#ZooKeys

ZooKeys 1216: 173-200 (2024)
DOI: 10.3897/zookeys.1216.129923

Research Article

Revision of the genus Digonis (Lepidoptera, Geometridae):
new species and new genera

Mario I. Ramos-Gonzalez'2®, Maria Francisca Venegas-Gonzalez', Carlos Zamora-Manzur®®,

Luis E. Parra’®

1 Departamento de Zoologia, Facultad de Ciencias Naturales y Oceanogrdficas, Universidad de Concepcion, Casilla 160-C, Concepcion, Chile

2 Programa de Doctorado en Sistematica y Biodiversidad, Facultad de Ciencias Naturales y Oceanogréficas, Universidad de Concepcion, Concepcion, Chile
3 Departamento de Ecologia, Facultad de Ciencias, Universidad Catélica de la Santisima Concepcion, Alonso de Rivera 2850, Concepcion, Chile
Corresponding author: Mario I. Ramos-Gonzdlez (marioramos@udec.cl)

OPEN Qaccess

Academic editor: Axel Hausmann
Received: 27 June 2024

Accepted: 27 September 2024
Published: 24 October 2024

ZooBank: https://zoobank.
org/1BBB8E67-1398-4D4F-B9CE-
B56B2B7A471C

Citation: Ramos-Gonzalez MI,
Venegas-Gonzalez MF, Zamora-
Manzur C, Parra LE (2024) Revision
of the genus Digonis (Lepidoptera,
Geometridae): new species and
new genera. ZooKeys 1216:
173-200. https://doi.org/10.3897/
zookeys.1216.129923

Copyright: © Mario |. Ramos-Gonzalez et al.
This is an open access article distributed under
terms of the Creative Commons Attribution
License (Attribution 4.0 International - CC BY 4.0).

Abstract

The taxonomic study of the Chilean Ennomini genera is still in its early stages. Within
this group, the maculation patterns of Chilean species are uniform and often inadequate
for distinguishing between many species, compounded by a lack of taxonomic revisions
focused on the genera within the tribe. In this study, the genus Digonis Butler, 1882, is
reviewed and redefined based on characteristics of wing patterns and genitalia. Digonis
comprises the following five species: D. aspersa Butler, 1882, D. cervinaria (Blanchard,
1852), D. punctifera Butler, 1882, D. gungnir Ramos-Gonzalez & Parra, sp. nov., and
D. apocrypha Ramos-Gonzalez & Parra, sp. nov. Additionally, D. cuprea Butler, 1882 is
synonymized with D. cervinaria (Blanchard, 1852), and all varieties of D. punctifera Butler,
1882, and D. cuprea are synonymized with their respective species. Furthermore, two
new genera are introduced: Phasmadigonis Ramos-Gonzalez & Parra, gen. nov., erected
for P alba (Butler, 1882), comb. nov., and Gugnelve Ramos-Gonzalez & Parra, gen. nov.,
established for G. butleri Ramos-Gonzalez & Parra, sp. nov. With the species and genus
descriptions, a comparative diagnosis, genitalia illustrations for all species, and wing
venation for each genus are provided.

Key words: Andean Region, Argentina, Chile, Ennominae, Ennomini, Nacophorini,
taxonomy

Introduction

Ennominae is the most diversified subfamily, comprising approximately half
of the described species within Geometridae (Gaston et al. 1995; Scoble 1999;
Pitkin 2002; Scoble and Hausmann 2007; Rajaei et al. 2022). However, due to
this extensive diversity (approximately 10,000 species), the taxonomic classifi-
cation at the tribal level has remained in constant flux, and there is still no con-
sensus on the taxonomic affinities of the recognized tribes, particularly when
considering taxa from the New World (Holloway 1993; Scoble 1995; Pitkin
2002; Beljaev 2006; Ounap et al. 2011; Brehm et al. 2019).

The most diverse tribes within the Chilean Ennominae, which are also better
studied, correspond to Odontoperini with almost 50 species (Rindge 1973,

173

Mario |. Ramos-Gonzédlez et al.: Revision of the genus Digonis (Lepidoptera, Geometridae)

1983; Parra and Henriquez-Rodriguez 1993; Pitkin 2002; Brehm et al. 2019),
and Diptychini with approximately 40 species (Rindge 1986; Parra 1999a,
1999b; Pitkin 2002; Parra et al. 2010; Parra and Hernandez 2010; Brehm et
al. 2019). In contrast, there have been few taxonomic studies conducted on
the other tribes (e.g., Gnophini: Parra and Vargas 2000; Macariini: Vargas et
al. 2005, 2020; Boarmiini: Vargas 2007, 2021; Nacophorini: Rindge 1973, 1983;
Parra 2018; Ennomini: Bocaz et al. 2016). Therefore, there is a great uncertainty
regarding the monophyly of many genera and their position in the systematics
of the group (Hausmann and Parra 2009; Brehm et al. 2019).

The tribe Ennomini is characterized by having a vinculum divided ventrally
by a membranous area, a paired m3 muscle inserted distally in the median
invagination in the basal portion of the juxta. Among Chilean genera of Enno-
mini, there is notable variability in wing maculation patterns, which has posed
challenges in correctly recognizing species. This is exemplified in genera such
as Syncirsodes Butler, 1882 (Bocaz and Parra 2005; Bocaz et al. 2016), Haso-
dima Butler, 1882 (Parra and Pascual-Toca 2003; Parra et al. 2009), Perusia
Herrich-Schaffer, 1855 (CZ-M, unpublished data), and Digonis Butler, 1882.

Digonis was described by Butler (1882) based on the wing shape and the
simple antennae of adults. The forewings exhibit a very acute apex, followed
by a concave outer margin that extends to vein M., where it terminates in a new
acute angle, giving the wings the appearance of having two pointed extensions
along their outer margin (Pitkin 2002).

Scoble (1999) recognized five species within Digonis: four originally de-
scribed by Butler (1882): D. aspersa, D. alba, D. punctifera, including the vari-
eties maculosa, acuminata, terranea, and fumosa, and D. cuprea, including the
varieties olivacea and fusca. Additionally, Scoble (1999) included one species,
D. cervinaria (Blanchard, 1852). Subsequently, in his review of neotropical En-
nominae genera, Pitkin (2002) identified the species comprising the genus as
D. aspersa, D. cervinaria, D. cuprea, and D. punctifera, excluding D. alba due to
significant differences in male genitalia.

Despite the general characterization of the genus provided by Pitkin (2002),
uncertainty still needs to be resolved surrounding the validity of the described
species and varieties. The maculation patterns are deceptively uniform and
insufficient for species recognition (e.g., Hausmann and Parra 2009). There-
fore, the aim of this contribution is to characterize the genus Digonis through
a taxonomic revision based on external morphology and genital structures. To
achieve this, we redescribe the genus Digonis, presenting its diagnostic charac-
ters; we provide an illustrated and annotated systematic species list, along with
new taxonomic changes, and describe two new genera and three new species.

Materials and methods

This study was based on 107 adult specimens deposited in the collections
mentioned below. The type material examined for each taxon is specified in the
Results section.

