Phulia phantasma Lamas, Willmott & Boyer, 2022

Pyrcz, Tomasz W., Willmott, Keith R., Lamas, Gerardo, Boyer, Pierre, Florczyk, Klaudia, Fåhraeus, Christer, Mahecha, Oscar, Cerdeña, José, Mrozek, Artur, Farfán, Jackie & Zubek, Anna, 2022, Considerations on the Systematics of Neotropical Pierina, with the Description of Two New Species of Phulia Herrich-Schäffer from the Peruvian Andes (Lepidoptera: Pieridae, Pierinae, Pierini), Neotropical Entomology 51 (6), pp. 840-859 : 845-850

Phulia phantasma Lamas, Willmott & Boyer , n. sp.

[ Hypsochila n. sp. Lamas (Lamas, 2004: 115, no. 315)]

Types: HOLOTYPE (male): PERU, Ancash, Cerro Amancaes, cerca Santo Toribio , [08°50′S, 77°54′W], 3000 m a.s.l., 22.v.[19]80, G. Lamas, MUSM-ENT-006767 , [ MUSM] ; PARATYPES (37 ♂ and 1 ♀): Ancash: 2 males: same data as holotype, MUSM-ENT-006759 , 6763 , [ MUSM] ; 2 males: [Cerro] Amancaes , N de Huaylas, [8°50′S, 77°54′W], 2900 m a.s.l., 6.v.[19]79, V. Pacheco, MUSM-ENT-006762 , 6768 , [ MUSM] ; 1 male: same data, but 3000 m, 8.v.[19]79, MUSM-ENT-006758, [MUSM] ; 1 male: Oriente de la Cordillera Negra, C[omunidad] C[ampesina] Shecta , 9°28′06″S, 77°35′43″W, 4131 m, 05.ii.2012, B. Medina, [MUSM] ; 6 males: road from Bamba, T. Pyrcz [CEP-UJ] ; 1 male: same data [PBF] ; 4 males: same data, J. Cerdeña & J. Farfan leg., [MUSA] ; Cajamarca: 1 male: Cascas-Contumazá p[oste] k[ilometrique] 100, [7°24′35″S, 78°48′4″W], 2750 m, 23.iii.2022, P. Boyer, [PBF] ; 1 male: Cascas-Contumazá “borne kilometrique” 100, [7°24′66″S,78°48′076″W], 2650–2750 m, 19.vi.2018, [PBF] ; 1 male: same data, [MUSA] ; 6 males: Chilasque , 06°01′S, 79°12′W, 1200 m, 13.vi.2000, G. Lamas, MUSM-ENT-006760 , 6761 , 6764 , 6765 , 6770 , 6771 , [ MUSM]; 1 male: same data, except 12.vi.2000, R.K. Robbins, MUSM-ENT-006769, [MUSM]; 1 female: Llacanora , [7°11′S, 78°25′W], 2720 m , 19.xii.2006, [R. Vila], [MUSM]; 1 male: Limón-Santa Rosa , 1800–3000 m , i-ii.1998, R. Marx, FLMNH-MGCL-147145, [FLMNH]; 1 male: same data, FLMNH-MGCL-147146, [FLMNH]; 1 male: same data, FLMNH-MGCL-147147, [FLMNH]; 1 male: same data, FLMNH-MGCL-147148; [FLMNH]; 1 male: same data, FLMNH-MGCL-147149; [FLMNH]; 1 male: same data, FLMNH-MGCL-147150, [FLMNH]; 1 male: vía Celendín-Balsas , El Choloque , [6°51′57″S, 78°4′35″W], 1800–2000 m a.s.l., local collectors, vi-vii.2006, [ MABO]; Amazonas: 1 male: Chachapoyas, [06°10′S, 77°38′W], [2343 m], 1889, M. de Mathan, Ex Oberthür Coll. Brit. Mus. 1927–3., MUSM-ENT-006766 , [ MUSM]; 1 male: Molinopampa — Granada , [06°23′S, 77°26′W, 2800–3000 m a.s.l.], B. Calderón leg., [ CEP-UJ]; ECUADOR: Azuay: 1 male: Oña , [03°28′27″S, 79°9′32″W], 2200 m a.s.l., 22.iii.[19]65, L.E. Peña, [ MUSM] .

