Just as some neotrichous epimere IV setae form in the protonymph, we envision setae *x* and *y* as precocious aggenital setae or their neotrichous derivatives.

Regardless of the source of the neotrichous setae, we cannot confidently identify any particular number as aggenital seta, and the specific attribution of six pairs to *Eulohmannia* by Balogh and Balogh (1992; p. 35) or 14 pairs for *E. bifurcata* by Fujikawa (2014) seems arbitrary.

**R14** — *Paraproctal setation*. The ontogeny of setation on paraproctal valves is rather simple and consistent in our material of *E. ribagai* (regardless of provenance) and *P. metzi*. Segment *PS* has four setae in the larva; following Grandjean's (1949b) concept, the most anterior is an inguinal seta, *psi* (*p*₄), which is lost when the segment joins the gastronotum in the protonymph. Such inguinal setae are common in early derivative oribatid mites and endeostigmatids (Grandjean 1942b, 1949b; Norton and Behan-Pelletier 2007 [their note 8]; Norton and Fuangarworn 2015 [their remark 11]). Segment *AD* appears in the protonymph with its full complement of four pairs of setae, and in *E. ribagai* (but not *P. metzi*) segment *AN* appears in the deutonymph, also with four pairs. The setal complements of *AD* and *AN* do not change in subsequent instars.

But according to Grandjean's (op. cit. p. 206) analysis, the ontogeny of both segments *PS* and *AD* is highly unusual in *E. ribagai*. Using a shorthand formula, he presented the setal ontogeny as 3–3444–444, which can be interpreted as follows. Segment *PS* (the first number) has three pairs of setae in the larva, but he was uncertain of the number in subsequent instars, where segment *PS* moves to the gastronotum; segment *AD* has three pairs when it appears in the protonymph, but four in subsequent instars; segment *AN* has four pairs when it appears in the deutonymph and retains this number. Except in taxa with neotrichy or paraproctal atrichosy, in which segments are glabrous when they first appear (Grandjean op. cit. p. 207), it is highly unusual for a segment to add setae after it forms. He cited only two examples of such delay in segment *AD* setae: *E. ribagai* and *Torpacarus omittens* (Lohmanniidae). As discussed above (justification of *Paedolohmannia* n. gen.) we feel he was wrong about *T. omittens*: a more parsimonious explanation is that the seta added in the deutonymph is an anal seta.

We also believe Grandjean proposed an incorrect ontogenetic formula for *E. ribagai* because he overlooked seta *ps*₁ in the larva of *E. ribagai*, as well as seta *ad*₁ in the protonymph. This might seem unlikely, considering that he was a renowned observer, and that in our material these are the largest setae on their respective segments (Figs 3C, D, 4A). But both these enlarged setae are highly prone to breakage during handling and clearing treatments; they are missing from many of our prepared specimens while the three smaller, more anterior setae of the respective segment are always intact. The large alveolus of a broken *ps*₁ or *ad*₁ can be found if searched for, but it is surprisingly inconspicuous: being close to the lateral edge and near the posterior end of the segment, it becomes almost vertically oriented in usual dorsoventral orientations and is covered by the edge of the gastronotum in lateral orientation. Both *ps*₁ and *ad*₁ (or the alveolus if broken) always are present in our abundant Nearctic material.

Grandjean did not mention the number of larvae or protonymphs he observed, but we studied one protonymph from Austria (Fig. 11C) and it is consistent with our Nearctic material: on one side the enlarged *ad*₁ is present, but on the other only the large alveolus remains. Also consistent are three protonymphs from Germany (Saxony and Berlin states) in the Senckenberg Museum für Naturkunde Görlitz studied by Dr. Ricarda Lehmitz (pers. comm. 2021): one had the large pair *ad*₁ present, the other two showed only their large alveoli. She also studied two larvae from the Czech Republic (Krkonoše Mountains, Šumava Mountains) and while setae *ps*₁ were broken, their large alveoli were present. Similarly, a larva from Norway (near Kumra, Nordland province) studied by Dr. Anna Seniczak (pers. comm. 2022) had the alveoli of *ps*₁ though the setae were lost.

