Liphanthus jenamro 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 : 7-12

Liphanthus jenamro Mir Sharifi & Packer , sp. nov.

urn:lsid:zoobank.org:act:276F8E65-769A-4429-B273-5B1BCE27DF9B

Figs. 1–4 , 10–19 , 124–125 , 131 , 134 , 135 , 163 , 169–170 , 173–174 .

Diagnosis. Males of this species can be differentiated from all others of the genus except L. sapos Mir Sharifi & Packer by the combination of two submarginal cells ( Fig. 10 ), metatibial spurs strongly curved at apices (as in Figs. 18–19 ) and metasomal sterna and reflexed portions of most metasomal terga yellow ( Fig. 12 ). Other species with two submarginal cells, except L. sapos , lack the combination of metasomal sterna yellow and metatibial spurs strongly curved apically. It can be differentiated from L. sapos by the subantennal sclerite black ( Fig. 11 ) except sometimes the extreme apex yellow whereas in L. sapos the lower 4/5 of the subantennal sclerite is yellow ( Fig. 21 ). If L. domeykoi Packer described below, is also considered to have strongly curved metatibial spurs (the sole specimen has them moderately curved), then L. jenamro can be differentiated from it by the almost entirely black-brown trochanters ( Figs. 10 & 12 ) which are yellow to yellow-brown ventrally in L. domeykoi ( Figs. 31 & 33 ). Females of L. jenamro can be differentiated from all other species known, except L. sapos , by the combination of two submarginal cells, metatibial spurs strongly curved at apices ( Figs. 18–19 ) and pronotal lobe yellow ( Fig. 16 ). Other species with two submarginal cells, except L. sapos , have a dark pronotal lobe or metatibial spurs not strongly curved apically. Females can be differentiated from those of L. sapos by metatibia black except yellow at the base and orange at the apex and inner metatibial spurs with narrow teeth, half as wide as long ( Figs. 18–19 ). For L. sapos females, the metatibia is yellow for the basal half dorsally and the inner metatibial spurs have broad teeth, as wide as long (Figs. 28–29).

Description: Holotype Male. Dimensions: Approximate body length: 3.92 mm; head width: 1.04mm, wing length: 2.63mm, intertegular width: 0.74mm.

Coloration: Black to dark brown with following parts yellow: labrum, mandible (except apex red and base suffused with brown), lower paraocular area below up to level of lower 1/3 of outer subantennal suture, clypeus (except upper 1/3 black), ventral surface of F4–F11 (F1–F3 yellow suffused with brown, rest of flagellum red-brown), pronotal lobe, apical ring on all femora, protibia dorsally, meso- and metatibia (except for brown dorsally incomplete ring around midlength), all basitarsi, metatibial spurs, metasomal sterna, ventrally reflexed and immediately adjacent nonreflexed portions of metasomal terga. Apical impressed areas of T1–T2 light brown, of T3–T5 yellow-brown.

Sculpture: Black portion of face densely and minutely tuberculate, dull (except lower ½ of subantennal sclerite weakly imbricate, shiny); yellow portion weakly imbricate, shiny; clypeus punctures large, weak, dense, i<d, those towards sides longitudinally effaced; moderately dense and rounded elsewhere, i=0.5–2d; yellow portion of lower paraocular area punctures smaller, sparse, i=1–3d; rest of lower paraocular and supraclypeal areas and subantennal sclerite punctures weak, irregularly spaced; upper paraocular, frontal and ocellocular areas, densely rugoso-punctate; ocellocular and vertexal areas punctures small, scattered and obscure; genal and hypostomal areas weakly imbricate, shiny; punctures weak, sparse i>d. Mesosoma strongly imbricate, dull; mesoscutum and scutellum with weak, moderately sparse punctures, i>d; metanotum more coarsely punctate; hypoepimeral area, rest of mesopleuron and metapleuron imbricate, dull; punctures obscure and sparse, i>2d except absent on hypoepimeral area; metapostnotum dorsal surface rugulose, posterior and lateral surfaces of propodeum imbricate, somewhat shiny. Metasomal terga increasingly weakly imbricate from T2–T6, T2 anteriorly dull, rest and remaining terga somewhat shiny; T1-T4 minutely sparsely punctate, T5 and T6 punctures shallow, moderately dense, i=1–1.5d; apical impressed areas imbricate, impunctate; metasomal sterna weakly imbricate, shiny; punctures small i=1–2d.

