Myrmarachnine jumping spiders of the new subtribe Levieina from Papua New Guinea (Araneae, Salticidae, Myrmarachnini)

Abstract A previously unreported radiation of myrmarachnine jumping spiders from New Guinea is described, which, although having few known species, is remarkably diverse in body forms. This clade is the new subtribe Levieina, represented by seven new species in three new genera. Within Levieagen. n. are three new species, L.herbertisp. n., L.lornaesp. n., and L.francesaesp. n., all of which are unusual among the myrmarachnines in appearing as typical salticids, not antlike. Papuamyrgen. n. superficially resembles Ligonipes Karsch, 1878 or Rhombonotus L. Koch, 1879 as a compact antlike spider, but lacks their laterally-compressed palp and bears an ectal spur on the paturon of the chelicera. Two species of Papuamyrgen. n. are described, Papuamyromhifosgasp. n. and P.pandorasp. n.Agorioidesgen. n., containing A.cherubinosp. n. and A.papagenasp. n., is antlike, with the carapace sunken inwards (concave) between the posterior lateral and posterior median eyes. Phylogenetic analysis of data from the 28S, 16SND1, and COI gene regions of 29 species of myrmarachnines shows that the three new genera form a clade that is sister to the subtribe Myrmarachnina (Myrmarachne sensu lato), with the subtribe Ligonipedina less closely related.


Introduction
The diverse and exquisitely antlike mymarachnine jumping spiders are found around the world, with hundreds of species in the Old World and a few in the Neotropics (Galiano 1969, Wanless 1978, Davies and Żabka 1989, Edmunds and Prószyński 2003, Ceccarelli and Crozier 2007, Edwards and Benjamin 2009, Yamasaki 2012, Yamasaki and Ahmad 2013, Yamasaki and Edwards 2013, Prószyński 2016, Pekár et al. 2017, World Spider Catalog 2018. Most belong to a group that we will treat as the subtribe Myrmarachnina ("Myrmarachne sensu lato"), consisting of Myrmarachne MacLeay, 1839, Belippo Simon, 1910, Bocus Peckham & Peckham, 1892 and several genera recently segregated from Myrmarachne (Prószyński 2016), all typically having forward-projecting male chelicerae and relatively delicate first legs. A smaller set of species falls into a cluster of genera with an embolus-bearing tegular groove, vertical chelicerae and enlarged first legs (including Ligonipes Karsch, 1878 and Rhombonotus L. Koch, 1879), which we treat as the subtribe Ligonipedina, and which forms the sister group to Myrmarachne sensu lato in recent molecular phylogenies Maddison 2012, Maddison 2016) and morphological phylogenies (Edwards and Benjamin 2009). The Ligonipedina is Australasian (Maddison 2015, World Spider Catalog 2018, as is the broader astioid diversification from which the myrmarachnines arose (Bodner and Maddison 2012), suggesting an original diversification of myrmarachnines in Australasia.
Accordingly, it is unsurprising that several distinctive new lineages of myrmarachnines have been found in New Guinea. We describe them here as three new genera, one of which (Leviea gen. n.) is unusual among myrmarachnines for being not in the least antlike. Even though the new levieines have body forms distinctive from other myrmarachnines, general body form is not necessarily a good clue to relationships in myrmarachnines because of their strong selective pressures for mimicry (Ceccarelli and Crozier 2007). Based only on body form, it would not be out of the question for the species described here to have been derived from within the Myrmarachnina. We therefore use molecular data to test the placement of our new species. The first molecular phylogeny of myrmarachnines with more than three species was that of Ceccarelli and Crozier (2007), who found two fairly distinctive clades among the Australian species of Myrmarachne. Subsequent molecular phylogenetic studies were done by Bodner and Maddison (2012), Merckx et al. (2015), Pekár et al. (2017), and Yamasaki et al. (2018), none fully incorporating data from the previous papers. We combine data from these studies to provide a sample of Myrmarachnina that is large enough to test whether the three new genera fall within that subtribe.
This paper began as a presentation in a symposium honouring Herbert W Levi in the 20 th International Congress of Arachnology (2016). Accordingly, it is dedicated to Levi and his service to arachnology. All of the specific epithets make reference to him and his family.

