Liphanthus sapos Mir Sharifi & Packer, 2019

Sharifi, Negar Mir, Graham, Liam & Packer, Laurence, 2019, Fifteen new species of Liphanthus Reed (Hymenoptera: Andrenidae) with two submarginal cells, Zootaxa 4645 (1), pp. 1-80 : 12-16

Liphanthus sapos Mir Sharifi & Packer , sp. nov.

urn:lsid:zoobank.org:act:64659156-3FC3-4D9A-93D4-1C7F70E71B2F

Figs. 20–30 , 128 , 136 , 138 , 175–176 .

Diagnosis: Males of this species can be differentiated from all others of the genus, except L. jenamro , by the combination of two submarginal cells ( Fig. 20 ), metatibial spurs strongly curved at apices (as in Figs. 28–29) and metasomal sterna yellow ( Fig. 20 ). Other species with two submarginal cells, except L. jenamro , have either the metasomal sterna brown or the metatibial spurs only weakly curved. It can be differentiated from L. jenamro by the first flagellar segment shorter or equal to its width and the clypeus and lower 4/5 of the subantennal sclerite yellow ( Fig. 21 ) whereas in L. jenamro the first flagellar segment is longer than wide and the subantennal sclerite is black ( Fig. 11 ), rarely with a small apical pale spot. As for L. jenamro , if L. domeykoi , described above, is also considered to have strongly curved metatibial spurs, then L. sapos can be differentiated from it by the almost entirely blackbrown trochanters ( Fig. 20 ) which are yellow to yellow-brown ventrally in L. domeykoi ( Figs. 31 &33 ). Females of this species can be differentiated from all other species except L. jenamro by the combination of two submarginal cells and metatibial spurs strongly curved at apices and pronotal lobe yellow (Fig. 26). Females can be differentiated from those of L. jenamro by the metatibia with reduced dark markings and the posterior metatibial spurs with broad teeth, as long as wide (Figs. 28–29). For L. jenamro females, the metatibia (except base yellow and apex reddish) is black and the posterior metatibial spurs have narrow teeth, twice as long as their basal width ( Fig. 18–19 ).

Description. Holotype Male: Dimensions: Approximate body length: 4.04mm; head width: 1.16mm, wing length: 2.63mm, intertegular width: 0.73mm.

Coloration: Black to dark brown with following parts yellow: labrum, mandible (except apex orange-red and extreme base dark brown), clypeus, lower paraocular area below from basal 1/4 of outer subantennal suture, subantennal sclerite (except upper 1/5), pronotal lobe, anterior spot on tegula, small apical mark on [procoxa] metacoxa and metatrochanter, apical ring on all femora, [pro-] and mesotibiae (except brown for most of ventral surface at midlength), metatibia (except small brown mark on posteroventral surface), all basitarsi and second tarsomeres (third yellow-brown, fourth and pretarsi orange-brown), ventrally reflexed portions of metasomal terga, T7 and S2–S6 (S1 red-brown). Ventral surface of flagellum yellowish, dorsal surface red-brown to orange-brown. Apical impressed areas of metasomal terga amber, narrowly margined with orange anteriorly.

Sculpture: Yellow portion of face and narrow strip on lower paraocular area adjacent to compound eye lacking microsculpture, shiny; black portion of face densely minutely tuberculate, dull except supraclypeal area below lower tangent of antennal sockets weakly imbricate shiny; clypeus and yellow portion of paraocular area punctures variable in size, mostly large, shallow, irregularly spaced, i=0.2–2d; subantennal sclerite with scattered, shallow punctures; punctures on rest of face and vertexal area shallow, difficult to detect among coarse imbrication, i>d; genal and hypostomal area weakly imbricate, shiny; punctures shallow, irregularly spaced, i=1–6d. Mesosoma imbricate, generally dull; mesoscutum punctures shallow, irregularly spaced, i=0.5–2.5d; scutellum imbricate, somewhat dull, sparsely and obscurely punctate, i>2d; metanotum punctures obscure, irregularly spaced, i=0.2–2d; hypoepimeral area impunctate; rest of mesopleuron and metapleuron punctures shallow, sparse, i>2d; propodeum distinctly and densely punctate below, i~d, obscurely punctate elsewhere. Metasomal terga strongly imbricate, dull becoming more weakly imbricate and somewhat shiny from T1–T7; punctures shallow, obscure among coarse imbrication; T7 punctures distinct, moderately dense, i=0.5–2d; metasomal sterna weakly imbricate, shiny; punctures minute, scattered.