Pubescence: Generally white, sparse and simple or with minute branches, ≤1.5 MOD, longer, ≤2.2 MOD on ventral surface of mesopleuron, posterior half of scutellum and metanotum. Dense, almost squamose ~0.5 MOD on dorsal surface of metapostnotum. Metasomal terga hairs short, 0.5 MOD, laterally oriented; hairs sparse and more erect on sides of metasoma.

Structure. Head: approximately as long as wide (92:90). Mandible 2 X as long as basal depth (38:19); gradually narrowing towards acute apex; outer ridge lamellate, lower margin at midlength with erect pencil of branched hairs where outer ridge meets ventral margin. Labrum rectangular, more than 1.5 X as broad as long (31:18). Clypeus 1.7 X as wide as long (51:32), apicolateral and apicomedial margins weakly convex. Outer subantennal suture outwardly convex, inner suture outwardly weakly concave, subantennal sclerite widest near midlength; epistomal suture ventrally concave between inner subantennal sutures. Anterior tentorial pit just below junction of outer subantennal and epistomal sutures. Frontal line narrowly depressed below and above, imperceptible around midlength. IAD> AOD (14:12). Inner margin of compound eyes convergent below, UOD:LOD 61:53. Facial fovea small, obscure, length ~0.5 MOD, comma-shaped, broadest below middle. IOC: OOC 20:18. Scape 1.6 X as long as greatest width (16:10), subequal to pedicel and F1 combined (17); pedicel shorter than wide (7:9), F1 as long as wide (10:10). F2 length subequal to width (6:7), remaining flagellomeres slightly longer than wide except F11 ~1.9 X as long as wide (21:11).

Mesosoma: Mesoscutum 1.25 X as wide as long (65:52), length of scutellum: metanotum: metapostnotum: 25:12:15. Marginal cell subequal in length to the distance between its apex and wing tip (60:58). 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 T2. T7 lacking pygidial plate. S1–S5 unmodified, S6 apicomedially translucent and weakly concave, premarginal line bearing few setae either side of concavity. S7. Apodeme moderately long and broad, anterior margin obtuse, weakly sinuate; apical lobe short, length and width subequal, broadly rounded, with sparse long-branched posteriorly oriented hairs almost as long as width of lobe. S8. Apodeme anterior margin sinuate, 1.5 X wider than long, lateral margins strongly diverging anteriorly; lateral lobe small, apex acute; apical lobe broadly rounded, tongue-shaped, lateral margins convex, short spicules on apical half of dorsal surface. Gonocoxa anterodorsal margin transverse, lateral margins diverging posteriorly, medial margin almost straight; gonostylus distinctly separated from gonocoxa by weakly sclerotized band, narrow, gradually narrowing to apex, posteromedially weakly curved, short hairs on medial and ventral surfaces; penis valve broad, laterally broadly convex with weak concavity near midlength; endophallus basal sclerotized fluting long and narrow, apical membranous portion extending close to apex of penis valve and surpassing the valve laterally ( Fig. 4 ).

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

Dimensions: Approximate body length: 4.56mm; head width: 1.16mm, wing length: 3.04mm, intertegular width: 0.91mm.

Coloration: Black to dark brown with following parts yellow: mandible (except apical half yellow-brown to redbrown), ventral surface of F3–F10 (rest of flagellum red-brown), pronotal lobe, apical ring on profemur, dorsal surface of protibia except apical 1/3, basal ring of mesotibia. Following parts orange to orange-brown: labrum, extreme apex of clypeus, all tibiae apicodorsally, basal ¼ of metatibia, metabasitarsus basally and ventrally, pro- and metabasitarsi, metasomal tergal margins between disc and apical impressed areas, metasomal sterna, darker towards apices of S3–S5.

Sculpture: Face below from midlength of subantennal sclerite weakly imbricate, shiny, rest of face acinose, dull; clypeus and lower paraocular areas punctures large, dense, i≤d; lower paraocular, subantennal and supra- clypeal areas punctures distinct, variable in size and spacing. Metasomal sterna impunctate anteriorly, posteriorly punctures increasing in size from S2–S5, S6 with few small subapical punctures.