Material and methods
The bulk of the material examined came from two expeditions to Papua New Guinea, one in 2008 (Maddison 2009, Maddison andZhang 2011), the other in 2013 (Leponce et al. 2016), although a few specimens date back to 1968 (Balogh 1971). Preserved specimens were studied from these collections: Spencer Entomological Museum of the University of British Columbia (UBC-SEM, curator Wayne Maddison), Royal Belgian Institute of Natural Sciences (RBINS, curator Wouter Dekoninck), and Hungarian Natural History Museum (HNHM, curator László Dányi).
Specimens were examined under both dissecting microscopes (Nikon S800, Olympus SZ61) and compound microscopes (Nikon ME600L, Nikon Eclipse E200) with reflected light. Drawings were made with a drawing tube on a Nikon ME600L compound microscope.
Terminology is standard for Araneae. All measurements are given in millimeters. Carapace length was measured from the base of the anterior median eyes not including the lenses to the rear margin of the carapace medially; abdomen length to the end of the anal tubercle. The following abbreviations are used: PLE, posterior lateral eyes; PME, posterior median eyes (the "small eyes"); RTA, retrolateral tibial apophysis.
Taxa included in the molecular phylogeny are listed in Table 1, which also lists taxonomic authority for the species. The taxon sample includes one specimen of each of Leviea, Agorioides, and Papuamyr, other myrmarachnines, a set of astioid outgroups and a few more distant species from the Salticoida. The sequenced specimens of Leviea francesae and Agorioides papagena are the holotypes; that of Papuamyr omhifosga is a paratype from the type locality. The Ligonipes specimen bears close resemblance to the male figured by Davies and Żabka (1989). The Rhombonotus specimen (from Australia, 26.235S, 152.640E) differs from the drawing in Davies and Żabka (1989) in having a slightly smaller spermophore loop. Except for the three new genera, most sequences come from already published works. It is not our purpose to resolve the phylogeny of Myrmarachne sensu lato, and so we include a small sample of 24 species, diverse enough to test whether the levieines are inside or outside the group. We include most Myrmarachne sensu lato used by Maddison (2016), and a selection of species from sequences obtained by Ceccarelli and Crozier (2007) (with identifications from Ceccarelli 2010), Jang andHwang (2011), Merckx et al. (2015), Pekár et al. (2017), and Yamasaki et al. (2018). From the published sequences we excluded specimens known only from COI because of its poor phylogenetic performance when analyzed alone (Hedin andMaddison 2001, Maddison et al. 2014). We also excluded duplicates of species and most lacking an identification to species or species group. The only new data from Myrmarachne sensu lato are from a Neotropical species (M. cf. mocambensis from Ecuador, 2.9962S, 78.4558W), Emertonius malayanus (from Malaysia, 4.04N, 114.816E), and Myrmarachne cornuta (from Malaysia, 4.0432N, 114.8110E).

Phylogeny
The phylogenetic tree (Fig. 1) shows three major groups of myrmarachnines: Ligonipes plus Rhombonotus, the Leviea group of genera, and Myrmarachne sensu lato. We consider these to represent three subtribes, as described below. The three subtribes and several subgroups of the Myrmarachnina are well supported according to bootstrap values ( Fig.  1) and independent support by both 28S (Fig. 2) and 16SND1 (Fig. 3). As expected from previous studies (Hedin and Maddison 2001;Maddison et al. 2008Maddison et al. , 2014Bodner and Maddison 2012), the gene COI on its own ( Fig. 4) has discordant results, scattering the myrmarachnines and with only 53% of myrmarachine nodes in agreement with the all genes tree (compared to 87% and 69% for 28S and 16SND1, respectively). Within the Myrmarachnina, a clade of Australian Myrmarachne species (the erythrocephala group) stands as sister group to the remainder. This deep split between a handful of Australian species and the rest of the Myrmarachnina is reflected as well in the results of Ceccarelli and Crozier (2007) and Pekár et al. (2017). These results suggest that the Myrmarachnini, and the Myrmarachnina, radiated initially in Australasia, as expected given that most astioid diversification is Australasian (Bodner and Maddison 2012). Our results differ from the morphological results of Edwards and Benjamin (2009) in placing Myrmapana outside the clade containing the bulk of Old World species, and in placing Belippo within Myrmarachne itself (as it had been conceived in 2009).
