Agraulis Boisduval & Le Conte, 1835

Penz, Carla M., 2022, Reinstated status of the butterfly genus Agraulis (Lepidoptera, Nymphalidae, Heliconiinae), Zootaxa 5209 (3), pp. 394-398 : 395-397

Agraulis Boisduval & Le Conte , reinstated status

* Agraulis incarnata (Riley)

** Agraulis incarnata incarnata (Riley)

** Agraulis incarnata nigrior Michener

* Agraulis vanillae (Linnaeus)

* Agraulis forbesi Michener

* Agraulis insularis Maynard

* Agraulis maculosa (Stichel)

* Agraulis lucina C. Felder & R. Felder

* Agraulis galapagensis Holland

* Agraulis dodona (Lamas & Farfán)

( Agraulis )

( Dione )

This study proposes, and provides a justification for, the reinstatement of Agraulis to generic status. Sister-taxon relationships between Agraulis and Dione have been supported independently by analyses using morphology ( Penz 1999) and multiple DNA markers demonstrated to be informative from species to family levels ( Wahlberg et al. 2009, Kozak et al. 2015, Zhang et al. 2019, Chazot et al. 2021). In support for the genus rank of Agraulis , morphological characters that distinguish Agraulis and Dione are listed in Tables 1 and 2 . Finally, Nuñez et al. (2022) provided intra and intergeneric mean genetic distances for 12 molecular markers that range from 0.2–2.1% among species of Agraulis , 0.87–5.12% among those of Dione , and 2.85–10.47% between these genera. Given their findings, a comprehensive taxonomic revision of Dione is warranted.

Tables

TABLE 1. Morphological characters that support the relationships between Agraulis and Dione plus those that distinguish

(L6).	Setae D1 on thoracic segments 1–3 equal to, or longer than, the head height in the first instar larvae. This character
was also observed in Dryadula phaetusa (Linnaeus) , Dryas iulia (Fabricius) , and Philaethria dido (Linnaeus) within
Heliconiiti, and also Speyeria mormonia (Boisduval) (Penz pers. obs.).
(L22).	Aboral region of mandibles enlarged in the fifth instar larvae. This character is also found in Euptoieta hegesia
(Cramer) and Heliconius doris (Linnaeus) .
L42.	Secondary setae distributed around the center of anal plate in the fifth instar larvae.
(P44).	Gold spots absent. This implies a loss of gold spots in the ancestor of ( Agraulis , Dione ) and a gain of this pattern in
the ancestor of Dione glycera and D. moneta . This hypothesis is equally parsimonious to two independent losses in
Agraulis vanillae and D. juno .
P52.	Distal end of antennae acute. In this clade the distal ends of the antennae do not reach the edge of the wing case, thus
producing the acute pattern.
(P53).	Apex of the head adorned with ridges or tubercles. This character was also found in Boloria selene (Denis &
Schiffermüller) (Penz pers. obs.).
(P58).	Dark pigment concentrated on the distal end of the tracheae. This character was also observed in Dryadula
phaetusa .
(P61).	Leg 3 not visible. Reversal in Dione juno .
P63.	Region along the dorsal suture of the mesothorax with a single keel: highest point along the suture in the mid-portion
of the mesothorax.
(P74).	Crown hooks bifid. This character was also observed in Actinote pellenea Hübner and Cethosia biblis (Drury) (Penz
pers. obs.).
(A81).	Distal segment of the female leg 1 tarsi with a groove formed by the swelling of the ventro-lateral portions. This
character is also observed in Euptoieta claudia (Cramer) (Penz pers. obs.).
A96.	Forewing cross-vein m2–m3 out-curved.
A118.	Juxta curved longitudinally upwards.Although in Agraulis vanillae both the juxta and the phallus are curved upwards,
the phallus of Dione did not show a correlated curvature with that of the juxta.
(A122).	Anterior edge of costae expanded beyond the pedunculi.
A134.	Filiform scales on stink clubs predominantly bifid.

( Agraulis , Dione )

TABLE 2. Morphological characters that distinguish Agraulis from Dione listed by Nuñez et al. (2022, Appendix I). Numbers were maintainedfrom theoriginal publication, which should be consulted for illustrations andfurther information. Comments following characters are mostly quoted verbatim from the original publication, but references were generally omitted. Character comparisons to Dione appear in parentheses after the characters listed for Agraulis.

1-	antennal club: 2 times as long as broad (3 times as long as broad in Dione ). Figured by Michener (1942), Agraulis
	antennal clubs are the broadest among Heliconiini .
2-	distribution of androconia: FW (FW & HW in Dione ). Condition shared only with Dryadula among the Heliconiini .
3-	FW outer margin: smooth, slightly concavous at mid portion (variously emarginated, deeply concavous at its mid portion in
Dione ). Emarginated in Cu1 and 1A and barely at M 3 in D. juno , emarginated in M2 and M 3 in D. glycera and D. moneta .
4-	Separation of HW anal veins ends: ≈ 2X distance 1A to Cu2 (≈ 2.5 X distance 1A to Cu 2 in Dione ).
5-	dorsal process of pulvillus: absent (present in Dione ). See A 87 in Table 1 .
6-	sexual dimorphism in origin of Cu veins at FW: present (absent in Dione ).
7-	crista on male genitalia valva: absent (present in Dione ). Nuñez et al. (2022) examined male genitalia of Agraulis
specimens at the ZSM representing incarnata (Rh1766) , insularis (Rh1754) , lucina (Rh1770, Rh1771), maculosa
(Rh1761) , nigrior (Rh1753, Rh1769) and vanillae (Rh1755, Rh1762). See A 125 in Table 1 .
8-	process on valva tip of male genitalia: short, with spines, posteriad pointed (elongated, without spines, anteriordorsal
pointed in Dione ). Modified from A 124 in Table 1 .
9-	dorsal process of valva: short, blunt and posteriad pointed (elongated, posterodorsal pointed, surpassing projection of tip
in Dione ).
10-	stink-club tip of female 8 th abdominal segment: rounded (squared in Dione ). Nuñez et al. (2022) examined female
	genitalia of Agraulis specimens at the ZSM representing incarnata (Rh1768) , insularis (Rh1772) , maculosa (Rh1767) ,
and vanillae (Rh1760) . See A 132 in Table 1 .
11-	scales on stink club: longly bifid (hairlike, tip simple or shortly bifid in Dione ).
12-	signum of female genitalia: constricted transversely near anterior end, spines large (without constriction, spines tiny in
Dione ).

Adult