NHMUK The Natural History Museum, London (UK)
MNHN Muséum National d’Histoire Naturelle, Paris (France)
MNNC Museo Nacional de Historia Natural, Santiago (Chile)

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Mario |. Ramos-Gonzdlez et al.: Revision of the genus Digonis (Lepidoptera, Geometridae)

MHNC Museo de Historia Natural de Concepcion, Concepcion (Chile)

MZUC-UCCC Museo de Zoologia dela Universidad de Concepcion, Concepcion
(Chile)

ZSM Zoologische Staatssammlung Miinchen, Munich (Germany)

For specimen identification and comparison, we used the original descrip-
tions and type material when available. Adult specimens were externally pho-
tographed using a Sony Cybershot DSC-HX300 compact camera. Wing and
genitalia slides were prepared following the methods outlined in Parra (1991).
Nomenclature for genitalia and external characteristics followed Klots (1970)
and Scoble (1995), respectively. All prepared slides were photographed using a
Motic SMZ-171-TL trinocular stereoscopic microscope equipped with a 5-Mpx
Moticam Motic2500 digital camera. These photographs were then used to cre-
ate detailed illustrations of the microscopic preparations.

Information on the flight period and species distribution was obtained from
the labels accompanying each examined specimen. This information was sup-
plemented with records from the citizen science platform iNaturalist (https://
www.inaturalist.org). All iNaturalist records cited were identified by the first au-
thor as part of the “Polillas de Chile” project (https://inaturalist.mma.gob.cl/
projects/polillas-de-chile).

All species were assigned to the biogeographical provinces proposed by
Morrone (2015). The newly described genera were diagnosed through compar-
ison with the type species of other closely related genera in the region, based
on both external and internal morphological similarities.

Results
Taxonomy

Tribe Ennomini Duponchel, 1845
Digonis Butler, 1882

Digonis Butler, 1882: 360; Bartlett-Calvert 1886: 333; Angulo and Casanueva
1981: 12; Scoble 1999: 229; Pitkin 2002: 248.

Type species. Digonis aspersa Butler, 1882. By original designation.

Diagnosis. Digonis resembles Digonodes Warren, 1895, Gonogala Butler, 1882,
and the newly proposed genus Phasmadigonis gen. nov. particularly due to the
mucronate shape of the wings. However, it is distinguished from Digonodes by the
presence of a mucronate outer margin on the M, of the hindwings and the presence
of filiform (not bipectinate) antennae. It is recognized as distinct from Gonogala by
the absence of bipectinate antennae and M, in the hindwings. It differs from Phas-
madigonis by the presence of two accessory areoles in the forewings, R, arising
from R,,,, and the absence of a vein connecting Sc+R, to the discal cell in the hind-
wings. The monophyly of Digonis is supported by the following genital characters:
a U-shaped gnathos with a plate or a pair of denticulate lobes, a concave, sclero-
tized process on the costa with an extended cucullus, a furca armed with spines,
an aedeagus without cornuti, and a strongly denticulate annular signum.

ZooKeys 1216: 173-200 (2024), DOI: 10.3897/zookeys.1216.129923 175

Mario |. Ramos-Gonzadlez et al.: Revision of the genus Digonis (Lepidoptera, Geometridae)

Redescription. Antennae serrate in males and filiform in females. Thorax
and abdomen with brown to grayish scales. Forewings castaneous yellow-
ish, gray, brown, or coppery; costal margin in the apical area slightly arched;
outer margin concave between apex and M, vein. Wing venation (Fig. 1).
Two accessory cells; Sc in contact with first accessory cell, R, originates
near apex of second accessory cell, R,,,,, from apex of second accessory
cell, R, and R, stalked, R, terminates at termen; M, equidistant between M,
and M.,, M, slightly arched and ending in small mucronate apex; CuA, origi-
nates 1/10 before end of cell, CuA, originates near the midpoint of the cell.
Hindwings paler than forewings, with multiple dark brown scattered spots;
medial band conspicuous or faint; outer margin slightly mucronate. Sc+R, in
contact with radial stem up to middle of cell, Rs originates 1/10 before end
of cell, M, absent. Male genitalia with conical uncus; gnathos “U” shaped,
with a pair of prominences or a denticulated plate; subrectangular valvae
with a strong sclerotized and concave costal process; spiny furca; aedea-
gus unarmed. Female genitalia with subpyriform corpus bursae, annular
and strongly denticulated signum.

Distribution. This genus is distributed between latitudes 30°S and 47°S,
spanning the provinces from Elqui to Capitan Prat in Chile.

SctR1
Rs
M1
M3
CuA1
CuA2
1A

Figure 1. Wing venation of Digonis Butler, 1882. Scale bar: 10 mm.

ZooKeys 1216: 173-200 (2024), DOI: 10.3897/zookeys.1216.129923 176

Mario |. Ramos-Gonzdlez et al.: Revision of the genus Digonis (Lepidoptera, Geometridae)

Digonis aspersa Butler, 1882
Figs 2A-D, 3

Digonis aspersa Butler, 1882: 361; Bartlett-Calvert 1886: 333; Angulo and Casa-
nueva 1981: 12; Scoble 1999: 229; Pitkin 2002: 248.

Material examined. Syntype. CHILE * 1 male; Pines Valley; “XII” [labeled]; “Chili,
82-107” [labeled]; “Type” [red labeled]; T. Edmonds leg.; NHMUK.

Other material examined. CHILE — San Antonio Prov. * Algarrobo; 10-I-1950;
n.n. leg.; MZUC-UCCC. — Diguillin Prov. + 1 male; Ninhue; 19-IV-2011; G. Moreno
leg.; MZUC-UCCC + Las Trancas; III-2013; -36.889694, -71.471611; “Mirg-006”
[genitalia slide]; MZUC-UCCC « Las Trancas; 16-IV-2010; G. Moreno leg.; “UCCC-
MZUC-Lep 0258” [Museum ID]; MZUC-UCCC - 1 female; Las Trancas; 8-II-2011;
G. Moreno leg.; “UCCC-MUZ-Lep-1056” [Museum ID]; “Mirg-022” [genitalia slide];
MZUC-UCCC « Las Trancas; 20-I-2012; G. Moreno leg.; “UCCC-MZUC-Lep 0377”
[Museum ID]; MZUC-UCCC. — Concepcion Prov: 2 males; Concepcion; 8-I-2001;
J. Artigas leg.; MZUC-UCCC + 1 male; Concepcion, Cerro Caracol; 15-Il-2001;
J. Artigas leg.; “Mirg-003” [genitalia slide]; MZUC-UCCC * 1 male; Concepcion,
Cerro Caracol; 30-II-2002; J. Artigas leg.; MZUC-UCCC + 1 male; Concepcion;
3-ll-1960; Trampas leg.; MZUC-UCCC * Concepci6n; 22-I-1960; Trampas leg.M
MZUC-UCCC + Concepcion; 1-IV-1960; Trampas leg.; MZUC-UCCC * Concepcion;
5-III-1959; Trampas leg.; MZUC-UCCC + Concepcion; 21-III-1962; Trampas leg.;
MZUC-UCCC + Concepcion; 30-IX-2001; J. Artigas leg.; MZUC-UCCC * Concep-
cion, Cerro Caracol; 30-II-2002; J. Artigas leg.; MZUC-UCCC * Concepcion; 7-I-
1961; Trampas leg.; MZUC-UCCC + 3 males; Concepcion, Barrio Universitario;
4-V-1985; Carrasco leg.; MZUC-UCCC * Concepcion; 29-XII-1958; Trampas leg.;
MZUC-UCCC + Concepcion; 16-III-1960; Trampas leg.; MZUC-UCCC. —Arauco
Prov. * 1 male; Lanalhue; 22-l-2018; L. Parra leg.; “Mirg-001” [genitalia slide];
MZUC-UCCC * Lanalhue; 23-I-2018; L. Parra leg.; MZUC-UCCC. — Biobio Prov.
* PN. Laguna del Laja; 6-XII-2008; G. Moreno leg.; MZUC-UCCC. — Malleco Prov.
* Nahuelbuta; Rio Picoyquen; 22-XII-1960; Fetis leg.; MZUC-UCCC * Collipulli; 24-
X-2014; E. Sepulveda leg.; “UCCC-MZUC-Lep 0308” [Museum ID]; MZUC-UCCC =
Rio Blanco; 19/25-II-1995; H. Thdény leg.; ZSM. — Valdivia Prov. 1 male; Valdivia;
30-IX-1987; Trampas leg.; MZUC-UCCC * 1 male; Valdivia; “5003” [Museum ID];
MNNC. — Llanquihue Prov. » Katalapi; 22-IV-2011; MZUC-UCCC. — Coyhaique
Prov. * Coyhaique; I-1934; E. Ureta leg.; “Museo 5004” [Museum ID]; MNNC = 1
female; Lago Verde, R.N. Coyaique; 21-Il-2007; MZUC-UCCC. — Aysen Prov. = 1
female; Rio Maca, Cuenca Cuervo; -45.114703, -73.016065; 21-II-2008; “Mirg-
020” [genitalia slide]; MZUC-UCCC. — Capitan Prat Prov. - Los Mellizos; 22-I-
2008; Munoz-Escobar leg.; MZUC-UCCC.