**R15** — *Vestigial seta d on genua I-III.* As part of a benchmark series of papers on the comparative chaetotaxy of oribatid mite legs, Grandjean (1942a) identified eight different ‘behaviors’ of genu seta *d* during ontogeny. Seven of these relate to patterns in which the seta is somehow diminished in size or suppressed in one or more instars. A pattern he called ‘larval
suppression' was considered unique to *E. ribagai* at that time, but it is shared by *Eulohmannia* sp. A and *Paedolohmannia metzi*. In each instance seta *d* is represented in the larva by a minute setal vestige inserted in a small pore-like alveolus (Figs 12B, 15G, 26A-C), but a normal seta forms in subsequent instars. A similar development occurs in *Perlohmannia* (Grandjean 1958A; Suzuki 1977) and *Nehypochthonius* (new observation; overlooked in original description, cf. Norton and Metz 1980).

This vestige easily might be confused with the so-called 'genual pore' (Grandjean 1940) that appears on these same segments in some Nothrina and Mixonomata (but not Eulohmanniidae). By contrast, this pore lacks an internal vestige, exists in all instars, and is always clearly independent of seta *d* (e.g., Grandjean 1954b; Norton and Sidorchuk 2014).

**R16** — *Leg IV setation in protonymph.* Leg IV of acariform mites forms in the protonymph with a diminished chaetome. Grandjean (1946b) noted many variations, but among oribatid mites the typical protonymphal setation includes seven setae on tarsus I, with all other segments being glabrous. The protonymph of *E. ribagai* and *P. metzi* have the usual tarsal setation, but tibia IV consistently forms seta *v''*. It is common, and perhaps plesiomorphic for *v''* to be retained on protonymphal tibia IV in Enarthronota and it forms in large species of Gehypochthoniidae (Parhyposomata) (Norton and Fuangarworn 2015), but these are the only taxa that share this trait, to our knowledge.

**R17** — *Setal pair (m) on tarsus I.* The 'poil monotrope', *m''*, is a eustasic larval seta of tarsus I located slightly distal to *pv''* in some early-derivative groups of oribatid mites (Grandjean 1962). It is common in Parhyposomata and Enarthronota (see also Norton and Fuangarworn 2015, their Remark #17) but we are not aware of any mixonomatan other than Eulohmanniidae that has this plesiomorphic seta, which was first reported in *E. ribagai* by Grandjean (1941b).

Seta *m'*, the pseudosymmetrical complement of *m''*, is less common and always developmental—deutonymphal in Eulohmanniidae (Figs 15I, 24D). We refer to pair (*m*) as monotropic setae even though the application is grammatically incongruous: derived from the Greek *mono* (one) and *tropos* (turning, direction), the name made sense originally because of the asymmetry of *m''*. Seta *m'* was not confidently identified in any oribatid mite until Grandjean (1962) became convinced that a developmental seta could appear distal to fundamental seta *pv'*, instead of proximal to it like a typical proximal accessory setae. The presence of *m'* in Eulohmanniidae is noted here for the first time. The pair (*m*) has a strong anterior (') disjunction, which also is true of *Gehypochthonius xarifae* Strenzke, 1963 (Parhyposomata); although publication delay made the chronology confusing, Grandjean (1962) considered the seta labeled *pv'* by Strenzke (1963, his Fig. 11) to be instead *m'*. This disjunction conforms to the usual anterior offset of the primiventral pair (*pv*) on tarsus I of oribatid mites (Grandjean 1958b, 1960), although in Eulohmanniidae (*pv*) have little or no disjunction. By contrast, in most of those Enarthronota that develop both *m'* and *m''* the pair has a posterior disjunction (Norton and Fuangarworn 2015).

The fate of pair (*m*) during development in Eulohmanniidae—*m''* remaining throughout ontogeny but *m'* becoming eupathidial—appears to be unique. The opposite occurs in several Enarthronota (Norton and Fuangarworn 2015), where the more distal *m''* becomes eupathidial and *m'* remains normal. In Grandjean's (1962) interpretation, both *m''* and *m'* become eupathidial in *G. xarifae*, in the tritonymph and adult, respectively.

