DHJ01 (P. argante of Brown, 1929)
2. An ongoing illustrative case study: Phoebis argante (Pieridae). As soon as barcoding was applied to the inventory, it became apparent that it exposes overlooked species (e.g. Hebert et al. 2004; Smith et al. 2006-2008, Burns et al. 2008). The large yellow pierid butterfly Phoebis argante (Fig. 2) is an ongoing example. P. argante is a well-known butterfly throughout the neotropics (DeVries 1987). The ACG inventory has reared it more than 350 times from caterpillars found on Zygia longifolia and six species of Inga (Fabaceae) in ACG dry forest, rain forest and intergrades. When barcoded, however, two very distinct lumps of barcodes appeared in the NJ tree of ACG Pieridae, each being so far apart that a similar species (Phoebis virgo) falls between them (Appendix SV). The two barcode lumps, P. argante DHJ01 and P. argante DHJ02, do not correlate with food plant, season or ecosystem. The caterpillars of both have been collected from the same plant at the same time, although the caterpillar of P. argante DHJ01 is about three times more commonly found than is that of P. argante DHJ02. The genitalia of the two species appear to be the same at the level of scrutiny normally accorded to pierid genitalia (and were first decreed by an experienced taxonomist to be "the same") but on closer study display slight but consistent differences (Fig. 2). Likewise, when sorted by barcode, the fine details of the wing colour pattern of both sexes became evident as a 100% reliable method for distinguishing these two species that are contained within ACG "P. argante" [e.g. note the difference in the pattern of black on the outer margin of the male forewing (Fig. 2)]. Prior to barcoding, these segregating wing traits had been viewed by the inventory and experienced taxonomists as variation not indicating a species-level dichotomy, and indeed both cryptic species are somewhat variable in these traits. UV light-visible wing reflectance patterns, long studied in Pieridae and interspecifically discriminatory (Silberglied & Taylor 1973; Allyn & Downey 1977; Rutowski & Macedonia 2008) are not different between the males of the two species of P. argante but do differ on the undersides of the females in ACG. Both species occur throughout Costa Rica, as based on morphological inspection of 66 net-caught adults in the INBio national inventory (IC). However, in contrast to ACG inventory rearing records, P. argante DHJ01 is the low-density species among these net-collected free-flying adults. Which, if either, of the two species of P. argante in ACG is actually P. argante? P. argante was described in 1775 from a Brazilian specimen and several so-called subspecies of P. argante have been described from Caribbean Islands, Peru, "America", and Mexico (Lamas 2004). All of these are candidates to match one, both, or none of the species in ACG and Costa Rica. However, a direct quote from a 1929 analysis of Phoebis argante Neotropical taxonomy (Brown 1929: 12) is relevant: "Two forms of the mainland race occur: the nymotypical argante and Cramer's hersilia. In studying a series of about two hundred specimens from over the entire range of this insect, it became apparent at once that hersilia is the dominant tropical form and argante the dominant form in the north and south. Form hersilia is gradually separating itself from the argante type and forming a purely tropical race that will be flanked on the north and south by forms similar to our present argante. In this case the difference in forms is apparent in the males - the marginal row of black dots having become confluent and having formed a black band in hersilia. However, only about forty per cent of the specimens from the northern part of South America show a complete band, but only one to two per cent show no trace of this band. There is a transitional group of about seventy per cent of all the males in the collections I have examined. The form hersilia ranges from Honduras to Bolivia - almost the entire range of the mainland race." If Brown had had our ACG barcode results, there is little doubt that he would have concluded that P. argante DHJ01 is P. argante and elevated P. argante DHJ02 to P. hersilia. However, when the holotypes behind these two names are examined in detail with a full barcode analysis of "P. argante" over all of its range, yet additional barcode groups may be found that correlate with morphological, geographical, ecological and/or behavioural traits, suggesting that P. argante is yet more than two species on the mainland.
Distribution and Larval Foodplants:
KS, TX (rare stray), E & W Mexico to Argentina, Trinidad
(From: A Catalogue of the Butterflies of the United States and Canada, Jonathan P. Pelham)
Genus Phoebis Hübner, 
Verz. bekannt. Schmett. (7): 98. Type-species: Phoebis cypris Hübner, , Verz. bekannt. Schmett. (7): 98, no. 1044 (and unjustified emendation of Papilio cipris Cramer, 1777, Uitl. Kapellen 2(9): 5, pl. 99, figs. E, F; (16): 148 (index); = Papilio argante Fabricius, 1775, Syst. Entomol.: 470, no. 116; synonym), by designation of Butler (1873), Lepid. Exot. (13): 155.