Additional records. CHILE. — Itata Prov. *- La Palma; -36.567368,
-72.689177; 9-IX-2021; observed by Claudio Maureira and submitted to iNat-
uralist in: https://inaturalist.mma.gob.cl/observations/100908464. — Con-
cepcion Prov. * Concepcion; 4-XIIl-2021; -36.834994, -73.011375; observed
by Flor Susana and submitted to iNaturalist in: https://inaturalist.mma.gob.
cl/observations/102608301. * Concepci6n; 5-V-2023; -36.8354793349,
-73.0287251249; observed by Luis Chavarriga and submitted to iNaturalist in:
https://inaturalist.mma.gob.cl/observations/159871318. * Coliumo; 9-II-2023;
-36.5526014, -72.9574049; observed by fpizarro and submitted to iNaturalist in:

ZooKeys 1216: 173-200 (2024), DOI: 10.3897/zookeys.1216.129923 177

Mario |. Ramos-Gonzédlez et al.: Revision of the genus Digonis (Lepidoptera, Geometridae)

Figure 2. Habitus of Digonis adults A Digonis aspersa (male in dorsal view) B Digonis aspersa (male brown-morpho in dorsal
view) C Digonis aspersa (male brown-morpho in ventral view) D Digonis aspersa (female in dorsal view) E Digonis cervinaria,
stat. rev. (male olivaceous-morpho in dorsal view) F Digonis cervinaria, stat. rev. (male olivaceous-morpho in ventral view)
G Digonis cervinaria, stat. rev. (male fuscous-morpho in dorsal view) H Digonis cervinaria, stat. rev. (female in dorsal view)
I Digonis cervinaria, stat. rev. (female olivaceous-morpho in ventral view) J Digonis cervinaria, stat. rev. (female brown-mor-
pho in ventral view) K Digonis punctifera (male in dorsal view; photo courtesy of A. Hausmann) L Digonis punctifera (female
in dorsal view) M Digonis gungnir Ramos-Gonzalez & Parra, sp. nov. (holotype in dorsal view) N Digonis apocrypha Ra-
mos-Gonzalez & Parra, sp. nov. (holotype in dorsal view) O Digonis apocrypha Ramos-Gonzalez & Parra, sp. nov. (holotype
in ventral view) P Phasmadigonis alba, comb. nov. (male in dorsal view) Q Gugnelve butleri Ramos-Gonzalez & Parra, sp. nov.
(holotype in dorsal view) R Gugnelve butleri Ramos-Gonzalez & Parra, sp. nov. (allotype in dorsal view). Scale bar: 10 mm.

https://inaturalist.mma.gob.cl/observations/148464516. * Concepcidn; 20-
XII-2023; -36.8263187829, -73.0265376168; observed by Antonio Maurei-
ra Navarrete and submitted to iNaturalist in: https://inaturalist.mma.gob.
cl/observations/144894518 * Concepcion; 15-XIIl-2022; -36.7784506232,

ZooKeys 1216: 173-200 (2024), DOI: 10.3897/zookeys.1216.129923 178

Mario |. Ramos-Gonzédlez et al.: Revision of the genus Digonis (Lepidoptera, Geometridae)

-73.0294558602; observed by Antonio Maureira Navarrete and submitted to
iNaturalist in: https://inaturalist.mma.gob.cl/observations/144446815. + Con-
cepcion; 20-XII-2023; -36.8263187829, -73.0265376168; observed by Antonio
Maureira Navarrete and submitted to iNaturalist in: https://inaturalist.mma.gob.
cl/observations/144894518 — Biobio Prov. + Polcura; -37.285095, -71.718547;
20-III-2022; observed by Flor Susana and submitted to iNaturalist in: https://
inaturalist.mma.gob.cl/observations/110011024. — Osorno Prov. + Osorno;
-40.565175, -73.161816; 19-VI-2021; observed by Ricardo Huenuanca and sub-
mitted to iNaturalist in: https://inaturalist.mma.gob.cl/observations/83672367
* Osorno; -40.565183, -73.161729; 02-V-2021; observed by Ricardo Huenuan-
ca and submitted to iNaturalist in: https://inaturalist.mma.gob.cl/observa-
tions/76765927 + Osorno; -40.565303, -73.162031; 09-I-2021; observed by Ri-
cardo Huenuanca and submitted to iNaturalist in: https://inaturalist.mma.gob.
cl/observations/67836032.

Diagnosis. This species can be easily distinguished from D. punctifera
(Butler), D. gungnir Ramos-Gonzalez & Parra, sp. nov., and D. cervinaria by its
brownish forewings crossed by slightly sinuous bands and bicolored spots in
the postmedial band at the level of veins R,, R,, R., M,, M,, M,, CuA,, CuA,, and
1A+2A. Externally, it differs from D. apocrypha Ramos-Gonzalez & Parra, sp. nov.
because its bicolored spots appear on both sides of both pairs of wings with an
equal proportion of white and black scales. It can be easily distinguished from
its congeners by three other genitalia characters: a tongue-shaped furca that
does not surpass the height of the transtilla, the presence of lateral spines in
the distal half of the furca, and anterior apophyses directed towards the tergum.

