Systematic position of Apodemia paucipuncta (Riodinidae), and a critical evaluation of the nymphidiine transtilla Author Penz, Carla M. Author Devries, Philip J. text Zootaxa 2006 2006-05-01 1190 1 50 journal article 27032 10.5281/zenodo.2646145 041d75f1-4053-4528-9d19-9efee178c59b 1175-5326 2646145 35C119EB-0729-41BF-A36B-E034E611CE38 Status of Adelotypa The 14 representative species of Adelotypa sorted into four separate lineages ( Fig. 3 ). Adelotypa eudocia paired with Apodemia paucipuncta (see above), A. zerna (Hewitson) and relatives constituted the sister group of Catocyclotis , and two other groups emerged within Calospila . Characters supporting these relationships are found in Table 2 . TABLE 2 . Character state changes assigned to groups A–I in Figure 3. Abbreviations follow terminology in MacClade (Maddison & Maddison 2000): ho, homoplasious outside; hao, homoplasious above and outside; u, unique, changing above; uu, unique, unchanged above. Unique character state changes are in bold. Groups A and E refer to Hallonympha and Harveyope , respectively.

group	character
A	10:1 (uu): In lateral view, uncus arched upward
29:6 (ho) Cornuti: multiple spines, short, thick, heavily sclerotized
70:1 (ho) Second segment of labial palpus, long, erect hair­like scales projecting ventrally
beyond the flattened scales present
B	27:1 (hao) Sculpturing of the vesica present
42:1 (ha) Dorsal processes of valva, when separated, not extended to form a sclerotized
bridge
48:0 (uu) When dorsal process separated, ventral process of valva extended at base to
form a sclerotized bridge
59:1 (ho) Ventral bracing of valvae forming a bridge between the valvae
C	1:1 (ha) Location of 3rd abdominal spiracle: at midline
3:0 (ho) Stn8 with terminal projection extending beyond edge of pleural membrane
13:0 (uu) Lateral margins of tegumen thickened at edges of lateral fenestra to form ribs
64:1 (hao) Number of spots in ventral hindwing cell cell Rs: 1
66:1 (hao) Dorsal forewing, two marginal lines present (continuous or broken)
D	11:1 (ho) In lateral view, dorsal outline of tegumen completely straight
20:0 (hao) Cuticular sculpturing prominent around subscaphium
43:1 (hao) Dorsal process of valva more sclerotized distally
60:1 (ho) In ventral view, anterior edge of valva projected
69:1 (hao) Ventral hindwing of much lighter color than ventral forewing
E	18:1 (ha) Sclerotized plate of the subscaphium uniformly broad
19:1 (ho) Sclerotized plate of the subscaphium extends to the end of subscaphium lobe
26:2 (ho) In ventrolateral view, vinculum laterally widened below tegumen to form a blade
that maintains its width towards saccus
28:2 (u) Sculpturing of the vesica: enlarged spines
29:1 (hao) Cornuti: a simple plate
54:1 (hao) In ventrolateral view, proximal region of ventral process slightly raised,
rounded
70:1 (ho) Second segment of labial palpus, long, erect hair­like scales projecting ventrally
beyond the flattened scales present
F	29:5 (ha) Cornuti: two spines
40:1 (u) Ventral process of valva extends posteriorly beyond dorsal process
62:0 (hao) In rear view, shape of the outline of the genitalic capsule (as formed by the
tegumen + vinculum + saccus): []
64:3 (ha) Number of spots in ventral hindwing cell cell Rs: 3
G	14:1 (ho) Notch in the anterior margin of tegumen present
15:0 (ho) Posterior end of subscaphium: one narrow lobe
29:3 (ho) Cornuti: a plate with two terminal spines of uneven size

TABLE 2 (continued)

Group	Character
31:0 (uu) When acutely pointed, aedeagus tip short
38:0 (ho) In lateral view, juxta arch extending beyond edge of valvae
46:1 (ho) Ridge of dorsal process at an angle with axis of valva
50:0 (ho) Ventral process of valva uniformly sclerotized
51:1 (ho) Distal portion of ventral process of valva not forming a point
57:0 (ho) In ventrolateral view, lateral fissure transverse to long axis of valva
62:3 (ha) In rear view, shape of the outline of the genitalic capsule (as formed by the
tegumen + vinculum + saccus): \_/
H	4:0 (ho) Stn8 simple, not divided
I	1 :2 (hao) Location of 3 rd abdominal spiracle: below midline
25:1 (u) Vinculum: distal edges fused to anterior edge of tegumen, remainder of vinculum
connected to tegumen by weakly sclerotized tissue
26:0 (hao) In ventrolateral view, vinculum relatively narrow throughout its entire length,
only mildly varying in width
29:7 (ha) Cornuti: multiple spines, long, thin, lightly sclerotized
