Generic placement of the Neotropical species of “ Phragmatobia” (Erebidae, Arctiinae), with a remarkable matrivorous species from the Peruvian Andes

Abstract Phragmatobia Stephens is briefly reviewed and a diagnosis is provided. The South American species currently placed in Phragmatobia Stephens are revised to two new genera, Andesobia Schmidt and De Freina, gen. n., and Patagobia Schmidt and De Freina, gen. n. (subtribe Spilosomina). Both Andesobia and Patagobia exhibit adaptations to high altitude habitats, including micropterous females in Andesobia (Patagobia females are unknown) and diurnal flight of males. The adults, immature stages, and mating behaviour of Andesobia jelskii (Oberthür, 1881) are described. Males of Andesobia jelskii enter the female cocoon to mate, and the micropterous, flightless females remain in the cocoon following oviposition where newly hatched larvae feed initially on the female’s body. Four species are included in Andesobia, Andesobia jelskii comb. n. (= Paracles imitatrix Rothschild, 1922, syn. n.), Andesobia flavata (Hampson, 1901), comb. n., Andesobia boliviana (Gaede, 1923), comb. n. (=Turuptiana flavescens Rothschild, 1933, syn. n.), and Andesobia sanguinea (Hampson, 1907), comb. n. Patagobia includes only Patagobia thursbyi (Rothschild, 1910), comb. n., and Patagobia thursbyi pluto Toulgoët is relegated to its synonymy. Patagobia shows affinities to Phaos Walker, 1855 of Australia, Metacrias Meyrick, 1886 of New Zealand, and Pseudophragmatobia Krüger, 2009 of South Africa, suggesting a common ancestry of circumantarctic origin. Phragmatobia karsholti Toulgoët, 1991 is transferred to Venedictoffia Toulgoët, comb. n., an unrelated genus that is removed from subtribe Arctiina and provisionally placed in the Phaegopterina. Phragmatobia oberthueri Rothschild, 1910, described from Tibet, is a synonym of Lachana alpherakii (Grum-Grzhimailo, 1891) [Erebidae: Lymantriinae], syn. n., comb. n.

Description of the immature stages and life history was based on four successive generations of laboratory rearings in 2009 and 2010 by JJD, obtained from live material from Peru, Junin region, Huicuash E of Tarma, 11°23'S, 75°53'W, 4100 m. All rearings were conducted indoors at ambient temperatures between 10° C and 23° C. Mortality was extremely low up to F3, but F4 larvae had higher mortality rates and females exhibited reduced fertility, both presumably symptoms of inbreeding. To obtain matings, newly emerged females were placed individually in wooden boxes screened at the top to allow air circulation. Copulation was achieved only under sunny conditions, probably because males are active only during warm, sunny periods under natural conditions. Larvae accepted both dandelion foliage (Taraxacum officinale L.) and grass (Poa sp.), with a preference for the latter.
We used the 658 bp 'barcode' region of the first subunit of the cytochrome oxidase (cox1) gene (Ratnasingham and Hebert 2007) of Andesobia jelskii to compare to Phragmatobia species and other New World genera of Spilosomina. DNA was extracted from one leg removed from a dried specimen, sent to the University of Guelph in dry Eppendorf tubes, and processed as part of the "All Leps Barcode of Life Campaign" (www.lepbarcoding.org). DNA extraction, amplification and sequencing protocols for the Barcode of Life (BOLD) initiative are detailed in Hebert et al. (2003). Haplotypes of all cox1 'barcode' fragments were compared with phylograms constructed using the neighbour-joining method as implemented on the BOLD website. Type locality. [Europe].