Redescription. Male (Fig. 2A—C). Head: antennae slightly serrate; palpi long,
one-third larger than eye diameter and slightly pointing upward; frons and vertex
covered with mottled brownish gray scales. Thorax: patagia covered with elon-
gated scales of same color as background; tegulae covered with piliform scales
of same color as background; tibial sour formula 0-2-4. Forewings: subtriangular
with acute apex and outer margin excavated between apex and M3, with a slight
mucronate extension; fovea absent; background color variable, ranging from yel-
lowish brown to dark brown with numerous small scattered blackish spots on the
surface; antemedial band dark brown to blackish, slightly arched and marked by
three bicolored spots (proximal half with whitish scales and distal half with black-
ish scales) at the level of radial, cubital, and anal veins respectively; medial band
diffuse, blackish to smoky brown, more noticeable along the costal margin; post-
medial band dark brown, slightly sinuous with the costal sector, strongly arched,
with bicolored spots (proximal half with blackish scales and distal half with whit-
ish scales) at the level of veins R,, R,, R., M,, M,, M,, CuA,, CuA,, and 1A+2A, also
visible on ventral side; subterminal band diffuse and marked only by dark spots
in subapical region, at the level of medial veins and along the anal margin; dis-
cal spot visible, punctiform, and blackish. Hindwings: subrectangular with small
mucronate apex at the level of M,; background color grayish brown; postmedial
band slightly smoky dark brown, marked by bicolored spots (proximal half with
blackish scales and distal half with whitish scales) at the level of veins, also vis-
ible on ventral surface; discal spot visible only on ventral surface. Male genita-
lia (Fig. 3A, B). Uncus conical, apically club-shaped; gnathos U-shaped with ex-
panded apex forming a pair of spiny lobes; valvae subrectangular, costa strongly
sclerotized with distal lobe before apex, crescent-shaped, slightly convex, and

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Mario |. Ramos-Gonzédlez et al.: Revision of the genus Digonis (Lepidoptera, Geometridae)

Figure 3. Genitalia of Digonis aspersa A male genitalia in ventral view B aedeagus in lateral view C female genitalia in
ventral view. Scale bar: 1 mm.

cucullus extending beyond apex of costa; transtilla truncated; saccus subrect-
angular; juxta emarginated anteriorly, with a median dimple, and furca straight,
short, does not surpass the height of the transtilla, tongue-shaped, spiny only
laterally in the distal half; anellus sclerite weak, pentagonal in shape. Aedeagus
tubular, straight; vesica without cornuti.

Female (Fig. 2D). Similar to male, but with simple antennae and well-defined
2-shaped antemedial band. Female genitalia (Fig. 3C). Ductus bursae one-sixth
the length of corpus bursae; lamella antevaginalis wide, strongly sclerotized;
corpus bursae membranous, subpyriform with an annular, hollow, strongly
denticulated signum anteriorly; posterior apophyses twice as long as anterior
apophyses, anterior apophyses slightly bent towards tergite.

Distribution. This species is found between the Chilean provinces of San
Antonio and Capitan Prat, with records in Neuquén, Argentina (Chalup 2020).
It is distributed in parts of the biogeographic provinces of Santiago, Central
Chilean subregion; Maule and Valdivian Forest, Subantarctic subregion, in the
Andean region.

Flight period. Specimens were captured or observed in January, February,
March, April, May, June, September, October, and December. There are no re-
cords for other months.

Remarks. Although the holotype stands out due to its yellowish tone, most
examined specimens exhibit a brownish coloration. There is no evidence to
suggest that this color variation reflects sexual dimorphism. The genitalia of
these brown specimens are congruent with those of the holotype, indicating
high variability in wing coloration.

ZooKeys 1216: 173-200 (2024), DOI: 10.3897/zookeys.1216.129923 180

Mario |. Ramos-Gonzédlez et al.: Revision of the genus Digonis (Lepidoptera, Geometridae)

Digonis cervinaria (Blanchard, 1852), stat. rev.
Figs 2E—J, 4

Ennomos cervinaria Blanchard, 1852: 89.

Drepanogynis eversaria Guenée, 1858: 93 (unnecessary replacement name)

Digonis cervinaria (Blanchard, 1852); Scoble 1999: 229; Pitkin 2002: 248.

Digonis cuprea Butler, 1882: 362; Bartlett-Calvert 1886: 333; Angulo and Casa-
nueva 1981: 12; Scoble 1999: 229; Pitkin 2002: 248. syn. nov.

Digonis cuprea fusca Butler, 1882: 362; Bartlett-Calvert 1886: 333; Angulo and
Casanueva 1981: 12; Scoble 1999: 229; Pitkin 2002: 248. syn. nov.

Digonis cuprea olivacea Butler, 1882: 362; Bartlett-Calvert 1886: 333; Angulo
and Casanueva 1981: 12; Scoble 1999: 229; Pitkin 2002: 248. syn. nov.

Material examined. Syntypes . CHILE * 1 female; Coquimbo; C. Gay leg.; “Digonis
cervinaria” [labeled]; MNHN * 1 female; Valparaiso; T. Edmonds leg.; “Digonis cu-
prea” [labeled]; “Chili, 82-107”; “Type” [labeled]; NHMUK * 2 males; same data as
for preceding but “var. fusca” [labeled]; “Type” [labeled] “Chili, 82-107” [labeled] and
“cuprea olivacea” [labeled]; “Type” [labeled]; “Chili, 82-107” [labeled]; all NHMUK.
Other material examined. CHILE — Valparaiso Prov. « Vina del Mar; 8-VIII-1954;
“Museo 5343” [Museum ID]; MNNC) « 1 female; Vifia del Mar; 18-IV-1953; “Museo
5018” [Museum ID]; “Mirg-015” [genitalia slide]; MNNC; * 1 male; Vifia del Mar;
15-X-1953; “Museo 5349” [Museum ID]; MNNC + 1 male; Vifia del Mar; 15-VIll-
1953; “Museo 5342” [Museum ID]; MNNC °« 1 female; Vifia del Mar; 12-IX-1953;
“Museo 5350” [Museum ID]; MNNC + 1 male; Laguna Verde; 10-X-1936; E. Ureta
leg.; “Museo 5014” [Museum ID]; MNNC. — Marga Marga Prov. > 1 male; Poza
Azul, Marga Marga; 14-XII-1953; “Museo 5346” [Museum ID]; MNNC. — Cordillera
Prov. - 1 male; La Obra; 26-III-1953; “Museo 5373” [Museum ID]; MNNC. — Cacha-
poal Prov. - 1 female; Termas de Cauquenes; 11-I-1953; E. Ureta leg.; “Museo
5006” [Museum ID]; “Mirg-013” [genitalia slide] » 1 male; Termas de Cauquenes;
11-I-1953; E. Ureta leg.; “Museo 5023” [Museum ID]; MNNC. — Talca Prov. 1 fe-
male; Panguilemo, La Calor; 1-II-2005; L. Parra leg.; “Mirg-009” [genitalia slide];
MZUC-UCCC. — Diguillin Prov. « 1 female; Las Trancas; 29-V-2011; G. Moreno leg.;
“Mirg-016” [genitalia slide]; MZUC-UCCC. — Concepcion Prov. - 1 female; Concep-
cidn; 6-I-1961; Trampas leg.; MZUC-UCCC. — Malleco Prov. - 1 female; Rio Blanco,
Curacautin; II-1995; H. Thony leg.; ZSM. — Cautin Prov. - 1 female; Termas de
Rio Blanco; IIl-1951; MZUC-UCCC. — Valdivia Prov. - 1 female; Valdivia; I|-1974;
Cameron leg.; MZUC-UCCC. — Osorno Prov. « 1 female; Puerto Octay; 11-III-1956;
Oehrens leg.; “Museo” [labeled]; MNNC. — Palena Prov. » Fjord Comau, Huinay
Station, -42.381111, -72.414722, 20 m; 8-Il-2008; A. Hausmann leg.; ZSM.
Additional records. CHILE — Itata Prov. * La Palma; -36.567339, -72.689122; 09-
VI-2021; observed by Claudio Maureira and submitted to iNaturalist in: https://inat-
uralist.mma.gob.cl/observations/85725251. — Concepcion Prov. * Concepcidn;
-36.7928173278, -73.0525010616; 13-XII-2022; observed by Antonio Maureira
Navarrete and submitted to iNaturalist in: https://inaturalist.mma.gob.cl/observa-
tions/144446817. — Cautin Prov. » Molco; -39.3393898652, -72.0921077239; 25-
XII-2022; observed by Michael Weymann and submitted to iNaturalist in: https://
inaturalist.mma.gob.cl/observations/145759642. + Molco; -39.3394188914,
-72.0921402457; 25-XII-2022; observed by Michael Weymann and submitted to
iNaturalist in: https://inaturalist.mma.gob.cl/observations/145759626. Osorno