**R18** — *Iteral setae.* Among oribatid mites the ontogeny of iteral setae (*it*), which are always developmental, varies greatly according to taxon, leg, and even between members of the pseudosymmetrical pair. As summarized by Grandjean (1964a) their ontogeny in *E. ribagai* is unique, with each leg being different: the development shown in Table 4 is summarized by his formula (n1—[n3, n2]—n3—o). In the context of his conceptual model of regressive evolution in these setae, the early appearance of (*it*) on tarsus I places it among the more plesiomorphic ontogenies known. The transformation of (*it*) on tarsus I to eupathidia is shared by a wide range of early-derivative oribatid mites, including members of Palaeosomata, Enarthronota, Parhyposomata, and some other families of Mixonomata.
Collohmanniidae (Norton and Sidorchuk 2014) setae of the *c*-row appear to follow Grandjean's rules for accessory setae (see R19) precisely, though they are not present in the nothrine taxa he used to initially formulate the rules (Grandjean 1958b). In these two families a pair of *c*-setae are added in each of the last three instars, with the respective notations (*c*<sub>n2</sub>), (*c*<sub>n3</sub>) and (*c*<sub>A</sub>).

In Eulohmanniidae only one series of proximal accessory setae forms. We identify them as *c*-setae because in their normal (non-eupathidial) form they align with (a) and because no accessory setae appear proximal to either (*pl*) or (*pv*)—i.e., they are not *l*- or *v*-setae. If either a *c*- or *a*-seta transforms to a eupathidium, alignment can be affected since setae tend to be slightly displaced distally and/or ventrally after transformation. We know of no other oribatid mites in which *c*-setae appear in the absence of other accessory setae. But the *c*-setae of Eulohmanniidae are unique, to our knowledge, in not precisely following Grandjean's rules (Table 5). Both *Paedolohmannia metzi* and *Eulohmannia* sp. A (Aborigen) have the same ontogeny in the *c*-row: a pair (*c*<sub>n2</sub>) forms in the deutonymph, as expected, and these have a null or slight posterior (") disjunction (Fig. 24D), but then a single posterior seta is added in the tritonymph and a single anterior seta added in the adult. If the latter two setae are eustasic, they would be *c*<sub>n3</sub>" and *c*<sub>A</sub>", respectively, which are the notations we apply herein. By contrast, *E. ribagai* forms a single *c*-seta in each of the last three instars, differing from the other two species in lacking seta *c*<sub>n2</sub>". We have seen no juveniles of *Eulohmannia* sp. B (Kashmir), but the adult has a single *c*-seta on each face, which in the other three species occurs only in nymphs; we interpret this as another manifestation of its paedomorphic tarsus.

While Grandjean's second rule of accessory setae applies to *c*-setae in Eulohmanniidae (they are present only on tarsus I), his third does not: setae remain in the presumed deutonymphal and tritonymphal verticils even though the adult verticil is partially lost. Also, the priority of setae seems to vary among the verticils, with the posterior (") seta being lost in the deutonymphal and adult verticils in *E. ribagai*, as expected from general priorities (see R19), whereas the tritonymphal verticil has lost the anterior seta ('). We have no explanation of these observed differences if the setae are eustasic.

An alternative hypothesis is that some *c*-setae instead have become amphistasic. In such a case, *c*-seta homologies could not be established based on the instar of appearance. For example, if the tritonymphal seta denoted *c*<sub>n3</sub>" in *E. ribagai* (Fig. 15J) were truly from the ancestral tritonymphal verticil (eustasy hypothesis), one would not expect it to have a more distal position than *c*<sub>n2</sub>' from the deutonymphal verticil. However, if the purported *c*<sub>n3</sub>" is actually a delayed *c*<sub>n2</sub>" (amphistasy hypothesis), then the positions of the two *c*-setae in the tritonymph would be exactly those of pair (*c*<sub>n2</sub>) in the deutonymph of *P. metzi* and *Eulohmannia* sp. A (cf. Figs 15J, 24D). At the moment, neither hypothesis seems stronger than the other.

**R21** — *Tarsus I eupathidia*. Eulohmanniidae have a moderate number—9, 10 or 11—of these probable gustatory chemosensilla (Alberti 1998) on tarsus I. This is far fewer than Perlohmanniidae (Grandjean 1958a) or Collohmanniidae (Grandjean 1966; Norton and Sidorchuk 2014), both of which have eupathidia among the abundant proximal accessory setae. Each of the three species for which we have developmental data has a slightly different adult complement and ontogeny of eupathidia (Table 5). We have no juveniles of the paedomorphic *Eulohmannia* sp. B (Kashmir) but based on a single studied adult there are nine, including (*p*), (*u*), (*it*), *s*, *m*' and one abaxial *c*-seta (not yet identified). Eulohmanniidae are almost unique among oribatid mites in having pair (*u*) eupathidial—according to Grandjean (1958a) this is shared only by some (unnamed) Damaeidae ('Belbidae').