Phragmatobia includes five species distributed in the Palaearctic and Nearctic regions (including one Holarctic species, P. fuliginosa (L.)), with the Neotropical species and one Asian species here transferred to other genera. As suggested by Forbes (1960), Phragmatobia is probably most closely related to the Nearctic genus Pyrrharctia Packard; male genitalic and molecular characters strongly support these two as sister taxa (Schmidt 2007). We examined all Phragmatobia species, i.e. Phragmatobia fuliginosa (Linnaeus, 1758), Phragmatobia amurensis Seitz, 1910, Phragmatobia placida (Frivaldszky, 1835, Phragmatobia lineata Newman & Donahue, 1966 and Phragmatobia assimilans Walker, 1855. Examination of the type material (BMNH) of Phragmatobia oberthueri Rothschild, 1910, described from Kuku-Noor, Tibet, revealed that it is a junior synonym of Lachana alpherakii (Grum-Grzhimailo, 1891) [Erebidae: Lymantriinae], syn. n., comb. n., a group recently revised by Trofimova (2008). Two other species sometimes placed in Phragmatobia in the recent literature have been transferred to other genera, namely Orontobia coelestina Püngeler, 1904(De Freina 1997 and Epatolmis luctifera ([Denis & Schiffermüller], 1775) (Kôda 1988). Watson and Goodger (1986) placed eight Neotropical species in Phragmatobia, three of which were transferred to other genera by Ferguson (1985). Phragmatobia modesta Maassen, 1890 was recently moved to Amastus Walker [Arctiini: Phaegopterina] by Vincent and Laguerre (2010), leaving four Neotropical species that are dealt with here. Diagnosis. Phragmatobia is a fairly homogeneous group characterized by the following combination of characters: male antennae simple; wings fully developed in both sexes, forewing transverse lines diffuse or absent; wing colours varying from pinkish red to dark vinaceous red with darkbrown to blackish markings. Microtymbal of metepisternum well developed (P. fuliginosa) to obsolete (P. assimilans). Male genitalia with apical process of valve finger-like and ovoid in cross section; clasper spade-like, oriented transverse to longitudinal axis of valve, originating from inner surface of valve and directed mesad (divided into a ventral and costal lobe in P. fuliginosa); apex of aedeagus with spinose plates; paired, intersegmental coremata present between sternites 7-8. Females with ductus bursae heavily sclerotized, dorso-ventrally flattened, and nearly as wide as width of abdomen; corpus bursae globose, with two signa consisting of small flattened spicules; dorsal pheromone gland paired, each duct with 3-4 branches, the apices of which are rounded. Etymology. The name is feminine in gender, formed by combining the words Andes and -obia from the generic name Phragmatobia.

Diagnosis.
Andesobia is related to Patagobia, but is distinguished by the following combination of characters: eyes reduced and ellipsoid, 1.4-1.6 × as high as wide, gena with broader unscaled area laterally; posterior antennal rami 1.2-1.5 × and anterior rami 1.1-1.5 × longer than segment length (longest anterior and posterior rami 3 × as long as segment in Patagobia); 2 nd labial segment short and stout, 1.1 × as long as wide, 2 × longer than apical segment; thoracic collar concolourous with dorsal thoracic vestiture (contrastingly paler ochre in Patagobia); thoracic vestiture sparse and shaggy, compared to dense and pilose vestiture in Patagobia; femur and tibia very stout, 3.0-3.5 × longer than wide compared to 4.5-5.6 × in Patagobia; metatibia of Andesobia with one pair of spurs, two pairs in Patagobia; medial line of forewing absent in Andesobia, present in Patagobia; postmedial line never double in Andesobia, often double in Patagobia; hindwing discal spot small and sharpor absent in Andesobia, diffuse and more elongate in Patagobia. Andesobia is endemic to the Puna grasslands of the high Andes of Peru and Bolivia.
Description. Male. Head -vestiture dark brown to black, shaggy appearance, setae long; antenna weakly bipectinate, ciliate ventrally; longest posterior rami 1.3-2.0 × segment length, longest anterior rami 1.1-1.8 × segment length; rami longest over middle third of antenna, decreasing in length toward base and apex; eye elliptical, 1.4-1.6 × as high as wide; labial palps short, not extending beyond vestiture of frons; 2 nd labial segment short and stout, 1.1 × as long as wide, 2 × longer than apical segment; haustellum reduced and poorly sclerotized, presumably non-functional. Thorax -vestiture of vertex and ventrum of thorax black brown; tegulae and patagia black brown; legs black brown, dorsum of femur ochre or dull pinkish red, co-varying with hindwing and abdomen ground colour; apex of prothoracic tibia with two subequal, blunt, triangular projections; two meso-and metathoracic tibial spurs, posterior spur slightly longer than anterior, length of spurs approximately equal to tibial width at apex; metepisternum with rounded ridge along anterior margin, metepisternal microtymbals absent. Forewing -relatively small for an arctiine, forewing length 8-13 mm, elongate with apex less rounded than in Paracles and Spilosoma, length:width