Morphological synapomorphies are not known for the Levieina, except possibly the ectal spur on the paturon of the chelicera, and yet this group is well supported by the molecular data. Its diversity of body forms arguably exceeds that of the much more species-rich Myrmarachnina, and its molecular divergences are as deep. These suggest the  Table 1) 1 phylogeny from all 4 genes concatenated; bootstrap percentages shown if ≥ 75% 2 phylogeny from 28S analyzed alone 3 16SND1 alone 4 COI alone.
levieines are an old radiation, with possibly many more species to be discovered in Australasia. Among forms we have seen (but do not describe here) are two more species of Leviea and three other species that may belong to Papuamyr, one of which is beetle-like.
The sister group relationship between Agorioides and Papuamyr is well supported in the molecular phylogeny, but we know of no morphological traits that support it. Indeed, some notable morphological characters are variable within the Levieina and discordant with the phylogeny (Ligonipedina, ((Leviea, (Agorioides, Papuamyr)), Myrmarachnina)). These include the antlike body (absent in Leviea but present in Ligonipedina, Agorioides, Papuamyr, Myrmarachnina, and absent outside the Myrmarachnini), the ectal spur on the paturon (present in Leviea, Papuamyr, and some Myrmarachnina), the RTA having a ventral flange (present in Ligonipes, Leviea, Papuamyr, and various Myrmarachnina, e.g., Wanless 1978 figure 3F), and the swollen first legs (absent in Leviea, femur-only in Agorioides, present in Papuamyr and the Ligonipedina). The lack of an antlike body in Leviea is most parsimoniously interpreted as a loss, given the antlike bodies in almost all other myrmarachnines, including its sister group, its first cousin (Myrmarachnina), and its second cousin (Ligonipedina). A loss in Leviea is perhaps not surprising, given that they are known from high elevation (2320-3700 m). In these cool mossy forests, ants are less visible and less diverse than at lower elevations (Orivel et al. 2018).

Tribe Myrmarachnini Simon, 1901
Toxeae, Toxeinae FO Pickard-Cambridge, 1900 (replaced due to synonymy of the type genus; see Maddison 2015). Myrmarachneae Simon, 1901 The Myrmarachnini are characterized by antlike bodies and distinctive genitalia (Edwards and Benjamin 2009). The palp has a round bulb and an immovable embolus whose terminal loop comes loose from the periphery and passes in front of (i.e., ventral to) the tegulum before terminating. The RTA is twisted and bent, frequently with a ventral flange that is sometimes developed into a distinct apophysis. The characteristic female genitalia (Wanless 1978 figure 3E) have separate atria that lie posteriorly, and lead to membranous copulatory ducts (Edwards and Benjamin 2009, character 8 state 1), sometimes irregular and voluminous, which eventually make their way to the posterior margin. The duct then narrows into a sclerotized spermathecal complex (Edwards and Benjamin 2009) that first takes the form of a narrow tube as it proceeds forward along the midline and before expanding to a bulb when it is are anterior to the atria. Prószyński (2016) considers the sclerotized tube to be part of the spermatheca, which are thus "pipe-like" in shape, while Edwards and Benjamin (2009) consider the tube to be part of the copulatory duct.
We have chosen to divide the tribe into three subtribes to reflect the group's diversity, and to provide a formal name for the clade that would otherwise have only the informal name "Myrmarachne sensu lato". A formal name for the clade is particularly urgent, now that the generic name "Myrmarachne" has lost its biological meaning following Prószyński's (2016) separation of twelve genera from Myrmarachne. Our results indicate that Myrmarachne in Prószyński's (2016) sense is not a monophyletic group. This is not surprising, given that Prószyński's dismantling of the genus was done without considering synapomorphies, quantitative analysis, or previously published phylogenetic analyses (Ceccarelli and Crozier 2007;Edwards and Benjamin 2009;Merckx et al. 2015), but rather by carving out some groups based on few characters. At present, the word "Myrmarachne" means merely "The set of Myrmarachnina species that remain in Myrmarachne, either because a more specific place for them has not been chosen, or because they are related to the type species Myrmarachne melanocephala MacLeay, 1839". (We do not have molecular data from M. melanocephala, and Prószyński's account does not guide us as to where in Fig. 1 it might fall.) Because of uncertainty about Prószyński's new arrangement, Yamasaki et al. (2018) chose to ignore it and describe new species under the old (broader) concept of the genus Myrmarachne. We are now in the unsatisfactory situation in which a generic name has two meanings: in our current taxonomy (e.g., as documented in the World Spider Catalog, 2018), it refers to an unnatural conglomerate, but informally (when we use the phrase Myrmarachne sensu lato), it refers to a natural group (when Bocus and Belippo are included). The informal use should be avoided, as it misleads us into thinking we are relieved of the responsibility of repairing Myrmarachne.