ZooKeys 1216: 173-200 (2024), DOI: 10.3897/zookeys.1216.129923 181

Mario |. Ramos-Gonzédlez et al.: Revision of the genus Digonis (Lepidoptera, Geometridae)

Figure 4. Genitalia of Digonis cervinaria, stat. rev. A male genitalia in ventral view B aedeagus in lateral view C female
genitalia in ventral view. Scale bars: 1 mm.

Prov. * Osorno; -40.565282, -73.161883; 31-l-2021; observed by Ricardo Huen-
uanca and submitted to iNaturalist in: https://inaturalist.mma.gob.cl/observa-
tions/68898181 * Osorno; -40.565277, -73.162024; 3-I-2021; observed by Ricar-
do Huenuanca and submitted to iNaturalist in: https://inaturalist.mma.gob.cl/
observations/67871022. — Llanquihue Prov. - Las Cascadas; -41.0799870456,
-72.6350197924: 6-V-2023; observed by Mario Ramos and submitted to iNaturalist
in: https://inaturalist.mma.gob.cl/observations/164957877. — Chiloé Prov. + Pun-
tra; -42.119003, -73.81765; 27-II-2022; observed by Waldo Moyano and submitted
to iNaturalist in: https://inaturalist.mma.gob.cl/observations/10784241 6.

Diagnosis. This species can be distinguished externally from other Digonis
species by having elongated whitish spots at the level of veins R3, R4, R5, M1,
M2, M3, CuA1, CuA2, and 1A+2A in postmedial region, also visible on ventral
view. Male genitalia are characterized by the presence of a needle-shaped furca
that surpasses the height of the transtilla, with only apical spines; costa strong-
ly sclerotized with distal lobe, expanded before apex with truncated edge, and
spatulate transtilla; aedeagus with a narrow and elongated caecum, one-third
of the total length of the aedeagus.

Redescription. Male (Fig. 2E—G). Head: antennae slightly serrate; palpi short,
subequal to eye diameter, porrect; frons and vertex covered with juxtaposed
brownish scales. Thorax: patagia covered with elongated scales of same color
as background; tegulae covered with piliform scales of same color as back-
ground; tibial spur formula 0-2-4. Forewings: subtriangular with acute apex and

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Mario |. Ramos-Gonzédlez et al.: Revision of the genus Digonis (Lepidoptera, Geometridae)

outer margin excavated between apex and M,, with slight mucronate exten-
sion; fovea absent; background color variable, ranging from olive-brown to dark
brown; antemedial band dark brown to blackish, slightly sinuous; medial band
diffuse, blackish to smoky brown, slightly oblique and in contact with postme-
dial band near anal margin; postmedial band dark brown, slightly sinuous with
the costal sector, near apex, strongly arched, and with elongated whitish spots
at the level of veins R,, R,, R., M,, M,, M,, CuA,, CuA,, and 1A+2A, also visible
on ventral view; subterminal band diffuse and marked only by dark spots in
subapical region; discal spot visible, punctiform, and blackish. Hindwings: sub-
rectangular with small mucronate apex at the level of M,; background color
grayish brown; postmedial band dark brown, slightly smoky, marked by elongat-
ed whitish spots at the level of veins, only visible on ventral surface; discal spot
visible only on ventral surface. Male genitalia (Fig. 4A, B). Uncus conical, api-
cally halberd-shaped; gnathos U-shaped with an expanded apex forming a pair
of spiny lobes; valvae subrectangular, costa strongly sclerotized with a distal
lobe before the apex, truncated, slightly convex, and cucullus extending beyond
the apex of the costa; transtilla spatulate; saccus membranous; juxta emargin-
ated anteriorly, with a median dimple, and furca straight, longitudinally striated,
long, surpassing the height of the transtilla, needle-shaped, spiny only apically;
anellus sclerite weakly defined. Aedeagus tubular, straight; caecum narrow and
elongated, one-third of the total length of the aedeagus; vesica without cornuti.

Female (Fig. 2H-J). Similar to male, but with simple filiform antennae,
2-shaped antemedial band, and zigzagging subterminal band. Female genitalia
(Fig. 4C). Ductus bursae one-fourth the length of the corpus bursae; lamella
antevaginalis, sclerotized; corpus bursae membranous, subpyriform with an
annular, hollow, strongly denticulated signum anteriorly; posterior apophyses
three times longer than the anterior apophyses.

Distribution. This species is found between the Elqui and Palena provinc-
es. It is distributed in parts of the biogeographic provinces of Coquimbo and
Santiago, Central Chilean subregion; Maule and Valdivian Forest, Subantarctic
subregion, in the Andean region.

Flight period. Specimens were captured or observed in January, February,
March, April, May, June, August, September, October, and December. There are
no records for other months.

Digonis punctifera Butler, 1882
Figs 2K,L, 5

Digonis punctifera Butler, 1882: 363; Bartlett-Calvert 1886: 333; Angulo and
Casanueva 1981: 12; Pitkin 2002: 248.

Digonis punctifera acuminata Butler, 1882: 363; Bartlett-Calvert 1886: 333;
Angulo and Casanueva 1981: 12; Pitkin 2002: 248. syn. nov.

Digonis punctifera fumosa Butler, 1882: 363; Bartlett-Calvert 1886: 333; Angulo
and Casanueva 1981: 12; Pitkin 2002: 248. syn. nov.

Digonis punctifera maculosa Butler, 1882: 363; Bartlett-Calvert 1886: 333; Angu-
lo and Casanueva 1981: 12; Pitkin 2002: 248. syn. nov.

Digonis punctifera terranea Butler, 1882: 363; Bartlett-Calvert 1886: 333; Angulo
and Casanueva 1981: 12; Pitkin 2002: 248. syn. nov.

ZooKeys 1216: 173-200 (2024), DOI: 10.3897/zookeys.1216.129923 183

Mario |. Ramos-Gonzélez et al.: Revision of the genus Digonis (Lepidoptera, Geometridae)

Material examined. Syntypes. CHILE * 1 male; Valparaiso; T. Edmonds Leg;
“Type” [labeled]; “Chili, 82-107” [labeled]; NHMUK « 1 female; Valparaiso; T. Ed-
monds leg.; “var. acuminata” [labeled]; “Type” [labeled]; “Chili, 82-107” [labeled];
NHMUK * 1 male; Valparaiso; T. Edmonds leg.; “var. fumosa” [labeled]; “Type”
[labeled]; “Chili, 82-107” [labeled]; NHMUK * 2 females; same data as for pre-
ceding but “var. maculosa” [labeled]; “Type” [labeled]; “Chili, 82-107” [labeled];
all NHMUK « 1 male; same data as for preceding but “var. terranea” [labeled];
“Type” [labeled]; “Chili, 82-107” [labeled]; NHMUK.