ratio averaging 2.2; ground colour ochre yellow, whitish to pinkish red or brownish grey; markings varying from obsoloete (A. jelskii) to well defined, grey-brown transverse bands; when present, darker pattern consisting of dark-brown basal area, sub-basal band, discal spot, postmedial band and marginal band; bands occasionally confluent along anal margin; ventrally with bands obsolete except for marginal band, and with a brighter yellowish or reddish ground colour. Hindwing -ground colour slightly richer yellowish or reddish than forewing, with dark-brown to grey-brown marginal band, varying from nearly obsolete (reduced to intermittent diffuse spots extending from apex halfway to anal angle), to broad and diffuse over distal third of wing; brownish, crescentic discal spot small but usually well defined, sometimes absent; ventrally with dark markings less saturated. Abdomen -Segments A1-A3 entirely brownish black, remaining segments ochre or reddish subdorsally, with brownish-black dorsal line, widest in A. flavata; ventrally, varying from entirely brownish black (A. sanguinea) to black with narrow ochre border on distal margin of sternites (A. flavata) or entirely ochre (A. jelskii); coremata highly reduced to paired patches of sparse, deciduous setae. Genitalia -highly simplified overall with massive, triangular dorsoventrally flattened uncus characteristic of subtribe; uncus as long as width of base, broadly joined to wide, band-like tegumen; dorsal margin of tegumen caudally recurved; valve simple and digitate, lacking processes or claspers, 1-1.7 × as long as uncus-tegumen complex; vinculum semicircular, saccus v-shaped, similar in length to uncus; juxta evenly convex and hemispherical, dorsal margin slightly narrowed; aedeagus relatively large and stout, 3 × longer than wide, 1.5 × as long as width of genital capsule, curving dorsad 25-30°, proximal end approximately ⅓ narrower than apex; coecum 1/10 length of aedeagus, directed slightly ventrad; vesica directed dorso-distad, globose, finely spiculate, with small basal and poorly differentiated apical diverticulum. Female (A. jelskii and A. sanguinea only; female of A. boliviana and A. flavata unknown). Head -antennae 0.5 × as long as that of male, finely biserrate; proboscis atrophied; vestiture of closely appressed, ochre scales, lacking long, shaggy black scales present in males. Thorax -vestiture similar to that of head, notably lacking 'shaggy' appearance of males; legs reduced, 2/3 as long as those of male. Forewing and hindwing -micropterous and highly reduced, forewing 1.5-2.5 mm long, fully scaled and concoulours with dull tan colour of thorax, but without any discernible wing pattern. Abdomen -light ochre gray with fine, short velvety hairs, tergites well sclerotized, black, giving dorsum of abdomen appearance of a broad, black medial band; ventrally with narrower, lighter grayish-black medial band; integument broad and membranous laterally, allowing for distension caused by ova. Genitalia (based on A. jelskii) -ostium and lamella antevaginalis membranous and poorly defined; lamella postvaginalis consisting of a broad, shallow sclerotized pouch; ductus bursae lightly sclerotized, dorsoventrally flattened, 2 × as long as wide; corpus bursae pear shaped, and relatively small, 2 × as longh as ductus bursae; diameter of distal, globose chamber 2 × width of ductus bursae; signum lacking; ductus seminalis wide and rugose, bulla seminalis large, diameter 1.5 × that of corpus bursae; posterior apophysis equal in length to papillae anales, anterior apophysis 0.6 × as long as papillae anales; each paired dorsal pheromone gland consisting of two tree-like subdivisions, each subdivision with 3-5 smaller diverticula.
Remarks. Structurally, Andesobia is quite homogeneous, the main species-level differences occuring in the length and shape of the male valve and the vesica. The highly simplified, digitate male valve and massive uncus-tegumen compex is shared with several other Neotropical genera including Paracles, Patagobia, Caribarctia Ferguson and Leichosila Schmidt. The mtDNA barcode sequence (A. jelskii) does not provide any additional resolution of relationships within this group, with minimum pairwise distances (uncorrected) between Andesobia, Paracles, Phragmatobia, Leichosila, Caribarctia and Phaos ranged from 6-8%. Sequences for Patagobia were not available.
Several Andean species are superficially similar to Andesobia and Patagobia, and require comment. Paracles herbuloti (Toulgoët), Paracles minuta Becker & Miller, and Paracles diminuta Becker & Miller are small species with a simple or highly reduced forewing pattern. Females of all three are unknown, but the structurally similar and probably congeneric Chilesia anguloi Ruiz, C. rudis (Butler) and C. watsoni Ruiz have micropterous females (Ruiz 1989;Vargas and Parra 2003). Despite these similarities to Andesobia (and Patagobia), the broader, more rounded wings, shorter, rounder saccus, greatly elongated tegumen, very short valva, and small vesica are consistent with those of other Paracles species, and not with Andesobia or Patagobia.