While we could synonymize all Myrmarachnina back into a single genus, we have chosen not to change the current generic arrangement pending further study. The subtribe is diverse enough that it will almost certainly be split into multiple genera eventually, though possibly very differently than in Prószyński's arrangement. (The erythrocephala group, for instance, could be its own genus, unusual among Myrmarachnina in retaining the excavate male chelicerae possibly ancestral for the tribe; see Pekár et al.'s (2017) figures 5H, 7E, and 13F.) By describing the subtribe, we provide a name for the clade that is a stable alternative to the phrase "Myrmarachne sensu lato". Because of the lack of clear meaning of "Myrmarachne", all of the species described by Pekár et al. (2017) and Yamasaki et al. (2018) would be better considered Myrmarachnina incertae sedis than members of Myrmarachne, but our nomenclature does not permit a species to be without generic placement. For this reason, we list their generic placement explicitly as "Myrmarachne (sensu lato)". Diagnosis and synapomorphies. Edwards and Benjamin (2009) indicate two distinct synapomorphies for the Ligonipedina: a bend in the duct-like portion of the spermathecal complex and medially placed atrial rims. These characters provide only weak support for the group as a whole, though if Damoetas were removed, the remaining genera would form a tight group well supported by a distinctive feature: the male palp laterally compressed, and with a groove in tegulum in which the last loop of the embolus rests as it passes over the tegulum. Ligonipes, Rhombonotus, and Judalana are also united by the first leg being swollen, especially patella and tibia, with ventral fringe of hairs.

Diagnosis and synapomorphies.
There are no unambiguous morphological diagnostic traits known for the levieines, though there are molecular traits. We tentatively suggest as a synapomorphy of this group the ectal spur on the male cheliceral paturon (white arrow in Figs 13,29,71,72,83,84). It is absent in Agorioides, in which, judging by phylogenetic position, the spur could be secondarily lost. Such a spur is seen also in some Myrmarachnina (see illustrations of Wanless 1978): Belippo spp., Hermosa andrewi (Wanless, 1978), Myrmarachne lesserti Lawrence, 1938, and Myrmele eugenei (Wanless, 1978). Given the apparently scattered distribution of this trait, it could be convergent with that in levieines. Other than this, no morphological traits are known to unite levieines. In the gene 28S, the three levieine genera are unique among myr-  Levi (or, Herb, as he humbly preferred to be known by) was one of the grand arachnologists of the twentieth century, describing over 1200 species of spiders, mentoring many subsequent leaders of the field, and curating one of arachnology's most important museum collections (Leibensperger 2016). Lorna collaborated in his work in many ways, coauthoring the classic book Spiders and their kin (Levi and Levi 1968), which introduced the first author of this paper to spider diversity. Frances accompanied them in the field and carried on an interest in woven creations. Their contributions, both personal and scientific, will long be remembered (Maddison 2014a, b;Leibensperger 2016). The Levis pronounced the vowels of their name approximately as their IPA equivalents (e as in Ed, i as in eat). The last three letters of Leviea are to be pronounced as three separate vowels (as their IPA equivalents, i-e-a). The name is to be treated grammatically as feminine.
Diagnosis. The form of the body is not in the least bit reminiscent of an ant, beetle or wasp, unlike other myrmarachnines. Instead, the body is of standard salticid form (e.g., Icius Simon, 1876, Salticus Latreille, 1804), somewhat glabrous, with chevron markings. Two features possibly retained from antlike ancestors are a female palp that is widened and somewhat dorso-ventrally flattened, and the many long macrosetae on the first tibia in two of the Leviea species. The male embolus is distinctive for ending broadly, not tapering to a point. As in Papuamyr, there is an ectal spur on the paturon (white arrow in Figs 13, 29). Diagnosis. Body somewhat smaller, less elongate, than the other species of Leviea, and with fewer (3 or 3.5 pairs) ventral macrosetae on first tibia. The robust dorsal branch of the RTA ends bluntly like a thumb, unlike the tapering and dorsally pointing tip of the other species. Bulb ca. 90° further rotated than in L. lornae and L. francesae, as indicated by the spermophores (Figs 5, 21, 38).