Other material examined. CHILE — Limari Prov. * Ovalle, Quebrada Seca; 28/29-
XI-1997; ZSM * 1 female; Ovalle, Los Molles; 16/19-X-1994; ZSM. — Petorca Prov.
* 1 male and 1 female; Cachagua, Quebrada Aguas Claras; 5-IIl-1997; ZSM. —
Valparaiso Prov. - 1 female; Vifia del Mar; 4-IV-1953; “Mirg-011” [genitalia slide];
MNNC. + 3 males and 1 female; Valparaiso; 10-III-2006; H. Thoeny leg.; ZSM. —
Diguillin Prov. * 1 male; Las Trancas; 8-Il-2011; G. Moreno leg.; MZUC-UCCC.

Additional records. CHILE. — Limari Prov. - PN. Fray Jorge; -30.658155,
-71.66439; 21-VII-2016; observed by Bastian Riveros and submitted to iNatural-
ist in: https://inaturalist.mma.gob.cl/observations/87956647.

Diagnosis. Externally, D. punctifera is characterized by the presence of a sinuous
postmedial band that follows the shape of the wing’s outer margin. The female
genitalia are distinguished by a ductus bursae that is half the length of the corpus
bursae and posterior apophyses five times longer than the anterior apophyses.

Redescription. Male (Fig. 2K). Head: antennae slightly serrate; palpi long, one-
third longer than the eye diameter and slightly pointing upward; frons and vertex
covered with mottled grayish brown scales. Thorax: patagia covered with elon-
gated grayish brown and dark brown scales; tegulae covered with piliform scales
of same color as patagia; tibial spur formula 0-2-4. Forewings: subtriangular with
acute apex and outer margin excavated between apex and M,, with slight mucro-
nate extension; fovea absent; background color variable, ranging from yellowish
brown to dark brown, with numerous small blackish spots scattered on surface;
antemedial band faint, brownish dark, 2-shaped; postmedial band dark brown,
sinuous following the shape of the wing's outer margin, with a strip or whitish
points toward the distal margin, not visible on ventral surface; subterminal band
diffuse and marked only by dark spots in subapical region, at the level of medi-
al veins and on anal margin; discal spot visible, punctiform, and blackish. Hind-
wings: subrectangular with small mucronate apex at the level of M,; background
color grayish brown; postmedial band dark brown, zigzagging, slightly smoky,
and marked by blackish spots at the level of veins, also visible on ventral surface;
discal spot visible, punctiform, and blackish. Male genitalia. Unknown.

Female (Fig. 2L). Similar to male but with simple filiform antennae and more
defined antemedial band, also X-shaped. Female genitalia (Fig. 5). Ductus bur-
sae half the length of the corpus bursae; lamella antevaginalis sclerotized; cor-
pus bursae membranous, subpyriform with an annular, hollow, strongly dentic-
ulated signum anteriorly, located to the right of the corpus bursae; posterior
apophyses five times longer than the anterior apophyses.

Distribution. This species is found between the Limari and Diguillin provinces.
It is distributed in parts of the biogeographic provinces of Coquimbo and Santi-
ago, Central Chilean subregion; Maule, Subantarctic subregion, Andean region.

Flight period. Specimens were captured or observed in February, March,
April, July, October, and November. There are no records for other months.

ZooKeys 1216: 173-200 (2024), DOI: 10.3897/zookeys.1216.129923 184

Mario |. Ramos-Gonzédlez et al.: Revision of the genus Digonis (Lepidoptera, Geometridae)

Figure 5. Female genitalia of Digonis punctifera. Scale bar: 1 mm.

Digonis gungnir Ramos-Gonzalez & Parra, sp. nov.
https://zoobank.org/2BE6459A-9B57-400B-91F1-844437B57E79
Figs 2M, 6

Type material. Holotype. CHILE — 1 male; pinned; Coquimbo, Elqui, Huanta;
1936 (year without more data); E. Ureta leg.; “Holotype Digonis gungnir” [red
handwritten label]; “5024” [Museum ID]; “Mirg-019” [genitalia slide]; MNNC.
Paratypes. CHILE — 1 male; pinned; Llanquihue Prov., Maullin; Il-1943; S. Bar-
ros leg.; MZUC-UCCC * 1 male; pinned; Magallanes Prov., Punta Arenas, Tres
Puentes; XII-1952; n.n. leg.; MZUC-UCCC.

Diagnosis. Externally, D. gungnir Ramos-Gonzalez & Parra, sp. nov. is charac-
terized by straw-colored wings and a straight postmedial band with pale points
bordered in dark brown at veins R,, R., M,, M,, and M, on the forewings. Male
genitalia feature a presence of a poorly defined juxta, a slightly arched spear-
shaped furca, and dense dorsal spines on the furca.

Description. Male (Fig. 2M). Head: antennae slightly serrate; palpi long, one-
third longer than the eye diameter, porrect; frons and vertex covered with juxta-
posed whitish scales. Thorax: patagia covered with elongated whitish scales;
tegulae covered with very pale yellowish piliform scales; tibial spur formula

ZooKeys 1216: 173-200 (2024), DOI: 10.3897/zookeys.1216.129923 185

Mario |. Ramos-Gonzédlez et al.: Revision of the genus Digonis (Lepidoptera, Geometridae)

A

Figure 6. Genitalia of Digonis gungnir Ramos-Gonzalez & Parra, sp. nov. (holotype)
A male genitalia in ventral view B aedeagus in lateral view. Scale bar: 1 mm.

0-2-4. Forewings: subtriangular with acute apex and outer margin excavated
between apex and M., with slight mucronate extension; fovea absent; back-
ground color straw-colored; antemedial band diffuse, marked by three light
brown points at the level of radial, cubital, and anal veins respectively; medial
band diffuse, light brown, slightly arched and more noticeable between the dis-
cal spot and the costa, zigzagging to the anal margin; postmedial band straight,
diffuse, light brown, with five white points bordered in dark brown at the level
of veins R,, R,, M., M., and M.,, visible only on ventral surface; subterminal band
diffuse and demarcated only by dark spots in subapical region, at the level of
medial veins and on anal margin; discal spot visible, punctiform, and blackish.
Hindwings: subrectangular with small mucronate apex at the level of M,; back-
ground color pale ashy; postmedial band light brown, slightly smoky, marked by
elongated light brown spots at the level of veins, only visible on ventral surface;
discal spot visible. Male genitalia (Fig. 6). Uncus conical, apex club-shaped;
gnathos U-shaped with an expanded apex forming a pair of spinous lobes;
valvae subrectangular, costa strongly sclerotized with a distal lobe before the
apex, rounded, and cucullus extended beyond the apex of the costa; transtilla
bifid; saccus subrounded; juxta poorly defined, pointed anteriorly, with a central
depression, and with a furca curved to the left, long, surpassing the height of
the transtilla, slightly arched spear-shaped, densely spiny dorsally, apex round-
ed; anellus sclerite weakly defined, only two subtriangular sclerites near the
base of the furca are visible. Aedeagus tubular, straight; vesica without cornuti.