Biology and distribution. Data on the biology of Andesobia is based primarily on A. jelskii and is discussed in more detail under the species account below. Andesobia is adapted to cold-temperate alpine habitats, males flying during sunny periods and the females being micropterous. Mating and oviposition occurs inside the female cocoon. The female-biased sex ratio of the broods reared during this study may indicate that females are capable of parthenogensis, as in some other cold-adapted flightless Lepidoptera (Suomalainen 1962). Adults emerge during the middle of the four-month wet season in the otherwise xeric grassland habitat. Andesobia is endemic to the Puna grasslands of the high Andes, occuring from central Peru south to the Lake Titicaca region of southern Peru/Bolivia. Diagnosis and re-description. Andesobia jelskii was omitted from the catalogue of Neotropical Arctiinae (Watson & Goodger 1982), probably because Hampson (1901) placed it in the Palaearctic genus Ocnogyna. This species is most similar to A. flavata and A. boliviana, but lacks any trace of transverse forewing lines and has an uninterrupted ochre costal band, which is interrupted by the antemedial and postmedial lines in other Andesobia species; the hindwing marginal band has a diffuse inner border and is much wider than in other species of Andesobia, extending at least to the discal area. Internally, the male valve is shorter and wider, appropximately 6 × longer than the narrowest diameter compared to 6.7-8.0 × longer in A. flavata.
Immature stages. Egg -almost spherical, poles only very weakly flattened; micropyle very weakly sculptured, barely visible; ivory white changing to dark greyish white prior to hatching. Larva -1 st instar larva initially translucent white, becoming opaque white; setae black, yellow orange prior to molting. 2 nd instar integument black, more densely setose than 1 st instar. 3 rd instar with jet black setae, except rusty brown on A2-A5; spiracles white. 4 th instar (Figs 15a, b) with verrucae more pronounced than in previous instar; setae jet black with silver sheen apically, somewhat lighter smoky grey subventrally; colour of setae polymorphic in last instar, either with A2-A5 yellowish orange and segments A6-A8 with subdorsal and lateral silvery-white setae mixed in (Fig. 15a), or with orange setae very dark brown to black (Fig. 15b); when mature, female larva about twice as large as male larva. Pupa -cremaster short, penicillate, head compact with short, stiff setae (Fig. 12). Cocoon spherical to ovoid, reddish brown to dull brown, consisting of a single, thin and flimsy layer with incorporated larval setae (Fig. 16). Biology and distribution. Eggs whitish, turning dark grey three days prior to hatching, hatching in about 10 days. First instar larvae initially feed on the tissue of the dead or dying female, then leave the cocoon in search of plant material. Duration of the first instar is six days, second instar five to six days. Taraxacum F.H. Wigg. and lawn grass (Poa L.) are both acceptible food plants in captivity, suggesting that larvae are polyphagous in nature. Notably, larvae emit an unpleasant odour of decay when disturbed. In late instars, female larvae are twice as large as male larvae. During the first three instars, larvae avoided sunlight, but the last two instars showed increased tolerance to sunlight, possibly to accelerate development. Cannibalism was not observed even at higher densities. Males pupated sooner than females, but the pupal stage is shorter in females lasting only a few days, so emergence of both sexes is more or less synchronous. Cocoons (Fig. 16) were spun between leaves of the food plants near the ground. The moths emerge in the morning, with relatively fast expansion of the wings. Females remain in the loosely-spun cocoon, and presumably emit mating pheromones from within the thin cocoon soon after emerging from the pupa, because males dig through the loose silk webbing to enter and mate inside the cocoon (Fig. 16). The pair remains in copula for several hours, after which the male leaves the cocoon, and the female deposits about 50 eggs inside the cocoon. Males are diurnal and fly rapidly during sunny periods. Reared cohorts of A. jelskii displayed an unequal sex ratio of about 5 female: 3 male; female microptery and a female-biased sex ratio is associated with parthenogenesis in other families (Heterogynidae, Psychidae, Lymantriinae: Teia Hübner), and Andesobia may also be capable of parthenogenesis, which has not been documented in the Arctiinae. Andesobia jelskii is currently known only from the Junin region of Peru (Fig. 17), at 4100 m elevation in the Puna grassland ecoregion of the central Andes (Fig. 18). Like other members of the genus, the flight period is early in the year (January), in the middle of the four-month wet season.