Description. Male (holotype). Carapace length 1.82; abdomen length 1.96. Carapace (Figs 12, 14, 15): Of typical salticid shape, without constrictions or thoracic hump. Chelicera (Fig. 13): Vertical, simple except for a small ectal spur on the paturon. At least five retromarginal teeth. Palp (Figs 5, 6). With round bulb. Embolus circling 1.8 times around, ending in broad tip. RTA appearing as a finger and thumb (dorsal apophysis and ventral flange, respectively), with the dorsal apophysis pointing somewhat ventrally. Legs unremarkable, first somewhat more robust. First tibia with three pairs of ventral macrosetae, of normal length (right tibia bears an extra small distal anterior macroseta). Colour in life (Figs 9-12): Markings are muted beige to reddish brown, with the digestive diverticula in the ocular area providing two pale stripes that continue as pale areas on the thorax and chevrons on the abdomen. Colour in alcohol ( Fig. 13-15): Body and appendages honey coloured, with indistinct dark annuli on the legs, and chevrons on the abdomen. Underside of abdomen with two longitudinal dark stripes.
Female (paratype, specimen PNG2008-0370). Carapace length 1.89; abdomen length 2.16. Carapace, legs, abdomen substantially as in male (Figs 16-18, 20), except for the less robust first legs. First leg tibia with seven ventral macrosetae (three pairs plus one prolateral-distal) (Fig. 19). Epigyne (Figs 7, 8  Diagnosis. Larger and more slender than L. herberti, with more (5 pairs) ventral macrosetae on first tibia. In these features it resembles L. francesae, from which it differs in details of the palp: dorsal branch of RTA slender and distal-pointing; bulb rounder than L. francesae, less rotated than L. herberti. Epigyne with openings further posterior than those of L. herberti, and smaller RTA pocket at rear margin.
Female (paratype in UBC-SEM). Carapace length 1.89; abdomen length 2.45. Carapace (Fig. 33) Etymology. Named in honour of Frances Levi. Diagnosis. Larger and more slender than L. herberti, with more (5 pairs) ventral macrosetae on the first tibia. In these features it resembles L. lornae. Differs from both L. herberti and L. lornae in details of the palp: dorsal branch of RTA swollen basally; bulb oval rather than circular, less rotated than L. herberti.
Description. Male (holotype). Carapace length 2.08; abdomen length 2.38. Carapace (Fig. 41): Like that of L. lornae. Chelicera: Ectal spur on paturon small. Five retromarginal teeth. Palp (Figs 38, 39): Bulb compressed slightly laterally to be oval rather than circular. Embolus thick, circling 1.7 times around. First leg tibia with five pairs of ventral spines. Colour in alcohol (Fig. 41) Diagnosis. Antlike, with concave-sided carapace, swollen first femur, a long ocular quadrangle, long fourth trochanters, and a spinose first tibia. The carapace is sunken inward (concave) between the PME and PLE, leaving the PLE on prominent tubercles, and yielding a constriction that resembles that of hisponine salticids. The femur of the first leg is shaped like a bird's lower leg ("drumstick"), swollen in the proximal half but thin distally. The length of the ocular quadrangle is distinctly more than half the length of the carapace. The fourth trochanter is unusually long, longer than either the coxa or the fourth tarsus. Unlike Leviea and Papuamyr, the paturon of the chelicera lacks an ectal spur, and the first tibia has many pairs of long macrosetae; both of these features can be found in some species of Myrmarachnina.
Remarks. The two species described are closely similar, but distinct in the form of the palp, shape of the carapace, and in colour. They were found only seven km apart, but at distinct elevations (570 m vs. ca. 1000 m). In other salticid genera, closely related species have been observed to segregate along such an elevational gradient in the same area (e.g., Cucudeta zabkai Maddison, 2009vs. Cucudeta uzet Maddison, 2009  Etymology. In the Levis' country home they hosted many animals, domesticated and not. One of these was a dog of multifarious ancestry, Cherubino, named after the character in Mozart's opera The Marriage of Figaro. The spider resembles the dog in having a hairy, grizzled appearance. Diagnosis. Differs from A. papagena in having the tibia of the male palp distinctly narrower than the cymbium (Figs 42 vs. 51), the cephalic area distinctly higher (Figs 44 vs. 53), and in the black body with a dusting of white setae. The bulb of the palp is rotated slightly more than in A. papagena (see Diagnosis for that species).