Pubescence: Generally white, sparse with short branches; ~1.5 MOD on most of head and mesosoma , longer on scutellum and mesopleuron, ~2 MOD; short and laterally oriented on dorsal surface of metapostnotum; simple, short, laterally oriented on metasomal terga.

Structure: Head: almost as long as wide (73:73). Mandible length to basal depth 51:24, lower margin at midlength with erect pencil of branched hairs, branches long and on only one side of rachis, outer ridge lamellate. Labrum rectangular ~ 1.5 X as broad as long (33:21). Clypeus ~ 1.7 X as wide as long (66:38); apicolateral and apicomedial margins almost straight. Outer subantennal suture strongly curved near base, otherwise straight, inner suture straight, sutures convergent below, subantennal sclerite widest near base; epistomal suture curved between inner subantennal sutures.Anterior tentorial pit below junction of outer subantennal and epistomal sutures by almost minimum diameter of F1. Frontal line imperceptible for most of length. IAD subequal to AOD (17:18). Inner margin of compound eyes convergent below, UOD:LOD 84:76. Facial fovea small, elongate, 4X longer than wide, subparallel to inner margin of compound eye. IOC: OOC 22.5:20. Scape 1.5 X as long as greatest width (20:13), shorter than pedicel and F1 combined (17); pedicel shorter than wide (9:10), F1 and F2 with length and width subequal, remaining flagellomeres slightly longer than wide except F11 ~ 1.5 X as long as width (19:13).

Mesosoma: Mesoscutum ~1.15 X as wide as long (72:63), length of scutellum: metanotum: metapostnotum: 27:15:18. Marginal cell longer than distance between its apex to wing tip (69:59). Metatibial spurs strongly curved at apices, subequal in length. Posterior metatarsal claw with axe-shaped tooth, anterior claw with tooth broad; claws of other tarsi cleft.

Metasoma: Broadest at midlength of T3. Pygidial plate absent. S1–S5 unmodified, S6 apicomedially translucent and weakly concave, premarginal line bearing few setae along sides of concavity and apically. S7. Apodeme long and broad, anterior margin obtuse; apical lobe slightly longer than wide, broadly rounded, posterior margin with sparse moderately short posteriorly oriented hairs, hairs less than half width of lobe. S8. Anterior margin sinuate and broad, anterolateral margins strongly diverging anteriorly; lateral lobe small, apex acute, recurved anteriorly; apical lobe broadly rounded, tongued-shaped, lateral margins convex, short spicules on apical half of dorsal surface. Gonocoxa anterodorsal margin transverse, lateral margins straight, diverging posteriorly, medial margin almost straight; gonostylus distinctly separated from gonocoxa by weakly sclerotized band, narrow, gradually narrowing to apex, posteromedially curved, short hairs on medial and ventral surfaces; penis valve broad, lateral margin broadly biconvex in dorsal view. Endophallus anterior sclerotized fluting long and narrow, apical membranous portion not extending to apex of penis valve ( Fig. 25 ).

Allotype Female: As in male except for usual sexually dimorphic features and as follows:

Dimensions: Approximate body length: 4.81mm; head width: 1.21mm, wing length: 3.13mm, intertegular width: 0.85mm.

Coloration: Black with following parts yellow: mandible (except apical 1/4 orange-red, third quarter orange), pronotal lobe, anterior spot on tegula, extreme apex of trochanters, apical ring on all femora, dorsal surface of basal 2/3 of protibia, basal ring on probasitarsus, basal ring on mesotibia and marks on reflexed portions of T3–T5. Following parts yellow-brown: apicoventral spot on F1–F3, ventral surfaces of F4–F10, probasitarsus medially, apical ring on mesotibia, base of mesobasitarsus, basal1/3 and apical ring on metatibia. Dorsal surface of F3–F10 red-brown. T3–T6 orange-brown, metasomal sterna orange. Apical impressed areas of metasomal terga amber, margined anteriorly by orange.