Most ontogenetic transformations are constant: setae (*p*) are eupathidial from the larva and each species transforms *s* and *u'* in the protonymph, (*it*) in the deutonymph, and *m'* in the tritonymph. If seta *c*<sub>n2</sub>" is present (*Eulohmannia* sp. A, *P. metzi*), it transforms in the tritonymph. We identified only one variation: the transformation of *u''* is either protonymphal, like *u'*, or delayed to the deutonymph (*P. metzi*).

Transformation of the antelateral setae is most diverse and lacks any recognizable pattern. Both *Eulohmannia* sp. A and *E. ribagai* transform *a'* and *a''*, but with reversed ontogenetic priority; *P. metzi* transforms *a''* surprisingly early (Pn) but *a'* remains normal throughout
ontogeny; in *Eulohmannia* sp. B neither a-seta transforms. We observed few intraspecific variations in eupathidial transformation, and all related to antelateral setae. In *Eulohmannia* sp. A, *a"* remained normal on one of four deutonymphal legs examined. In *E. ribagai* seta *a'* transforms in the deutonymph, except on one of 10 tarsi examined from the New York population it remained normal. The transformation of *a"* in *E. ribagai* is more variable, perhaps differing among populations (but there are few observations). In three tritonymphs from New York and one from Germany, only *a'* was eupathidial, suggesting that *a"* would have transformed in the adult. In another tritonymph from New York, two from Austria (near the type locality), and two from Canada (Cape Breton and Reindeer Station) *a"* had transformed, so both (*a*) were eupathidial. While variable among specimens, in each instance the state of *a"* was symmetrical in a given mite.

Eulohmanniidae may be unique regarding eupathidial transformation in the monotropic pair (*m*). In some in Enarthronota (see Norton and Fuangarworn 2015) *m"* becomes eupathidial while *m'* remains normal, and in *Gehypochthonius xarifae* (Parhyposomata) both *m'* and *m"* become eupathidial, with *m"* transforming first (Grandjean 1962). But we are unaware of any species outside Eulohmanniidae in which *m'* transforms to a eupathidium while *m"* remains normal.

**R22** — *Complement of solenidia*. Eulohmanniidae have a relatively rich complement of solenidia (15 per side), with a development pattern widely shared among early-derivative oribatid mites (Grandjean 1964c). Like most Parhyposomata, Mixonomata and Enarthronota, they have a single (larval) solenidion on tibia I, having lost the developmental *φ*₂ that typifies most taxa in the more highly derived Nothrina and Brachypylina. An opposite pattern is seen on genu I, where a developmental solenidion is added to the two larval solenidia. This is known elsewhere only in Palaeacaridae (Palaeosomata; several other palaeosomatans with three solenidia have unknown development) and Nanohystricidae (Enarthronota); in these instances, the third solenidion is tritonymphal or (in *Nanohystrix*) possibly deutonymphal (Grandjean 1964c; Norton and Fuangarworn 2015), whereas that of Eulohmanniidae appears in the protonymph (Grandjean 1942a). If regression has shaped the solenidial complements of oribatid mites, then that of Eulohmanniidae is the least regressive known for genu I.

**R23** — *Coupling of solenidia*. In the large majority of oribatid mites, seta *d* is coupled to a solenidion on at least some genua and tibiae or shows evidence that this was true in their ancestors (Grandjean 1942a). In Eulohmanniidae seta *d* is distant from any solenidion on these segments and is one of the few families outside Palaeosomata and Parhyposomata in which this plesiomorphic separation persists. Among mixonomatans Perlohmanniidae is the only other family lacking such coupling entirely.

However, in all instars seta *l'* of genu I is coupled to solenidion *σ'*. This same association is found in the early-derivative mixonomatan family Nehypochthoniidae, in which there are no *d*-solenidion couplings on leg I, though they exist on other legs (Norton and Metz 1980). An *l'*-*σ* coupling also occurs on genu I of Ptyctima, both Phthiracaroidea and Euphthiracaroidea (e.g., Grandjean *op. cit.*; Hammen 1963; Lions and Norton 1998), in which *d*-solenidion couplings do exist. We are not aware of *l'*-solenidion couplings on genu I outside Mixonomata, but on genu II of *Mucronothrus* (Nothrina) a weak coupling develops during ontogeny (Norton *et al.* 1996).