Andesobia flavata (Hampson, 1901), comb. n. http://species-id.net/wiki/Andesobia_flavata Maenas flavata Hampson, 1901: 512, pl. 51, f. 7 Type material. The three examined male syntypes of Turuptiana flavescens exhibit variation in the extent of the forewing markings, two specimens closely approaching the appearance of the A. flavata holotype (see also Remarks under A. flavata). The third syntype labeled "type" with a round, red-bordered label and a blue label reading "genitalia slide no. 3422" is here designated as lectotype; it is an almost exact match to the two specimens illustrated here (Fig. 2).
Remarks. Subsequent to its description, Estigmene boliviana (Fig. 2b) disappeared from the literature. Although appearing in the print version of the Zoological Record for 1923, it is absent from the digital version of Zoological Record and the card index of the BMNH (Global Lepidoptera Names Index 2011). It was also omitted by Watson and Goodger (1986). The holotype label data is as follows: "La Paz / Bolivia / 95 Garlepp"; "als Puppe / Nov. 8. larva"; "1855a"; "Coll. / Staudinger"; "Estigmene / boliviana G. / E. Rschau [Entomologische Rundschau] B40 S20."; "Type". (Hampson, 1907) Type material. Hampson (1907) based his description on what he believed to be a male and a female, but both specimens (BMNH) are males. One syntype is in rather poor condition with rubbed, partially broken wings, lacking antennae and abdomen. The second syntype, in excellent condition, labeled "Type" with a round, red-bordered label and a blue label reading "genitalia slide no. 3423" is here designated as lectotype to ensure the stability of the name. Diagnosis. Andesobia sanguinea is the only member of the genus with red colouration, prevalent on the hindwing and often the forewing, the latter varying from whitish pink (Fig. 3a) to whitish tan (Fig. 3b). Females are micropterous and are similar to A. jelskii, but with a more yellowish colour (Fig. 6). The biology is unknown. It appears to be sympatric with A. boliviana, and is known only from the Lake Titicaca region (Fig. 18).  Rothschild, 1910. Etymology. The name is derived from a combination of the words Patagonia and Phragmatobia.

Andesobia sanguinea
Diagnosis. Although Patagobia shows similarities to the Holarctic Phragmatobia in some external aspects, it differs in having longer, symmetrical rami of the male antenna, ochre thoracic collar, lack of a male clasper, pale tan forewing pattern (usually), and a restricted distribution to the Chilean Andes of South America. The wing colour and pattern is also similar to Andesobia, but structurally Patagobia has a more robust build with denser thoracic vestiture, equally long posterior and anterior male antennal rami (anterior rami shorter than posterior in Andesobia), male antennal rami up to 3 × longer than antennal segment length (up to 2 × in Andesobia); 2 nd labial segment elongate, 1.8 × as long as wide, 1.5 × longer than apical segment; thoracic collar contrastingly paler ochre (conconcolourous with dorsal thoracic vestiture in Andesobia); thoracic vestiture dense and pilose (sparse and shaggy in Andesobia); femur and tibia elongate, 4.5-5.6 × longer than wide (very stout, 3.0-3.5 × longer than wide in Andesobia); metatibia with two pairs of spurs (one pair in Andesobia); forewing medial line present (absent in Andesobia); postmedial line usually double (absent in Andesobia); hindwing discal spot diffuse and elongate (sharp or absent in Andesobia). The male coremata betwen the 7 th and 8 th sternite are moderately developed in Patagobia, very reduced in Andesobia.