Diagnosis: Phulia phantasma n. sp. can be recognized from species of similar size previously placed in Hypsochila , Tatochila , or Theochila by its more compact appearance resulting from its less elongate wings, being similar in this respect to some species of Hesperocharis C. Felder, 1862 , such as H. marchalii (Guérin-Méneville, [1844]) , which is actually syntopic with P. phantasma n. sp. These two species of similar size and flight pattern can be easily confused in the field. Particularly pale individuals of P. phantasma with reduced black in the DFW apex and reduced (or almost absent) dark VHW markings are very similar to some individuals of P. maenacte from southeastern Brazil, but can be readily distinguished (apart from distribution) by having the base of vein R 4+5 on the FW much closer to the distal margin than it is to the base of vein M 1, whereas in P. maenacte it is approximately equidistant. Otherwise, P. phantasma n. sp. can be distinguished from P. maenacte by the dark scaling that lines the VHW veins broadening and fusing towards the distal margin, rather than tapering or remaining similar in width, and by the presence of a line of dark postdiscal spots in cells M 1 -Rs to CuA 2 -2A on the HWV, which may be shaped like distally pointing arrows, or just present as diffuse dark scaling, always absent in P. maenacte but similar to species previously placed in Hypsochila or several species of Tatochila . From the majority of other species of Phulia , P. phantasma n. sp. may be distinguished by the lack of any dark scaling at the end of the FW discal cell, with the dark markings in the distal half of the FW present as a simple black triangular patch which variably fills the apex and may be almost absent. Other superficially similar species (such as those formerly placed in Hypsochila , e.g., P. microdice or P. huemul (Peña, 1964)) , have at least the FW discocellular veins lined with black, and a variable line of black FW postdiscal spots.

Description: MALE ( Fig. 3A–B ): Head: eyes brown, bare, with narrow fringe of white scales at base and a yellow dorso-lateral spot; antennal shaft mixed black and white dorsally (24 antennomeres) with conspicuous rounded club (8 antennomeres) which is pale yellow except for proximal dorsal half which is black; labial palpi white with sparse long, black hair-like scales; top of head and frons white. Thorax: dorsal surface with black scales and long white hair-like scales from sides and near wing bases, ventral surface white, legs with sparse white scaling except for femur with denser white scaling and long white hair-like scales, mid- and hindlegs lacking tibial spurs (present in P. maenacte ; Field 1958). Wings: Forewing (length 27–28 mm, n = 5) triangular, hindwing an elongate oval with slightly straighter margin in middle of wing and angled tornus; FW with four radial veins, R 4 and R 5 fused, with veins R 4+5 and R 3 originating relatively close to the apex (R 4+5 approximately half length of R 3+4+5), M 2 originating independently of base of M 1 + R 3+4+5. FWD ground color white, variably present scattered blackish scaling filling apex with uneven basal edge indented in middle of each cell, scattered blackish scaling at very base of wing. HWD similar to forewing except lacking black apical scaling. FWV similar to dorsal surface except lacking dark scaling at wing base and dark apical scaling slightly paler, tinged yellowish, with very indistinct paler yellowish scaling forming intervenal stripes within darker apical area, costa with scattered pale grayish scales. HWV ground color pale yellow, slightly darker orange-yellow anterior of discal cell and more prominently along edge of costa, dark orange-yellow spot at base cell 2A-Cu 2; scattered dark grayish scaling lining edges of veins and a forming a “Y”-shaped marking in middle of discal cell, line of indistinct dark gray postdiscal markings in cells 2A-Cu 2 to Rs varying from distally pointing arrow shapes to indistinct spots, grayish scaling lining veins broadening and fusing towards margin. Abdomen: dorsal surface black, ventral surface white. Genitalia ( Fig. 6 ): similar in overall form to other members of the genus, for example Phulia wagenknechti (type species of Hypsochila ) as figured by Field (1958), notable features include long uncus (similar in length to tegumen), valvae with a blunt distal point, aedeagus of even width, and opening ventrally. In comparison with P. maenacte , there are several differences in the male genitalia, including in P. phantasma n. sp. a much longer uncus, greatly reduced in P. maenacte , and a tapering, downward-pointing aedeagus (flaring and upward-pointing in P. maenacte ). FEMALE ( Fig. 3C–D ): similar to male, except as follows: forewing (length 25 mm, n = 1) and hindwing more rounded, ground color slightly yellowish, dark DFW apical marking more extensive and with intruding pale scaling middle of each cell; ventrally similar to heavily patterned males, FW with trace of indistinct yellowish lines in middle of each cell in apical half, HW with stronger yellowish tinge to pale areas. Genitalia not examined.