52:3 (u) Distal portion of ventral process of valva bent upward
62:2 (hao) In rear view, shape of the outline of the genitalic capsule (as formed by the
tegumen + vinculum + saccus): ()
71:0 (hao) Frons scales: mixed broad and thin, converging from sides to midline
72:1 (hao) Color of the cuticle of the ventral mesothorax: orange­brown

Species in the zerna ­group constitute the sister lineage to Catocyclotis based on five character state changes ( Table 2 D ), but seven changes set them apart from Catocyclotis , one of which is unique (sculpturing of the vesica: enlarged spines, 28:2; Table 2 E ). Adelotypa zerna and close relatives can therefore be considered a well­defined species group within a larger assemblage including two other Nymphidiini genera. Bremer support for this group is moderate ( Fig. 3 ). These results demonstrate the monophyly of the zerna ­ group outside of the remaining Adelotypa . Adelotypa bolena (Butler) ( type species of Adelotypa ), borsippa (Hewitson) and penthea (Cramer) grouped with six species of Calospila , including parthaon (Dalman) ( type species of Calospila ). This grouping is supported by four character state changes, one of which is unique (ventral process of valva extends posteriorly beyond dorsal process, 40:1; Table 2 F ). Bootstrap and Bremer support values for this group are low ( Fig. 3 ). The amasis ­group is strongly supported by ten character state changes, one unique and universal (tip of aedeagus acutely pointed and short, 31:0; Table 2 G ), and bootstrap and Bremer support values for this group are high ( Fig. 3 ). The association of the amasis ­ group with Calospila emylius (Cramer) + cilissa (Hewitson) is supported by a single character state change ( Table 2 H ), and bootstrap and Bremer support values for this group are low ( Fig. 3 ), underscoring the need for a phylogenetic reassessment of these groups. Status of Calospila The nine species representing Calospila appear to be paraphyletic with respect to two lineages of Adelotypa . These relationships are strongly supported by eight character state changes, including two that are unique (distal edges of vinculum fused to anterior edge of tegumen, remainder of vinculum connected to tegumen by weakly sclerotized tissue, 25:1; distal portion of ventral process of valva bent upward, 52:3; Table 2 I ). Bootstrap and Bremer support values are moderately high ( Fig. 3 ). This group is not fully resolved in the strict consensus tree because of the uncertain position of Calospila apotheta (Bates) ( Fig. 2 ). In five of six equally parsimonious trees C . apotheta appeared as sister species to the amasis ­group, but in one tree it was basal to the group including Calospila parthaon and Adelotypa bolena (trees not illustrated). Successive approximation weighting favored the latter topology ( Fig. 3 ), but instability in tree topology points to the need for further examination of C. apotheta in the context of increased taxon and character sampling. Descriptions of new genera Our analysis demonstrated that paucipuncta and eudocia together constitute a monophyletic group well separated from the remaining Adelotypa . Placing the paucipuncta ­group as a sister clade to any of the three Adelotypa clades in Fig. 3 produces a considerable increase in tree length ( bolena + borsippa + penthea : 21 steps: amasis ­ group: 16 steps; zerna ­group: seven steps). Based on the shortest, most parsimonious tree, we describe a new genus for eudocia and paucipuncta . Furthermore, subsuming the paucipuncta ­group into any of the existing genera studied here (e.g., Nymphidium ) would require an unwieldy number of generic changes. Our results showed that the zerna ­group is monophyletic, and that it does not group with other lineages of Adelotypa ( Fig. 3 ). Placing the zerna ­group as a sister lineage to bolena + borsippa + penthea or the amasis ­group increases tree length by 30 and 18 steps, respectively. The zerna ­group differs substantially from Catocyclotis in color pattern; in the former the forewing and hindwing are similar in dorsal coloration, whereas in the latter they differ (see character 68 in Appendices 1 and 2, and Figs. 1 and 5 ). Based on the shortest, most parsimonious tree and differences in color pattern, we describe a new genus for the zerna ­group. Differences in wing coloration make the new genus easily recognizable from Catocyclotis . Hallonympha