Description. Male (female unknown). Head -vestiture dark brown to black, setae long; antenna bipectinate, ciliate ventrally; longest posterior rami 1.5-3.0 × segment length, longest anterior rami 1.1-3.0 × segment length; rami longest over middle third of antenna, decreasing in length toward base and apex; eye elliptical, 1.2-1.5 × as high as wide; labial palp short, not extending beyond vestiture of frons; haustellum reduced and poorly sclerotized, presumably nonfunctional. Thorax -vestiture of vertex and ventrum of thorax black brown; tegulae entirely black brown or black brown edged with yellowish brown; patagia yellowish brown or rarely black brown; leg vestiture brownish ochre, dorsum of femur yellow or red, co-varying with hindwing and abdomen ground colour; apex of prothoracic tibia with two subequal, blunt, triangular projections; mesotibia with two apical and two subapical spurs, length of apical spurs 1.5 × and supapical spurs 0.6 × tibial width at apex; two metatibial spurs, posterior spur slightly longer than anterior; metepisternum lacking microtymbals. Forewings -forewing length 12.9-13.2 mm (mean 13.1 mm; n = 4), length:width ratio averaging 2.1; ground colour pale ochre yellow but with broad, sometimes entirely confluent dark-brown transverse bands (Fig. 4c); pattern elements consisting of dark basal area and sinuous, diffuse dark-brown transverse lines (Fig. 4a) discal spot indistinct dorsally, but well defined ventrally; ventrally with bands obsolete except for marginal band. Hindwing -ground colour pinkish red or rarely yellow (Fig. 4d), with broad dark-brown marginal and costal band; well-defined brown, crescentic discal spot; similar ventrally but with discal spot better defined. Abdomen -Vestiture brownish black and pinkish red or yellow subdorsdally, ventrally with segmental margins yellowish ochre; abdomen entirely dark brown in melanic specimens (Fig. 4c). Coremata between sternites 7-8 in shallow pockets, scent scales approximately 0.5 × as long as sternite length. Genitalia -highly simplified, with large, triangular, dorsoventrally flattened uncus characteristic of subtribe; uncus 1.5 × longer than width of base, broadly joined to wide, band-like tegumen; dorsal margin of tegumen caudally recurved; valve simple and digitate, lacking processes or claspers, 1.5 × as long as uncus-tegumen complex; vinculum semicircular, saccus v-shaped, similar in length to uncus; aedeagus large, 5-6 × longer than wide, 2 × as long as width of genital capsule, curving dorsad slightly; coecum 1/10 length of aedeagus, directed slightly ventrad; vesica extremely large, diameter when inflated 2 × that of genital capsule; vesica directed right-laterad, globose, finely spiculate, with poorly differentiated basal chamber and large apical chamber (Fig. 11b). Patagobia thursbyi (Rothschild), comb. n. http://species-id.net/wiki/Patagobia_thursbyi Turuptiana thursbyi Rothschild, 1910: 176, comb. n Diagnosis. The taxon pluto Toulgoët has been treated as a subspecies distinct from nominate thursbyi based on the nearly unicolourous forewing, resulting from the confluence of the transverse bands. Genitalic structure of both taxa is identical (Toulgoët 1987). Examination of series of specimens from a single locality (Fig. 4b-d) shows that there is considerable variation in the extent of forewing banding, and also in the hindwing ground colour. We therefore treat pluto as a synonym of nominate P. thursbyi.

Discussion
In his global review of Arctiina genera (as Arctiini), Ferguson (1985) divided the genera among five groups, namely "Neoarctia-Grammia", "Holomelina" (now Virbia Walker; Zaspel and Weller 2006), "Arctia-Hyphoraia", "Spilosoma" and "Phragmatobia-Ocnogyna". The latter two do not form natural groups, but are monophyletic as a whole and were treated as subtribe Spilsomina by Schmidt (2007), with the exclusion of Kodiosoma Stretch, which is related to the Euchaetes group (Schmidt 2007;Schmidt & Opler 2008). Ferguson considered the Andean "Phragmatobia" species to be closely related to and possibly congeneric with Metacrias Meyrick, 1887 of New Zealand. Metacrias is probably congeneric with Phaos Walker, 1855 of Australia / Tasmania (Patrick et al. 2003). Patagobia is structurally more similar to Metacrias/Phaos than to Andesobia, the latter apparently representing a more ancient split from Patagobia + Metacrias / Phaos (PMP) lineage. This implies that the Andean group is not monophyletic, and that the PMP group has a circumantarctic distribution. The biology of the PMP group provides additional evidence for a common ancestry; all inhabit mesic, cool-temperate habitats, particularly grassland and tussock tundra, with specialized adaptations such as diurnal flight (males) and flightless females. Land connections between southern South America and Australia existed as late as the Late Eocene (~35 MYA) (McLoughlin 2001), but the biogeography of New Zealand Metacrias is more difficult to explain, since Zealandia separated from other Gondwanan landmasses about 80 MYA (McLoughlin 2001). Transoceanic dispersal events would be highly unlikely for the flightless females, and early instar larvae are not known to exhibit ballooning behaviour (wind dispersal by silk strands) (Gibbs 1962). The zoogeography of this fascinating group awaits further study.