Etymology: The name is a neuter Latin noun in the nominative singular meaning a ghost or phantom, in reference to the pale markings of this species and its mysterious rarity in collections.

Bionomics and distribution: This species is known from a large area of the central tropical Andes, extending from southern Ecuador to central Peru ( Fig. 13 ). It is rare in collections which does not reflect its status in the field. The perceived rarity results from this species occurring in the areas which are seldom visited by lepidopterists, dry Andean valleys, and especially being confused with common white pierids, especially with Hesperocharis marchalii and Leptophobia aripa (Boisduval, 1836) . Additionally, P. phantasma n. sp. is a very active, patrolling butterfly, only sporadically seen visiting flowers, such as of yellow and purple Onoseris albicans ( Asteraceae ) flowers ( Fig. 12E ), and not seeking humid areas where mud-puddling other pierids are frequently found. The flight is fast and swift. Although it has been recorded from a wide altitudinal range from 1200 to 4131 m, it seems that its optimal elevational range is between 2000 and 2800 m a.s.l. All individuals were collected between December and July, representing the wet season and first half of the dry season.

Molecular phylogeny

Our study provides strong support for the monophyly of the expanded concept ( Zhang et al. 2021) of Phulia in both the COI (bs: 94) ( Fig. 8 ) and the concatenated 3-genes tree (bs: 100) ( Fig. 9 ). The internal topology of this large clade differs, however, between analyses based on only COI or on all three genes.

In the COI tree, former Phulia cluster with former Piercolias , Pierphulia , and Infraphulia , as well as with former Hypsochila and three species formerly placed in Tatochila , P. orthodice ( Weymer & Maassen 1890) , P. mercedis , and P. theodice , and forms a sister clade to P. autodice . Phulia xanthodice (Lucas 1852) and Phulia sp. , in turn, form a sister clade to former Phulia and other species plus P. autodice , whereas P. homoeodice is situated on a long external branch relative to all previously mentioned species. Finally, Phulia maenacte and Phulia phantasma cluster together and are sister to all the other taxa. However, the COI tree is not fully resolved because the support values on deeper nodes are low. On the other hand, the support of terminal branches is high but they refer only to single, or exceptionally two species, simply meaning that the species identification based on external morphology fully agrees with barcode data.

In the concatenated tree, the former genera Phulia , Pierphulia , Piercolias , and Infraphulia cluster together in a weakly supported clade (bs: 41), sister to Phulia mercedis + Phulia wagenknechti , but branch support is low (bs: 48). Phulia orthodice is situated in an external position relative to all the abovementioned taxa, on a long branch, but again the common node support is low (bs: 36). Finally, Phulia maenacte and Phulia phantasma n. sp. form a weakly supported clade sister to all the other taxa. As in the COI tree, all the small internal clades composed of one or two species are strongly supported. In the concatenated tree, only a few species of former Tatochila + Hypsochila were included, which hampers understanding of the relationships of species formerly included in these genera in relation to other Pierina, and in particular the species of the Phulia sensu stricto clade. The only interesting noticeable difference is the position of Phulia orthodice as sister to the Phulia s. s. clade on the concatenated tree, and well-rooted in the Phulia sensu stricto clade in the COI tree. Otherwise, the topology of both trees is roughly similar, in particular the sister status of the Phulia phantasma n. sp. + Phulia maenacte clade in relation to the remaining genera.