Pubescence: Longest on ventral surface of mandible and tibial scopa ~2.5 MOD; apical margin of clypeus medially ~2 MOD; subapical fringes of S2–S5 ~1 MOD. T6 and prepygidial fimbria hairs ranched, brown <1.5 MOD.

Structure. Head: length subequal to width (73:72). Mandible ~ 3 X as long as basal depth (64:21), gradually narrowing towards rounded apex. Labrum rectangular, <1.5 X as wide as long (27: 20), raised portion U-shaped. Clypeus <~2.3 X as wide as long (68:29). IAD= AOD (16:16). Inner margin of compound eyes subparallel, UOD: LOD 70:71. Facial fovea sinuate, length to width: 20:4, outer margin parallel to inner margin of compound eye. IOC: OOC 21:18. Scape 2.5 X as long as greatest width (28:11), ~1.5 X as long as pedicel and F1 combined (17); pedicel as long as wide, F1 longer than wide (11:9); F2 shorter than wide (6:9), F3 (7:9), remaining flagellomeres slightly longer than wide except F10 1.6 X as long as wide (16:10).

Mesosoma: Length of scutellum: metanotum: metapostnotum: 28:15:23. Tarsal claw teeth short.

Metasoma: Metasomal terga and sterna unmodified. T2 fovea 3X longer than greatest breadth (30:10) which is subapical. Pygidial plate triangular, sides weakly convex, forming an angle of ~50˚, apex rounded, plate narrowly raised medially.

Material examined. Holotype male, allotype female, 14 male and 101 female paratypes as follows: holotype male, allotype female, 6 male and 28 female paratypes: CHILE, Region II, ~ 13 km S. of Paposo , {S25˚ 09.67 W 70˚26.54}, 28–29.ix.2002, J. Grixti & A. Zayed, pan trap; 5 females same data except 28.x.2002 and with gps co- ordinates included on label and four of them ex Nolana ; 3 males and 22 females, same data except 29.ix–4.x.2002. Three female paratypes as follows: CHILE, Region II, 13 km S. of Paposo, 4.x.2002, J. Grixti & A. Zayed. Nine females, two males, same data but 28.ix.2002, six of the females from Nolana sp., two males and two females from Oxalis bulbocastanum Phil. ; 11 females and two males, same data but 4.x.2002, one male and two females from O. bulbocastanum , one male, two females from Nolana paradoxa Lindl. , seven females, Nolana sp.; 22 females same locality except 4.x.2002, with previous gps coordinates on label, 17 ex Nolana , 5 ex O. bulbocastaneum ; one female, CHILE, Region II, 5km N. of Paposo , 25.x.2000, L. Packer. One male paratype: CHILE, Region II, N of Tal-tal, Blue Pans, 20.xi. {20}02, J. Grixti & A. Zayed. All specimens are at PCYU except for single male and mul- tiple female paratypes at each of the following institutions: AMNH, MNHN, PUCV and female paratypes at BBSL, CTMI and SMNH. The holotype and allotype will eventually be deposited at MNHN.

Etymology. This species is named after the collectors, Jennifer Grixti and Amro Zayed and the specific epithet should be considered as a noun in apposition.

Variation. For males, apex of mandible reddish to pale yellow; subantennal sclerite varies from entirely black to black with small yellow spot apically; clypeus varies from entirely yellow to apical 2/3 yellow; anterior spot on tegula ranges from yellow to yellow-brown; subapical bands on metasomal terga variable in color (pale yellow to orange).

For females, metasomal terga apical impressed areas and metasomal sterna range from pale yellowish to brown.

DNA barcodes. Two full length DNA barcode sequences are available, one of them from the allotype, with the BIN AAV8057 and listed as Liphanthus n.sp. 2 in Packer and Ruz, 2017, Genbank accessions KX820350 and KX820641 . The two sequences differ by 0.31% from each other and 2.6% from their nearest neighbour, L. sapos described below.

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. 28 for females. Both couplets have the option of metatibial spurs curved and vertex concave or straight versus metatibial spurs straight and vertex convex. Liphanthus jenamro has the metatibial spurs curved but the vertex convex.