Description. Male (holotype). Carapace length 2.16; abdomen length 2.27. Carapace (Figs 44-50): Strangely shaped, as described for the genus, with a constriction between the PME and PLE. Thoracic hump prominent (Fig. 44). Ocular quadrangle occupies more than half of the length of the carapace. Clypeus extremely narrow. Chelicera: Vertical, simple. Teeth not examined for fear of damaging the specimen. Palp (Figs 42, 43): Embolus wrapping around bulb more than once; RTA simple and unbranched. Legs with relatively few short setae, except a greater density yields weak fringes beneath metatarsi and tarsi 2-4 and tibiae 3-4. First tibia with seven pairs of long ventral macrosetae; first metatarsus with two pairs. First femur distinctly swollen in proximal half, shaped like a bird leg's drumstick (Figs 47, 49). Fourth trochanters distinctly longest, longer than the fourth tarsi (Figs 44, 48). Colour in life (Figs 48-50): Black body with fully or partly erect white setae on the sides and back of the carapace, the abdomen, and the femora of the fourth legs. First and fourth femora black; other segments translucent white to honey with darker lines or patches. Colour in alcohol (Figs 44-47): Carapace brown with black around eyes. Abdomen black except paler around the constriction and anteriorly. Clypeus dark, with only a few setae. Chelicerae honey coloured. Palp black. Legs pale except black on proximal half of first femur, brown to black fourth femur, and black lines on anterior faces of legs 2-4 and posterior faces of legs 3 and 4.
Female. Unknown. Additional material examined. One penultimate instar male (specimen PNG2008-2765, in UBC-SEM), with same data as holotype, also from leaf litter. It has a black body and leg markings as in holotype, but is more glabrous, with only a few white setae. Etymology. The Levis' love of opera was reflected in their animals' names, including their dog Papagena, named after the character in Mozart's The Magic Flute. The spider does not look like the dog or the opera character. Although the holotype is a male, the species is named for the female that remains to be found.

Agorioides papagena
Diagnosis. Differs from A. cherubino in having the tibia of the male palp almost as wide as the cymbium (Fig. 51), flatter profile of the carapace (Fig. 53), and a distinctly orange body and legs. The bulb of the palp is rotated less in A. papagena than in A. cherubino, as seen by the orientation of the spermophores in Fig. 51 versus Fig. 42. Although this could be due to a slight expansion of the palp of the one known specimen of A. papagena, suggested by the offset of the tip of its embolus from the cymbial apical groove, this offset is of lesser angle than the difference in spermophore orientation.
Description. Male (holotype). Carapace length 2.84; abdomen length 3.24. Structure of body, legs as in A. cherubino , with first leg having swollen femur (Figs 57,58). Chelicera (Fig. 54): Vertical, though robust. Four retromarginal teeth. Palp (Figs 51,52): Embolus wrapping around bulb more than once; RTA simple and unbranched. Tibia distinctly wider than in A. cherubino. Colour in life (Fig. 58): Orange, darkening to brown on the abdomen and with black around the eyes. Swollen femur of the first leg particularly bright orange. Some partially-erect white setae on carapace sides beneath the eyes, on thorax, and on fourth femora, but not as dense or distinct as in A. cherubino. Colour in alcohol (Figs 53,54,56,57): Except for black around the eyes, carapace, legs and palpi are orange, darkest on the thorax and palest at the leg tarsi. Abdomen a muted orangish gray.
Female. Unknown. Additional material examined. One juvenile (specimen PNG2008-1676, in UBC-SEM, Fig. 59), similarly coloured, with same data as holotype, on leaf litter. 90 Description. A single large female from the Muller Range is clearly an Agorioides by carapace shape and long fourth trochanters (the first legs are missing), but is not formally described here because the specimen is missing most of its legs. It seems likely to represent a distinct species, as it has a carapace that is flatter (Fig. 65) than that of the two described above. It is notably larger than A. cherubino (carapace 3.00, abdomen 3.43). Its epigyne is typical myrmarachnine, with the RTA pocket displaced anteriorly (Figs 62, 63 Diagnosis. Small antlike salticids with somewhat swollen male first legs, vertical and excavated male chelicerae with an ectal spur on the paturon, a round bulb on the palp, and a relatively long RTA. In contrast, the Ligonipes group of genera has the bulb of the male palp compressed laterally (Davies and Żabka 1989: figs 7 and 9), while most Myrmarachne males have projecting chelicerae and thin first legs. An apparent synapomorphy uniting the two Papuamyr species described below is a complex fold in the embolus at the point where it crosses over previous loops before terminating (see black arrows in Figs 70 and 78).