Our results infer a divergence of the Phulia sensu lato clade from Ganyra josephina in the middle Miocene (Tortonian), ~ 8.8 Mya (4.97–11.8 HPD 95%) ( Fig. 10 ), followed by the subsequent divergence of the two main clades of the Phulia group in the late Pliocene (Piacenzian), ~ 3.3 Mya (2.4–4.2 HPD 95%). The radiation of Phulia sensu lato occurred throughout the Pleistocene, with the divergence of P. maenacte and P. phantasma n. sp. at ~ 2.5 Mya (1.72–3.03 HPD 95%), and that of other clades of Phulia sensu lato slightly later, at ~ 2.1 Mya (1.55–2.51 HPD 95%). The two clades which form the Phulia sensu stricto group originated at ~ 1.8 Mya (1.34–2.07 HPD 95%). In addition, Phulia garleppi diverged from the rest of Phulia sensu lato at ~ 1.9 Mya (1.46–2.18 HPD 95%). The clades ( Phulia stoddardi n. sp. + Phulia (“ Pierphulia ”) sp.) + two species previously associated with Piercolias ( P. cf. forsteri + P. coropunae )) diverged at ~ 1.6 Mya (0.52–1.88 HPD 95%). The group comprising two species previously associated with Pierphulia ( P. rosea + P. nysias ( Weymer & Maassen, 1890)) + Phulia nymphula ) separated at~ 1.4 Mya (0.39–1.62 HPD 95%). The group comprising the species previously placed in Infraphulia ( P. ilyodes )+ Phulia sensu stricto ( P. nannophyes Dyar, 1913 + P. paranympha Staudinger, 1894 )) at ~ 1.2 Mya (0.36–1.51 HPD 95%), and the clades (( P. rosea + P. nysias ) + P. nymphula ) + ( P. ilyodes +( P. nannophyes + P. paranympha )) diverged at ~ 1.5 Mya (1.1–1.83 HPD 95%) ( Fig. 10 ).

Figures

Fig. 3 Adults: A Phulia phan- tasma male holotype, dorsal (Cerro Amancaes); B idem, ventral; C P. phantasma female paratype, dorsal (Llacanora); D idem, ventral; E Phulia (Theochila) maenacte maen- acte male (Sao Francisco de Paula), dorsal; F idem, ventral; G Phulia (Theochila) maenacte itatiayae male (Itatiaya), dorsal; H idem, ventral

Fig. 6 Male genitalia of Phulia phantasma n. sp.: A lateral view; B vertical view; C aedea- gus lateral view; D aedeagus vertical view

Fig.13 Distribution of Phulia phantasma n. sp. and Phulia stoddardi n. sp

Fig.12 Habitat of Phulia phan- tasma n. sp.: A. A cliff above the Contumazá – Cascas road B. Asteraceae plants along the road; C view on the valley of Cascas; D xerophytic vegetation along the Contumazá–Cascas road, 2700 m; E flowers of Onoseris albicans (Asteraceae) frequently visited by numerous species of pierids, including P. phantasma n. sp.; F patches of cloud forest below the Con- tumazá–Cascas road, 2700 m (photos: P. Boyer)

Fig. 8 COI ML tree of Phulia, bootstrap values above 60 indicated at ▸ nodes. All the generic names used previously are shown in order to underline their relative position in the phylogeny. The genera Ascia and Ganyra are used as outgroups

Fig. 9 Concatenated (COI, GAPDH, RpS-5) ML tree of Phulia, bootstrap values above 60 on the nodes. All the generic names used before are shown in order to underline their relative position in the phylogeny. The genera Ascia and Ganyra are used as outgroups

Fig. 10 A hypothesis of divergence timing of the genus Phulia (generic names used previously in parenthesis)