Sculpture: Face below antennal socket and upper paraocular area adjacent to compound eye mostly lacking microsculpture, shiny; rest of face acinose to minutely tuberculate, dull; clypeus punctures variable in size, moderately dense, i=1–1.5d; lower paraocular area punctures more distinct, moderately sparse, i=1–2d; subantennal and supraclypeal areas punctures mostly small, spacing variable, i=0.5–2d; genal and hypostomal areas weakly imbricate, shiny, with scattered shallow punctures, i>2d. Mesoscutum imbricate, dull, punctures obscure, i=1–3d; scutellum more weakly imbricate, punctures more distinct, variable in size and spacing, i=1–4d; metanotum imbricate, dull, punctures small and obscure; sides of thorax imbricate, somewhat dull, hypoepimeral area rugoso-punctate anteriorly, rest of mesopleuron minutely, sparsely punctate, i>2d; metepisternum obscurely punctate. Metapostnotum dorsal surface rugulose, Propodeum imbricate, somewhat shiny. T1–T5 increasingly weakly imbricate, T1 dull, T5 somewhat shiny, with minute, scattered punctures except T5 distinctly punctate, i>d basally, i~d apically; metasomal sterna weakly imbricate, shiny, punctures small, i>d.

Pubescence: Seemingly abraded on most of head and mesosomal dorsum, otherwise longest on tibial scopa ~2.5 MOD; <1.5 MOD on genal area and sides of mesosoma . Dorsal surface of metapostnotum hairs plumose, short, ~0.5 MOD. Prepygidial fimbria pale straw, ~1.6 MOD.

Structure: Head: slightly wider than long (86:82). [Mandible length not measured due to degree of abrasion of apex of pollex]. Labrum ~1.5 X as wide as long (37:26), raised area sides straight, convergent below, curved to transverse apex. Clypeus more than 2 X as wide as long (80:35). Outer subantennal suture straight, inner subantennal suture briefly outwardly concave near antennal socket. Frontal line narrowly depressed below, absent above. IAD= AOD (15:16). Inner margin of compound eyes subparallel, UOD:LOD 72:73. Facial fovea shallow,> 5X longer than greatest width, 24:4.5, which is above midlength, lateral margin straight and parallel to inner margin of compound eye, medial margin convex. IOC: OOC 21:18. Scape ~ 3 X as long as greatest width, (26:9), longer than pedicel and F1 combined (18); pedicel as long as wide (9:9), F1 length subequal to width, (9:8), F2–F9 slightly longer than wide to length and width subequal, F10 longer than wide 19:12. FIGURES 26–29. Liphanthus sapos sp. nov., allotype female, 26: lateral habitus, 27: head frontal view, 28: apex of tibia to show curved metatibial spurs, 29: ESEM of metatibial spurs. Scale bars Figs. 26–27 1 mm, Fig. 28 0.25mm, Fig. 29 50μm.

Mesosoma: Mesoscutum ~1.2 X as wide as long (85:70), length of scutellum: metanotum: metapostnotum: 25:16:17. Tarsal claw teeth short.

Metasoma: Metasomal terga and sterna unmodified. [Pygidial plate narrowly rounded, sides concave].

Etymology. The species is named after the type locality for which the nearest named site is Los Sapos, approxi- mately 13.5km to the East. “Los Sapos” means, “the toads”, perhaps referring to an area of unusual dampness near where the absolute desert reaches further inland and at higher altitude than anywhere else in the countery.

Material studied. Region III: Holotype male, allotype female and 4 male and 1 female paratypes as follows : Holotype male: CHILE, Region III , 13.5km W. of Los Sapos -28.019, -70.554, 488m, 12.x.2010, L. Packer, CCDB B09997- H1 . Allotype female and two male paratypes: CHILE, Region III , Atacama, SW of Pedernales , -26.44190, -70.35372, 2687m, 1.xi.2015, L. Packer, CCDB 28312 -B09, B11 and H05. Paratype male: same data as holotype except CCDB B09997- G11 ; paratype male: CHILE, Region III , W. of El Salado , -26.395, -70.388, 118m, 16.ix.2010, L. Packer, CCDB B09866 C09 (specimen badly damaged) . Paratype female: CHILE, Region III , Cuesta Montandon , -26.45214, -69.33440, 2771m, 19.x.2015, L. Packer, Pans. All specimens are at PCYU , the holotype and allotype will eventually be deposited in MNHN .

DNA barcodes. Five full length DNA barcode sequences are available with the BIN AAO3541, they include both the holotype and the allotype. One of these was listed as Liphanthus n.sp. 3 in Packer and Ruz (2017) with Genbank accession KX820807 . The sequences differ by an average of 0.36% with a maximum divergence of 0.77%. They differ from their nearest neighbour, L. jenamro described above, by an average of 2.34%.