Penz & DeVries , NEW GENUS Type species: Apodemia paucipuncta Spitz, 1930 Description ( Fig. 4 ) Character state changes from our analysis are indicated by numbers in parentheses. Head : predominantly brown. Frons brown, bordered in white and covered mostly with long hair­like scales projecting anteriorly. Head apex with long scales projecting anteriorly. Second segment of labial palpus laterally white ( paucipuncta ) to brown + white ( eudocia ), with long hair­like scales projecting ventrally (character 70:1); first and third segments brown. Antenna brown with white rings, club orange at tip. Body : dorsal body scales brown, ventral body and leg scales silvery beige ( paucipuncta ) to brown ( eudocia ). Wings : FW length 10–11 mm (n=7). Sexes similar. See Fig. 4 for wing shape, venation, and location of spots. Dorsal wing color pattern : brown with darker brown spots contained inside cells, and a band across the distal edge of both FW and HW discal cells. Dark brown spots with neighboring bright white spots ( paucipuncta ), or lacking such white spots ( eudocia ). FW and HW similar in color and pattern. Ventral wing color pattern : slightly paler than dorsal surface. Brown spots with neighboring bright white spots ( paucipuncta ), or bordered dirty white ( eudocia ). Male terminalia : Sternite 8 not extended posteriorly beyond the pleural membrane. Genitalic capsule short and compact. Uncus arched dorsally (character 10:1), small central indentation vestigial or absent. Sclerotized plate of subscaphium broad, diamond­shaped. Gnathos sickle­shaped with a blunt tip. Vinculum not continuous through anterior edge of tegumen. Aedeagus (=phallus) with a long, acute tip. Coecum penis small. Cornuti comprising multiple short, thick and heavily sclerotized spines (character 29:6). Valvae clearly divided into two processes; dorsal processes separated laterally, ventral processes forming a bridge above aedeagus. Tip of ventral process reduced and fused to bridge ( paucipuncta ), or reduced and not visible ( eudocia ). Lateral, rounded extension of ventral process protruding outward to form a small flap. Remarks on female genitalia We observed the following female genitalic characters in paucipuncta ( Fig. 4 ) and eudocia : (1) antrum broad and well­sclerotized; (2) ductus bursa with a mildly sclerotized, internally spined enlargement anterior to corpus bursa in paucipuncta , ductus bursa simple in eudocia ; (3) corpus bursa moderately elongated; (4) paucipuncta with signa symmetrically placed in corpus bursa, asymmetrically positioned in eudocia . Differences in female genitalic morphology between paucipuncta and eudocia appear to be comparable to those found between Catocyclotis aemulius (Fabricius) and adelina (Butler) (see illustrations in Penz & DeVries 2004 ). Natural history Hallonympha paucipuncta is an ant­associated, polyphagous species endemic to the Brazilian cerrado ( DeVries et al. 2004 and references therein). The larvae have balloon setae, vibratory papillae, and tentacle nectary organs similar to other Nymphidiini ( DeVries et al. 2004 ) , but are unique within Riodinidae by possessing a cervical gland used in myrmecophily. The A3–7 spiracles of paucipuncta are located dorsally, a character state that is also present in Catocyclotis adelina . FIGURE 4. Hallonympha paucipuncta : adult male in dorsal and ventral views; wing scheme showing venation and location of dark (heavy stippling) and white markings (open); male genitalia in lateral and ventral views; male sternum 8 (note extended membranous area with a small, ventrolateral cluster of setae); female genitalia (inset: ventral view of ostium bursa). Scale bars: 1 mm. The relationships between eudocia and paucipuncta suggest that the larva of eudocia (currently unknown) may also possess balloon setae, dorsal spiracles and a cervical gland. In Mexico , eudocia inhabits a habitat similar to that of paucipuncta (A. Warren pers. com.). Natural history studies of this species will be of much interest. Etymology We name this new genus after J. P. W. Hall in recognition for his contribution to riodinid systematics. Species in the genus Based on our phylogenetic analysis ( Fig. 3 ) and generic definition (above), we place the following species in Hallonympha : Hallonympha paucipuncta ( Spitz, 1930 ) [ type species], new combination Hallonympha eudocia (Godman & Salvin, 1897) , new combination