Agorioides sp.
Remarks. Papuamyr lacks the morphological diagnostic characters of the Ligonipedina and Myrmarachnina, but we have not found morphological characters that place it more clearly. The molecular data, however, are clear that Papuamyr is within the Levieina and a close relative of Agorioides (Fig. 1).

Etymology.
A combination of letters derived from the first letters of words in a statement of Herbert Levi's from approximately 1985. He had forgotten to do some bureaucratic task, and in dismay, he exclaimed "Och, my head is full of spider genitalia!" Truly, it was, and for that knowledge which he conveyed to us, arachnology is forever enriched. The name is particularly apt for this species: its genitalia are distinctive and elegant, the palp having a long transparent RTA and sharply bent embolus, the epigyne an RTA pocket displaced far to the anterior.
Diagnosis. Distinct for its unusual genitalia and its orange and black bicoloured body. The embolus tip extends onto the retrolateral side of the cymbium before looping back to terminate ventrally, much as seen in many amycoid salticids such as Tartamura (Bustamante and Ruiz 2017). Most notable, however, is the long RTA whose posterior branch extends almost to the tip of the cymbium (Fig. 67). Accordingly, the RTA pocket of the epigyne is displaced far anteriorly (Fig. 68) constriction (Fig. 72). Carapace glabrous and shiny, without evident microsculpture. Ocular quadrangle less than half the length of the carapace. Clypeus extremely narrow. Chelicera (Fig. 71): Excavated medially, and with an ectal spur on the paturon. Five very small retromarginal teeth. Palp (Figs 66,67,70): Bulb round, with embolus with a sharp bend near the terminus (black arrow in Fig. 70), at the point where it passes a previous loop, just before extending to the retrolateral side of the cymbium. Ventral lobe of RTA projects ventrally; dorsal lobe extremely long. Legs with relatively few, short setae. First legs distinctly more robust than others. First tibia with two short anterior ventral macrosetae in the distal third, and two extremely small posterior ventral macrosetae in the proximal two thirds. Colour (Figs 71-75): Orange (vibrant in life, pale in alcohol) except for the dark cephalic region, the black posterior half of the abdomen, and dark lines on legs 3 and 4.
Etymology. From the Greek "all giving" or "all gifted", referring primarily to the gifts given us by the Levis, most memorably the 1254 species that Herb described, and the wonderfully abundant Thanksgiving celebration dinners that Lorna put on for their students each year. It is also the name of the dog Lorna had when Herb and Lorna were married.
Diagnosis. Distinct in having the thoracic hump higher than the ocular area in both sexes (Figs 84,87,89). Males can be recognized by the shape of the RTA, with dorsal branch shorter than in P. omhifosga and twisted, and a very short ventral flange (Fig. 80). Females can be recognized by the lack of anterior pocket, the relatively large spermathecae (Fig. 81), and the long lateral extension of the sclerotized ducts near the posterior margin (Fig. 82).
Description. Male (holotype). Carapace length 1.52; abdomen length 1.41. Carapace (Figs 83-85): Thoracic slope with a large hump which is higher than the flat ocular area (Fig. 84). Ocular quadrangle with rugose integument (Fig. 85), approximately as long as wide, occupying half of the carapace (0.75 long). Clypeus extremely narrow (  looped twice around it, with a twist on the prolateral side of the bulb (black arrow Fig.  78) and a slight bend at the end. Ventral flange of RTA small, a slight hump; dorsal lobe well developed, with an elongated "S" shape. Legs: Leg I robust, each segment is at least twice as wide as other legs. Other legs slender. Abdomen: pear shaped, with a slight con-striction at the anterior fourth/third. Colour (Figs 85-87 in alcohol): Body dark brown with a transverse pale band in the middle of the carapace and abdomen. Coxa I, leg II, proximal half of coxa IV, patella III-IV, metatarsus III, and all tarsi pale yellow. Trochanter and femur I dark brown, patella and metatarsus I dark yellow with a dark ventral side. Leg III-IV black except the pale yellow segments mentioned earlier (Figs 85-87).