**R24** — *Pretarsal ambulacrum*. Grandjean (1969) considered the ancestral adult oribatid mite ambulacrum to be tridactylous, with strong empodial and paired lateral claws. He viewed the combination of strong lateral claws and a regressed central (empodial) claw—the ‘*Acaronychus*'-type' of tridactyly (herein type-A)—as an 'exceptional secondary character', shared among members of three of his major groups: Palaeosomata (Archeonothroidea), Parhyposomata (*Parhypochthonius*) and Mixonomata (*Eulohmannia*). To these we can now add Elliptochthoniidae (Norton 1975), Nehypochthoniidae (Norton and Metz 1980) and some Gehypochthoniidae (Strenzke 1963, Martinez and Bernava Laborde 2000). Grandjean considered a bidactylous ambulacrum, with the absence of the empodial claw (herein type-B), as a different 'exceptional character', found in some primitive Enarthronota—Atopochthoniidae
and Phyllochthoniidae (and subsequently in Arborichthoniidae; Norton 1982)—plus some unnamed *Gehypochthonius* species (probably *G. rhadamanthus* Jacot, which he had studied previously; Grandjean 1943). At least in Gehypochthoniidae, it seems clear that the type-B ambulacrum represents the end of a regressive trend, since the reduced empodium is either present or absent, according to species. With one exception, all Eulohmanniidae discussed herein have a type-A ambulacrum: the purported absence of an empodial claw in *E. bifurcata* is an error, as discussed above. Only in *Eulohmannia* sp. B (Kashmir) is the empodial claw large, and this seems a clear example of neoteny (see above).

In our view, type-A tridactyly was ancestral in Novoribatida, since it is present in all known genera and most known species of Parhyposomata; therefore, we consider the type-A ambulacrum in the mixonomatan families Nehypochthoniidae and Eulohmanniidae as plesiomorphic, rather than secondary. While there are no known extant Enarthronota with a type-A ambulacrum, it existed in the extinct family Protochthoniidae (Norton et al. 1988), so a similar regressive loss may have resulted in the type-B ambulacrum of the enarthronote families noted above. Since the type-A ambulacrum also characterizes Archeonothroidea, which we consider the most basal group of Palaeosomata (Norton and Fuangarworn 2015), it may be ancestral in Sarcoptiformes.

Among mixonomatans, Perlohmanniidae and Epilohmanniidae adults are monodactylous, suggesting paedomorphosis at some point in their evolution, like that of *Eulohmannia* sp. B. Collohmanniidae are homotridactylous (Norton and Sidorchuk 2014) and Ptyctima are rather plastic in ambulacral structure (e.g., Lions 1964; Lions and Norton 1998), but in no instance is the empodium reduced. Unlike Eulohmanniidae, these other mixonomatans have ambulacra that collectively mirror those of Nothrina and Brachypylina. If the type-A ambulacrum is ancestral in oribatid mites, these more widespread adult forms could have evolved from it by the fixation of a large, paedomorphic empodial claw, while lateral claws either remained large or became susceptible to regression.

**R25** — *Rutellar comb* (= brush, ciliary comb). In Nothrina and Brachypylina, the dorsal face of the rutellum usually bears a comb comprising one (usually) or two rows of thin 'cilia' that presumably act to clean small particles from the lateral (abaxial) face of the chelicera (Grandjean 1957, Hammen 1968). The comb is secondarily absent—a loss—from many Brachypylina with specialized mouthparts, and probably so in most Astigmata (Norton 1998). Palaeosomata, Enarthronota, and Parhyposomata also lack the comb, but this is an ancestral state.

In mixonomatans—none of which have significantly specialized mouthparts—the comb can be well developed, as in Eulohmanniidae, or have structures that may be homologs of the comb. Such a distribution suggests that the comb evolved within Mixonomata. Grandjean (1958a) considered irregular dorsal ridges on the rutellum of *Perlohmannia* to represent a reduction (i.e., a loss) of the comb's cilia, but this seems unlikely. If the comb functions by removing debris, what advantage would be gained by their loss from an unspecialized rutellum? The opposite trend seems more likely: a rutellum dorsally roughened by ridges (seen also in Nehypochthoniidae and Epilohmanniidae) represents the primitive form of a cleaning surface, with comb-like projections from the ridges having evolved subsequently.

