Agraulis Boisduval & Le Conte, 1835
publication ID |
https://doi.org/ 10.11646/zootaxa.5209.3.8 |
publication LSID |
lsid:zoobank.org:pub:FA768332-156C-47D9-A115-84443D948177 |
DOI |
https://doi.org/10.5281/zenodo.7334259 |
persistent identifier |
https://treatment.plazi.org/id/03B68171-FFD2-BA08-FF50-A5A70DD7FBDA |
treatment provided by |
Plazi |
scientific name |
Agraulis Boisduval & Le Conte |
status |
|
Agraulis Boisduval & Le Conte View in CoL , reinstated status
** Agraulis incarnata incarnata (Riley)
** Agraulis incarnata nigrior Michener
* Agraulis vanillae (Linnaeus)
* 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 View TABLE 1 and 2 View TABLE 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.
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 |
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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. |
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 |
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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 View 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 View 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 View 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 View 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
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