Different numbers of individuals were collected in differently coloured pan traps despite there being an equal number of each colour placed at the type locality (Grixti personal communication). The data are as follows: nine males and 40 females (including allotype) in yellow pan traps, two males (including holotype) and eight females in blue pan traps, three male and seven females in white pan traps.

There is a number of undescribed species of Liphanthus in which the males have a lamellate outer ridge to the mandible and where this ridge meets the ventral surface of the mandible there are hairs, arranged in a tuft, which have long branches on one side of the rachis ( L. incasicus and L. quadrifasciatus also have these two characteristics) and most have yellow markings on the metasoma. Except for the four species described herein ( L. jenamro and the three following new species), these have three submarginal cells. These species with three submarginal cells are not conspecific with the species being described here as they all differ in numerous characteristics such as colour pattern, head shape and sculpture (Packer unpublished data). One of these undescribed species with three submarginal cells is from the same political region as L. jenamro but from at least 2500m in altitude in the Andes, whereas all specimens of L. jenamro are from just above sea level along the coast.

Most males of most of the species with the aforementioned mandibular features have angularly reflexed sides to the metasomal terga and transversely concave sterna. Most of these specimens come from traps containing propylene glycol and it is possible that the method of temporary preservation caused this concavity. However, hundreds of other congeneric specimens have been collected using this method with very few of them having a ventrally concave metasoma whereas this concavity is almost universal among L. jenamro and the species related to it, whether they have two, or three, submarginal cells.

This species has not been found since 2002 despite extensive pan trapping in areas that encompass the type locality and time of year.

Floral hosts are known for this species: Nolana paradoxa Lindl. , Solanaceae and Oxalis bulbocastanum Phil. Oxalidaceae .

Figures

FIGURES 1–3. Unusual morphological features of some Liphanthus species. 1: L. jenamro sp. nov. ESEM of frontal area of holotype male to show minutely and densely tuberculate microsculpture, 2: L. jenamro ESEM of frontal area of allotype female to show acinose (bracket) to minutely and densely tuberculate microsculpture, 3: L. jenamro mandibles of a paratype male to show tuft of specialized hairs at the junction of the outer ridge and ventral margin, note the long branches on one side of the rachis only. Scale bars Figs. 1–2 100μm, Fig. 3 0.25mm.

FIGURES 4–9. Apicomedial portion of genital capsule ventral view (unless otherwise stated) to show elaboration of the endophallus, 4: L. jenamro sp. nov. holotype, apex of extension indicated by red arrow, 5: L. molavi sp. nov. paratype to show broadly triangular sclerotized (lateral apex indicated by red arrow) portion of endophallus, 6: L.amblayensis sp. nov. holotype to show absence of elaboration; 7–9: L. centralis sp. nov. paratypes, 7: to show very complex structure of cleared structures as seen with light microscopy in ventral view, 8: detail of genitalia to show structure of apical portion of penis valve in uncleared material, oblique view. 9: ESEM to show details of apical portion of penis/penis valve oblique view, showing minutely tuberculate/papillate microstructure, red arrow points to the apex of the penis valve. Scale bar Figs. 4–8 0.125mm, Fig. 9 100 μm.

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 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.

FIGURES 124–127. 124: Liphanthus jenamro to show pale pronotal lobe, 125: L. jenamro head, frontal view to show pale lower paraocular area, 126: L. amblayensis to show dark pronotal lobe, 127: L. ancashensis head, frontal view to show dark lower paraocular area. Scale bars 0.5mm.

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 133–134. Liphanthus spp. mesosomal venters, oblique ventral view: 133 L. domeykoi holotype to show pale ventral surface of all trochanters (arrows), 134: L. jenamro to show dark ventral surface of all trochanters (arrows). Scale bars 0.5mm.

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 163–164. Liphanthus sp. Side of mesosoma to show colour of pronotal lobe. 163: L. jenamro with pale pronotal lobe, 164: L. amblayensis with dark pronotal lobe. Scale bars 0.5mm.

FIGURES 169–172. 169: L. sapos metatibial spurs to show strong curvature; 170: L. jenamro to show IOD (red line)> OOD (blue line), 171: L. centralis metatibial spurs to show weak curvature, 172: L. centralis to show IOD (red line) <OOD (blue line). Scale bars Fig. 169 100μm, Figs. 170 & 172 0.5mm, Fig. 171 50μm.

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 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 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.