In Collohmanniidae the rutellum has dorsal ridges that lack cilia, but there are 1-3 (usually two) small spines (Norton and Sidorchuk 2014). Grandjean (1966) thought these spines represented a special form of the comb; if so, it seems an intermediate form, rather than representing the culmination of a trend. Eulohmanniidae have a comb (Fig. 16K; seen by transparency in Figs 10A, C, 21C) as fully developed as those in Nothrina and Brachypylina, so it seems the most highly-derived form among mixonomatans.

**R26** — *Palp segmentation*. The plesiomorphic state in oribatid mites is to retain full independence of the five ancestral articulating segments: trochanter (*Tr*), femur (*Fe*), genu (*Ge*), tibia (*Ti*) and (*Ta*) tarsus. All Palaeosomata have this state, but fusions between segments are known in some families of all other major groups. Among mixonomatans, Nehypochthoniidae, Perlohmanniidae and Collohmanniidae retain the plesiomorphic state. Eulohmanniidae have
a complete Fe-Ge fusion (misidentified as Tr-Fe by Haumann 1991 and Weigmann 2006) in all instars, with (Paedolohmannia) or without (Eulohmannia) a crease-like vestige of the articulation. In Epilohmanniidae there usually are two apparent segments (Tr-Fe-Ge-Ti fusion) but in some there are seemingly three, as the tibia is only partly fused. Most diverse are the Ptyctima, with three, four or five functional segments (Grandjean 1954a; Hammen 1963; Märkel 1964; Lions and Norton 1998).

The Fe-Ge fusion found in Eulohmanniidae seems the most common type: it exists in all the mixonomatan examples mentioned above in which the palp has either four or three segments. It is also the fusion present in those few Enarthronota with fewer than five palpal segments (Norton et al. 1983). Elliptochthoniidae (Parhyposomata) is a rare exception, in which the single fusion is clearly Tr-Fe. [Haumann 1991 considered Phthiracaridae to have four segments with a Tr-Fe fusion, but this was an error; see Grandjean 1954a; Hammen 1963]. Since the Fe-Ge fusion clearly has evolved independently multiple times, it probably is the optimal location to limit movement, if a less articulated appendage is somehow beneficial.

**R27** — *Cheliceral internalization and emargination*. Chelicerae of acariform mites ancestrally attach to the soft cheliceral frame in a simple fashion, such that the entire appendage extends from the body wall. This is true of Palaeosomata and Parhyposomata, but in Nothrina and Brachypylina (also Astigmata; Norton 1998) the attachment has encroached onto the cheliceral body, such that about the proximal third of the chelicera is internalized (inserted), while the regressed trochanter remains external (Grandjean 1947, his Fig. 2A; Norton and Sidorchuk 2014, their Fig. 11A, B). In most mixonomatans there is no, or very little insertion, but in Collohmanniidae and Eulohmanniidae it is conspicuous.

Another trait seems to correlate well with such insertion: the proximal half of an inserted chelicera is broadly emarginated ventrolaterally on the adaxial face. No functional studies have addressed this, but the combination of insertion and emargination may allow more varied directionality of retractor muscles and therefore cheliceral movements. Another correlation is with Trägårdh's organ, which seems to be well-developed only in groups where the chelicera is internalized and emarginated (R28).

**R28** — *Trägårdh's organ*. This is an elongated, tapering, finger-like oncophysis that evaginates and extends distad from the soft cuticle on the emarginated adaxial face of the chelicera, at the juncture of the cheliceral frame with the trochanter and principal cheliceral segment (Grandjean 1959; Hammen 1968; Alberti et al. 2011). A fully developed Trägårdh's organ is present in all known Nothrina (except Malaconothridae) and Brachypylina. Grandjean (1959) thought the absence of the organ in Malaconothridae represented a loss, presumably because the family is otherwise clearly a member of 'Nothroidea' (in his sense) and Norton (1998) considered their similar absence in Astigmata to be a regressive synapomorphy shared with that family.