ZooKeys 1216: 173-200 (2024), DOI: 10.3897/zookeys.1216.129923 186

Mario |. Ramos-Gonzédlez et al.: Revision of the genus Digonis (Lepidoptera, Geometridae)

Female. Unknown.

Etymology. The species name is a noun in apposition, referring to Odin’s
spear (the chief god in Norse mythology), due to its longer and more armed
furca within the genus. Gungnir is treated here as a neuter noun.

Distribution. This species is found between the provinces of Elqui and Ma-
gallanes. It is distributed in parts of the biogeographic provinces of Coquimbo
and Santiago, Central Chilean subregion; Maule, Valdivian Forest, and Magel-
lanic Forest, Subantarctic subregion, Andean region.

Flight period. Specimens were captured in December and February. There
are no records for other months.

Digonis apocrypha Ramos-Gonzalez & Parra, sp. nov.
https://zoobank.org/458C9CE8-3466-4EEE-99A0-35F4E244557C
Figs 2N, O, 7

Type material. Holotype. CHILE: 1 male; pinned; Nuble, Las Trancas; 31-III-2011; G.
Moreno leg.; “Holotype Digonis apocrypha” [red handwritten label], “UCCC_MZUC_
Lep_0300” [Museum ID], “Mirg-021” [genitalia slide] (MZUC-UCCC). Paratypes.
CHILE — Diguillin Prov. - 1 male; pinned; Nuble, Recinto; 06-II-2011; G. Moreno
leg.; MZUC-UCCC + 1 male; pinned; same data as holotype but “UCCC_MZUC_
Lep_0304” [Museum ID] (MZUC-UCCC) « 2 males; pinned; same data as holotype
but 12-IV-2013; “UCCC_MZUC_Lep_1729” and “UCCC_MZUC_Lep_1730” [Museum
ID]; MZUC-UCCC + 2 males; pinned; same data as holotype but 20-III-2011; “UCCC_
MZUC_Lep_0330” and “UCCC_MZUC_Lep_0320” [Museum ID]; MZUC-UCCC = 1
male; pinned; same data as holotype but 16-IV-2010; “UCCC_MZUC_Lep_0303”
[Museum ID]; MZUC-UCCC « 2 males; pinned; same data as holotype but 7-IV-2010;
MZUC-UCCC 1 male; pinned; same data as holotype but “UCCC_MZUC_Lep_0301”
[Museum ID]; MZUC-UCCC = 1 male; pinned; same data as holotype but 29-V-2011;
“UCCC_MZUC_Lep_0302” [Museum ID]; MZUC-UCCC « 1 male; pinned; same data
as holotype but 18-IIl-2012; “UCCC_MZUC_Lep_1160” [Museum ID]; MZUC-UC-
CC. — Valdivia Prov. +» 1 male; pinned; 09-IV-2010; Huilo-Huilo, Mocho-Choshuen-
co volcano; -39.911943; -71.969167; L. Roa and D. Vergara leg.; MZUC-UCCC. —
Capitan Prat Prov. + 1 male; pinned; Cochrane; 11-IV-2008; Parra and Alvarado leg.;
“sampled in scrub of Notro-Nirre” [labeled] (Embothrium coccineum (Proteaceae)
- Nothofagus antarctica (Nothofagaceae)); MZUC-UCCC.

Diagnosis. This species is distinguished from the other species in the genus
by the presence of elongated blackish marks with small whitish point at the
level of veins in the postmedial band of the forewings, which are not visible
in ventral view. In the male genitalia, it differs from D. aspersa by the elongat-
ed and striated furca, more similar to that of D. cervinaria. It also differs from
D. cervinaria because it has spines around the distal half of the furca. This spe-
cies stands out for the presence of a U-shaped gnathos with an expanded apex
like a denticulate plate, transtilla with a digitiform process at the apex, and ae-
deagus with a fine and elongated caecum, half the total length of the aedeagus.

Description. Male (Fig. 2N, 0). Head: antennae slightly serrate; palpi long, one-
third longer than the eye diameter, porrect; frons and vertex covered with grayish
brown scales. Thorax: patagia covered with elongated scales of same color as
background; tegulae covered with piliform scales of same color as background;

ZooKeys 1216: 173-200 (2024), DOI: 10.3897/zookeys.1216.129923 187

Mario |. Ramos-Gonzadlez et al.: Revision of the genus Digonis (Lepidoptera, Geometridae)

5

that
a

B

Figure 7. Genitalia of Digonis apocrypha Ramos-Gonzalez & Parra, sp. nov. (holotype)
A male genitalia in ventral view B aedeagus in lateral view. Scale bar: 1 mm.

tibial sour formula 0-2-4. Forewings: subtriangular with acute apex and outer mar-
gin excavated between apex and M., with slight mucronate extension; fovea ab-
sent; pale ashy background with a large number of small blackish spots scattered
over the surface; antemedial band diffuse, slightly zigzagging, marked by three dark
brown points at the level of radial, cubital, and anal veins respectively; medial band
light brown, diffuse; postmedial band, diffuse, light brown, with bicolored spots
(elongated blackish mark with the center marked by a whitish point) at the level
of veins R,, R,, M,, M,, M,, CuA,, CuA,, and 1A+2A, not visible on ventral surface.
Hindwings: subrectangular with small mucronate apex at the level of M,; pale ashy
background, without visible bands or spots; discal spot visible. Male genitalia (Fig.
7). Uncus conical, apex club-shaped; gnathos U-shaped with an expanded apex like
a denticulate plate; valvae subrectangular, costa strongly sclerotized with a distal
lobe before the apex, rounded, and cucullus extended beyond the apex of the cos-
ta; transtilla bifid; saccus membranous, weakly defined; juxta poorly defined, with
a central depression and furca, long, surpassing the height of the gnathos, club-
shaped with two longitudinal stripes, densely spiny towards the distal half; anellus
sclerite weakly defined. Aedeagus tubular, straight; fine, elongated caecum, half
the total length of the aedeagus; vesica without cornuti.

Female. Unknown.

Etymology. The species name is an adjective from Greek aprocryphos (“not
genuine”), referring to the deceptive maculation pattern, superficially resem-
bling Digonis aspersa Butler.

ZooKeys 1216: 173-200 (2024), DOI: 10.3897/zookeys.1216.129923 188

Mario |. Ramos-Gonzdlez et al.: Revision of the genus Digonis (Lepidoptera, Geometridae)

Distribution. This species is found between the provinces of Diguillin and
Capitan Prat. It is distributed in parts of the biogeographic provinces of Maule,
Valdivian Forest, Magellanic Forest, Subantarctic subregion, in the Andean region.

Flight period. Specimens were captured in February, March, April, and May.
There are no records for other months.

Phasmadigonis gen. nov.
https://zoobank.org/F092281 5-E334-4247-9CE0-9F98FDODABEO

Type species. Digonis alba Butler, 1882.