= ‡Colias Hübner, 
Verz. bekannt. Schmett. (7): 99. Type-species: Papilio sennae Linnaeus, 1758, Syst. Nat. (ed. 10) 1: 470, no. 72, by designation of Hemming (1939), Proc. R. Entomol. Soc. Lond. (B)8(7): 136. Preoccupied by Colias [Fabricius], 1807, in: Illiger, Mag. f. Insektenk. 6: 284.
= Callidryas Boisduval & Le Conte, 
Hist. Lépid. Amér. sept. (8): 73. Type-species: Papilio eubule Linnaeus, 1767, Syst. Nat. (ed. 12) 1(2): 764, no. 102 (= Papilio sennae Linnaeus, 1758, Syst. Nat. (ed. 10) 1: 470, no. 72; synonym), by designation of Blanchard (1840), in: Castelnau, Hist. Nat. Ins. 3: 432.
= ‡Metura Butler, 1873
Lepid. Exot. (13): 155. Type-species: Callidryas bracteolata Butler, 1865, Proc. Zool. Soc. Lond. 1865(2): 458, pl. 26, fig. 6 (= Colias neocypris Hübner, , Samml. exot. Schmett. 2: pl. , figs. 1, 2; synonym), by designation of Scudder (1875), Proc. Am. Acad. Arts Sci., Boston 10(2): 218, no. 695. Preoccupied by Metura Walker, 1855, in: Allen's Nat. Libr., Lepid. 2: 229.
= Parura W. F. Kirby, 1896
In: Allen, Nat. Libr. Lepid. 2: 229. Type-species: Callidryas bracteolata Butler, 1865, Proc. Zool. Soc. Lond. 1865(2): 458, pl. 26, fig. 6 (= Colias neocypris Hübner, , Samml. exot. Schmett. 2: pl. , figs. 1, 2; synonym), as a replacement name. Proposed to replace Metura Butler, 1873, preoccupied (Code Articles 60.3, 67.8).
Phoebis argante (Fabricius, 1775)
Phoebis argante argante (Fabricius, 1775)
Syst. Entomol.: 470, no. 116.
Original Combination: P[apilio]. D[anaus]. C[andidus]. Argante
Type Locality: “Brasilia” Fabricius cited “Mus. Banks” and L. Miller and F. Brown (1981), Mem. Lepid. Soc. (2): 191, state that only Rio de Janeiro was visited by Banks.
Types: Syntype in BMNH.
= cipris (Cramer, 1777)
Uitl. Kapellen 2(9): 5, pl. 99, figs. E ♀ D, F ♀ V; (16): 148 (index).
Original Combination: Pap[ilio]. Dan[aus]. Candidi Cipris
Type Locality: “Surinamen” “Suriname”
Types: Syntypes (#665064, 665065) in BMNH.
= volcanica (Perry, 1811)
Arcana 1: , pl. .
Original Combination: Papilio Volcanica
Type Locality: “Rio de la Plata” [Argentina]
Types: Types probably lost.
= ‡cnidia (Godart, 1819)
In: Ency. méth. 9(Ins.)(1): (86), 93-94, no. 14; cited Papilio cipris Cramer, 1777, Uitl. Kapellen 2(9): pl. 99, figs. E, F: (16): 148 (index), and Herbst (1792), in: Jablonsky, Natursyst. Ins., Schmett. 5: pl. 111, figs. 3, 4.
Original Combination: Colias Cnidia
Proposed to replace Papilio cipris Cramer, 1777, erroneously thought preoccupied.
= ‡cypris (Hübner, )
Verz. bekannt. Schmett. (7): 98, no. 1044; cited Papilio cipris Cramer, 1777, Uitl. Kapellen 2(9): pl. 99, figs. E, F; (16): 148 (index).
Original Combination: P[hoebis]. Cypris
An unjustified emendation of Papilio cipris.
= aurantia Larrañaga, 1923
Escrit. Larrañaga. publica Instit. Hist. Geogr. Uruguay: 418.
Original Combination: Papilio aurantia
Type Locality: “frequens in floribus civitatis” [frequent at flowers in the city of Montevideo, Uruguay]
Types: Type(s) lost.
= ‡”albante” F. Brown, 1929
Am. Mus. Novit. (368): 13.
Original Combination: Phoebis argante argante female form albante
Type Locality: “Tegucigalpa, Honduras”
Types: “Holotype” in AMNH.
‡”alba” Scott, 1986
Butts. N. Am.: 205, no. 52.
Original Combination: [Phoebis argante ♀] form alba
Proposed as a “form” with no nomenclatural standing; a nomen nudum.