Fujikawa (2014) purportedly found Trägårdh's organ in *E. bifurcata*, but we failed to find it in the holotype (see above) and doubt its existence in any Eulohmanniidae. Among Mixonomata, a fully developed Trägårdh's organ is known only in Collohmanniidae (Norton and Sidorchuk 2014), a family that some molecular studies inferred is closely related to, or even within, Nothrina (e.g. Xue *et al.* 2017; Pachl *et al.* 2020). Smaller, less conspicuous, and delicate structures having a rather similar position are found in some Ptyctima (reviewed by Lions and Norton 1998); these structures remain poorly known and whether they represent early, plesiomorphic forms of the organ or independently derived structures remains equivocal. In all questionable cases, the chelicera is neither inserted nor adaxially emarginated (R27), conditions that otherwise correlate well with Trägårdh's organ. The fact that the chelicera of Eulohmanniidae is both inserted and emarginated leaves open the possibility that Trägårdh's organ was lost, as in Malaconothridae.

Lohmanniidae appear to be the only family of Enarthronota to possess a structure that has been referred to as Trägårdh's organ (Ermilov 2017; Ren *et al.* 2017). Grandjean (1950b) first noticed an elongated ridge on the adaxial face of the chelicera in this family, and this ridge ends in a slightly elevated 'boss'. He dismissed it as a possible Trägårdh's organ, though he reserved
judgement on whether there was some degree of homology. Since it is not a hollow, flexible oncophysis and does not evaginate from the juncture of the chelicera with the cheliceral frame, we believe this ridge on the wall of the chelicera is an independently evolved structure and should not be identified as Trägårdh's organ.

**R29** — *Thelytoky*. Usually referred to by the misapplied term parthenogenesis, thelytoky has been the reproductive mode associated with Eulohmanniidae in the literature, which is understandable since our limited knowledge has been associated exclusively with *E. ribagai*. But given newly discovered sexual populations of *E. ribagai*, the probably sexual *Eulohmannia* sp. A (Aborigen; see above), and the sexual type species of *Paedolohmannia*, the ancestral reproductive mode of Eulohmanniidae should be reconsidered. Other extant mixonomatans are mostly sexual (Norton *et al.* 1993): it is the only known mode in Perlohmanniidae and Collohmanniidae and it dominates in Ptyctima (Euphthiracaroidea, Phthiracaroidea). Although Epilohmanniidae include some common thelytokes, probably most species are sexual. Nehypochthoniidae, a monogeneric and phylogenetically relictual taxon, is now the only mixonomatan family that seems to be wholly thelytokous. Outside Mixonomata, a reversal from thelytoky to sexuality was suggested for a group in the nothrine family Crotoniidae (Domes *et al.* 2007), and two others in Enarthronota (Pachl *et al.* 2021) but these are diverse groups and there is both morphological and molecular evidence for reversal. Since Eulohmanniidae is morphologically isolated and species-poor, we have no convincing way to test for reversal within it, but the most parsimonious explanation is that Eulohmanniidae was ancestrally sexual and this mode remains in species and populations from northwestern North America and northeastern Asia, where further diversity should be sought.

## Acknowledgements

We are grateful for the kind help of many individuals who contributed to various parts of this work. Dr. Satoshi Shimano (Hosei University, Tokyo, Japan) translated relevant parts of papers written in Japanese and facilitated our study of type specimens of *Eulohmannia bifurcata*. Alexey A. Gubin (Tyumen State University, Tyumen, Russia) produced most of the scanning electron micrographs. Important loans or donations of materials were provided by Drs. Valerie Behan-Pelletier and Wayne Knee (Agriculture Canada, Ottawa); Dr. Derek S. Sikes (University of Alaska Museum, Fairbanks); Dr. Ken-ichi Okumura (National Museum of Nature and Science, Tokyo); Dr. Heinrich Schatz (Leopold-Franzens University of Innsbruck, Austria); and Dr. Ricarda Lehmitz (Senckenberg Museum, Görlitz, Germany). Drs. Lehmitz and Anna Seniczak (University Museum of Bergen, Norway) also made important observations of juvenile *E. ribagai* from Germany and Norway, at our request. Robin Andrews (University of Alaska, Fairbanks) collected important material from Alaska and helped us interpret glacial refugia. Dr. Jun Chen (Institute of Zoology, Chinese Academy of Sciences, Beijing) clarified Chinese distribution records. We are particularly grateful for constructive comments by Dr. V. Behan-Pelletier on a draft manuscript and for the helpful review by self-identified referee Dr. H. Schatz. This research was partially supported by a grant from the Russian Foundation for Basic Research, project No. 20-54-56054.

## ORCID

Roy A. Norton https://orcid.org/0000-0001-9051-1450

Sergey G. Ermilov https://orcid.org/0000-0002-0913-131X

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