Comments. Using the key in Ruz and Toro (1983), if treated as having three submarginal cells, this species fails at the first couplet for such taxa, couplet 3 for males and 28 for females. Both couplets have the option of metatibial spurs curved and vertex concave or straight versus metatibial spurs straight and vertex convex. Like L.jenamro , L. sapos has the metatibial spurs curved but the vertex convex.

On the date of capture of the holotype, the type locality was the only area with flowering plants within a substantial distance, the vegetation being dense and quite lush because a small quebrada had been at least partially blocked as a result of road and bridge construction ( Fig. 30 ).

This species is part of the same species group as L. jenamro described above and L. domeykoi and L. discolor described below, all of which have males with a mandibular hair tuft at the end of a strongly lamellate outer ridge and minutely tuberculate frontal areas.

While the males are from low lying areas, the females are from higher altitudes. However, the DNA barcodes suggest they are conspecific and the distance between the localities is as small as 100km. Furthermore, whereas the most divergent individual is from lower elevations, one of the other low elevation individuals is less than 0.3% divergent from one of the high altitude ones. Consequently, we believe the sexes to have been associated correctly.

Figures

FIGURE 30. Type locality of L. sapos sp. nov., to the east of the Pan American highway, west of Los Sapos.

FIGURES 20–25. Liphanthus sapos sp. nov., male, 20: paratype lateral habitus, 21: holotype head frontal view, 22: paratype upper paraocular and frontal areas to show facial fovea (red arrow), 23: S7, 24: S8, 25: genital capsule. Scale bars Figs. 20–21 1mm, Fig. 22 50μm, Figs. 23–25 0.25mm.

FIGURES 128–132. 128: L. sapos metasoma oblique ventral view to show yellow sterna and ventrally reflexed portions of terga, 129: L. discolor metasoma in ventral view to show sterna suffused with brown and pale marks on reflexed portions only on more apical terga (arrows), 130: L. molavi—metasoma in lateral view to show dark metasomal sterna and lateral margins of terga, 131: L. jenamro mandible to show tuft of specialized hairs where lamellate outer ridge (red arrow) meets ventral margin, 132: L. molavi mandible to show unmodified outer ridge (red arrow) and absence of tuft of specialized hairs. Scale bars Figs.128–130 0.5mm, Figs. 131–132 0.25mm.

FIGURES 135–136. Liphanthus spp. head, frontal view: 135: L. jenamro to show dark marked base of clypeus and dark sub- antennal sclerite, 136: L. sapos to show lack of dark marking at base of clypeus and mostly pale subantennal sclerite. Scale bars 0.5mm.

FIGURES 137–138. Liphanthus sp. ventral view of metasoma: 137: L. discolor holotype to show brown suffused sterna and lack of pale marking laterally on T2 (arrow); 138: L. sapos to show lack of brown suffusion on metasomal sterna and yellow lateral tergal markings that extend also to T2 (arrow). Scale bars 0.5mm.

FIGURES 173–176. 173: L. jenamro metatibia to show narrowly yellow base and orange apex, 174: L. jenamro metatibial spur to show teeth longer than basal width, 175: L. sapos Metatibia to show broadly yellow base and yellow apex, 176: L. sapos metatibial spur to show teeth no longer than basal width. Scale bars Figs. 173 & 175 0.5mm, Figs. 174 & 176 50μm.

FIGURES 10–15. Liphanthus jenamro sp. nov., male. 10: male holotype lateral habitus, 11: male paratype head frontal view, 12: male holotype metasoma oblique ventral view to show colouration, 13: S7, 14: S8, 15: genital capsule. Scale bars Figs. 10–12 1mm, Figs. 13–15 0.25mm.

FIGURES 31–33. Liphanthus domeykoi sp. nov., holotype. 31: lateral habitus, 32: head frontal view, 33: posterior part of meso- and most of metasoma oblique ventral view to show yellow ventral surface of trochanters (red arrows) and concave and yellow metasomal sterna. Scale bars 1mm.

FIGURES 16–19. Liphanthus jenamro sp. nov., female allotype. 16: lateral habitus, 17: face frontal view, 18: apex of tibia to show curved metatibial spurs, 19: ESEM of metatibial spurs. Scale bars Figs. 16–17 1mm, Fig. 18 0.25mm, Fig. 19 100μm.