lessertinella Denis, Linyphia Latreille, Maro O. Pickard-Cambridge, Microlinyphia Gerhardt, Neriene Blackwall, Oilinyphia Ono &amp; Saito, Pityohyphantes Simon, Porrhomma Simon, Sisicus Bishop &amp; Crosby.</p><p>5— Micronetinae: Agyneta Hull, Microneta Menge, Syedrula Simon, Tennesseellum Petrunkevitch, Theonina Simon.</p><p>6— Mynogleninae: Afroneta Holm, Cassafroneta Blest, Haplinis Simon, Hyperafroneta Blest, Megafroneta Blest, Metafroneta Blest, Metamynoglenes Blest, Novafroneta Blest, Parafroneta Blest, Poecilafroneta Blest, Promynoglenes Blest, Protoerigone Blest, Pseudafroneta Blest, Trachyneta Holm.</p><p>“Miscellaneous genera” (previously Drapetiscinae)— Allomengea Strand, Aphileta Hull, Arcuphantes Chamberlin &amp; Ivie, Asthenargus Simon &amp; Fage, Doenitzius Oi, Drapetisca Menge, Fageiella Kratochvíl, Helophora Menge, Jacksonella Millidge, Laminacauda Millidge, Neomaso Forster, Saaristoa Millidge, Sintula Simon, Stemonyphantes Menge, Typhlonyphia Kratochvíl, Wubana Chamberlin.</p><p>Supplementary Materials. The following supporting information can be downloaded at the DOI landing page of this paper.</p><p>SUPPLEMENTARY FILE 1. Phenotypic Matrix.</p><p>SUPPLEMENTARY FILE 2. Morphological character list (Originally published in Silva Moreira &amp; Hormiga, 2022).</p><p>TABLE S1. Geographic information and voucher number of the species used for the first time in this study.</p><p>SUPPLEMENTARY FIGURE 1. Tree hypotheses summaries.A, Consensus of all hypothesis 1 trees. B, Consensus of all hypothesis 2 trees. B, Consensus of all hypothesis 3 trees. Arrows point to the Outgroups. Clades are colored as follows: Light pink— Pimoidae; Brown— Stemonyphantinae; Red— Labulla thoracica; Yellow— Pocobletinae; Green— Linyphiinae (new circumscription); Blue—ConoSur Clade; Orange—Clade A; Dark Pink—Clade B. The genera Labulinyphia and Turinyphia are considered incertae sedis and not considered when summarizing the topologies.</p><p>SUPPLEMENTARY FIGURE 2. Hypothesis 1 ( Pocobletinae + ( Linyphiinae + Clade B + Clade A + ConoSur clade))trees summary. Letters indicate the analysis and the recovered topology.Asteriscs (*) indicate unique topology. Arrow points to outgroups. Clades are colored as follows: Light pink— Pimoidae; Brown— Stemonyphantinae; Red— Labulla thoracica; Yellow— Pocobletinae; Green— Linyphiinae (new circumscription); Light Blue— ConoSur Clade; Orange—Clade A; Dark Pink—Clade B. The genera Labulinyphia and Turyniphia are considered incertae sedis and not considered when summarizing the topologies.</p><p>SUPPLEMENTARY FIGURE 3. Hypothesis 2 (Clade B + ( Linyphiinae + Pocobletinae + Clade A + ConoSur clade)) trees summary.Letters indicate the analysis and the recovered topology.Asteriscs (*)indicate unique topology. Arrow points to outgroups. Clades are colored as follows: Light pink— Pimoidae; Brown— Stemonyphantinae; Dark Blue— Pecado impudicus; Red— Labulla thoracica; Yellow— Pocobletinae; Green— Linyphiinae (new circumscription); Light Blue—ConoSur Clade; Orange—Clade A; Dark Pink—Clade B. The genera Labulinyphia and Turinyphia are considered incertae sedis and not considered when summarizing the topologies.</p><p>SUPPLEMENTARY FIGURE 4. Hypothesis 3 ( Linyphiinae + ( Pocobletinae + Clade B + Clade A + ConoSur clade)) trees summary.Letters indicate the analysis and the recovered topology.Asteriscs (*)indicate unique topology. Arrow points to outgroups. Clades are colored as follows: Light pink— Pimoidae; Brown— Stemonyphantinae; Red— Labulla thoracica; Yellow— Pocobletinae; Green— Linyphiinae (new circumscription); Light Blue— ConoSur Clade; Orange—Clade A; Dark Pink—Clade B. The genera Labulinyphia and Turinyphia are considered incertae sedis and not considered when summarizing the topologies.</p></div>	https://treatment.plazi.org/id/7D34D83FFFA47131FF70B1114CFDCDE1	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.		MagnoliaPress via Plazi	Silva-Moreira, Thiago Da;Kulkarni, Siddharth;Hormiga, Gustavo	Silva-Moreira, Thiago Da, Kulkarni, Siddharth, Hormiga, Gustavo (2025): A total evidence phylogenetic analysis of the spider family Linyphiidae (Araneae, Araneoidea). Zootaxa 5685 (1): 1-78, DOI: 10.11646/zootaxa.5685.1.1, URL: https://doi.org/10.11646/zootaxa.5685.1.1