Diagnosis. Phasmadigonis bears resemblance to Digonodes Warren, 1895, Di-
gonis Butler, 1882, and Gonogala Butler, 1882, particularly due to the mucronate
shape of the wings. However, Phasmadigonis can be distinguished by the pres-
ence of vein Sc connected by a vein to the single accessory cell in the forewings
and by having vein Sc+R1 connected to radial trunk by a weak vein in the hind-
wings. Phasmadigonis is distinguished by the following genitalia characters: gna-
thos V-shaped with the absence of lobes or spines, spatulate transtilla, broad shov-
el-shaped juxta, furca armed with small sagittal spines and a dimple in the sclerite
at its base, aedeagus with a digitiform apex, and vesica armed with a large spine.

Description. Antennae serrated in males and filiform in females. Thorax and
abdomen with grayish scales. Forewings gray-lilac reticulated with white, lack-
ing bands, subterminal region delimited only by a series of blackish spots at the
level of R2, R3, R4, M1, M2, M3, CuA1, CuA2, and 1A+2A; costa margin in apical
zone is slightly arched; outer margin is concave between apex and M3. Wing
venation (Fig. 8): one accessory cell; Sc connected with accessory cell through
a short vein, R1 and R2 arise from accessory cell, R3 and R4 are pedunculate,
R5 terminates at termen; M2 equidistant from M1 and M3, M3 slightly arched
and ending in small mucronate apex; CuA1 arises 1/10 before end of cell, CuA2
arises near middle of cell. Hindwings paler than forewings, with subterminal
region delimited by series of dark spots at the level of veins. Sc+R1 connected
to radial trunk by a weak transverse vein, Rs arises 1/6 before end of cell, M2 is
absent. Male genitalia with rod-like uncus; gnathos V-shaped; valvae suboval;
transtilla spatulate; furca spiny; vesica with prominent spine.

Etymology. The generic name is formed by combining the Greek phasma
(meaning phantom or apparition), with Digonis in reference to the false re-
semblance to moths of the genus Digonis Butler and the wing coloration; its
gender is neuter.

Distribution. Similar to its sole species, P. alba (Butler)

Phasmadigonis alba (Butler, 1882), comb. nov.
Figs 2P 9

Digonis alba Butler, 1882: 361; Bartlett-Calvert 1886: 333; Angulo and Casanue-
va 1981: 12; Scoble 1999: 229; Pitkin 2002: 248.

Material examined. Syntype. CHILE * 1 female; pinned; Mountains of the hacienda
of Cauquenes; T. Edmonds leg.; “Chili, 82-107” [labeled]; “Type” [labeled]; NHMUK.

ZooKeys 1216: 173-200 (2024), DOI: 10.3897/zookeys.1216.129923 189

Mario |. Ramos-Gonzadlez et al.: Revision of the genus Digonis (Lepidoptera, Geometridae)

GEA

Figure 8. Wing venation of Phasmadigonis Ramos-Gonzalez & Parra, gen. nov. Scale bar:
10 mm.

Other material examined. CHILE — ° 1 male; Chile; “Museo 5022” [Museum
ID]; MNNC. — Diguillin Prov. * 1 male; Nuble, Las Trancas; 18-III-2012; G. More-
no leg.; “UCCC-MUZC-Lep1173” [Museum ID]; “MMA1173” [genitalia slide];
MZUC-UCCC. — Araucania Region : 1 male; Araucania, II-1888, coll. n.n, “Museo
5029” [Museum ID]; MNNC; * 2 males; same data as for precedings but “Museo
5027” [Museum ID] and “Museo 5028” [Museum ID]; all MNNC. — Chiloé Prov.
* 1 male; Mocopulli, Ruta 5 Sur km 1170, -42.368000, -73.728833, 182 m; 3-ll-
2017, leg. M. Ramos-G, M. Ramos-SM & C. Rose.

ARGENTINA — * 1 male; Rio Negro Prov., Bariloche; 29-I-1991; H. Ibarra-Vidal
leg.; “HIV-0034” [Museum ID]; MHNC.

Additional records. CHILE. — Talca Prov. - Cipreses hydroelectric plant;
-35.7867249, -70.8078157; 16-XII-2023; observed by César Picar and submit-
ted to iNaturalist in: https://inaturalist.mma.gob.cl/observations/196336300. *
Cipreses hydroelectric plant; -35.7865833685, -70.8078496903; 21-I-2024: ob-
served by César Picar and submitted to iNaturalist in: https://inaturalist.mma.
gob.cl/observations/197310545.

Diagnosis. As for the genus.

Redescription. Male (Fig. 2P). Head: antennae slightly serrated; short pal-
pi, subequal to eye diameter, porrect; frons and vertex covered with juxtaposed
brownish scales. Thorax: patagia covered with elongated scales of same color as
background; tegulae covered with piliform scales of same color as background;
tibial spur formula 0-2-4. Forewings: subtriangular with acute apex and outer

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Mario |. Ramos-Gonzdlez et al.: Revision of the genus Digonis (Lepidoptera, Geometridae)

Figure 9. Genitalia of Phasmadigonis alba, comb. nov. A male genitalia in ventral view
B aedeagus in lateral view. Scale bar: 1 mm.

margin excavated between apex and M,, with a slight mucronated extension; fovea
absent; ground color gray-lilac reticulated with white; bands absent, only observ-
able on wing surface are blackish points in subterminal area at the level of veins
R,,R,,R,, M,,M,,M,, CuA,, CuA,, and 1A+2A, also visible on ventral side; discal spot
visible, punctiform, and blackish. Hindwings: subrectangular with small mucronat-
ed apex at the level of M,; ground color ash-white, termen grayish; bands absent,
only observable on wing surface is a row of blackish dots in subterminal area at the
level of veins Sc+R,, Rs, M,,M,, CuA,, CuA,, and 1A; discal spot visible. Male genita-
lia (Fig. 9). Uncus straight, apex rod-like; gnathos V-shaped; valvae suboval, valvula
and cucullus slightly setose, costa sclerotized; transtilla spatulate with sharp, pro-
jected vertices; saccus obcordate; juxta shovel-shaped, dorsally flattened and ex-
tended, with finger-shaped furca curved to the left, short, not surpassing the height
of the transtilla,, densely spiny sagittally; anellus sclerite weakly defined. Aedeagus
tubular, straight, apex digitiform; vesica armed with a large spine.

Female. Unknown.

Distribution. This species is found between the provinces of Cachapoal and
Chiloé (Chile) and Rio Negro (Argentina). It is distributed in parts of the biogeo-
graphic provinces of Santiago, Central Chile subregion; Maule and Valdivian
Forest, Subantarctic subregion, in the Andean region.

Flight period. Specimens were captured in December, January, February, and
March. There are no records for other months.

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Tribe Nacophorini Forbes, 1948

Gugnelve gen. nov.
https://zoobank.org/5D5F1234-1907-4D23-A970-C051EF3D8DEA

Type species. Gugnelve butleri Ramos-Gonzalez & Parra, sp. nov.

Diagnosis. Externally, Gugnelve resembles Euangerona Butler and Dectochi-
lus Warren. All three genera have an oblique curved band in the central area of
the forewings. However, Euangerona and Dectochilus have a wavy termen, un-
like Gugnelve, which has a smooth termen and a slightly falcate apex. The wing
shape and antennae are similar to those of Laninia Orfila & Schajovskoy and
Macrolyrcea Butler, but they differ significantly in various genital structures,
such as the gnathos with a posteriorly directed apex and distally lobulated
costa in Laninia and the spatulate uncus and wide oval valvae in Macrolyr-
cea. The V-shaped gnathos with a longitudinal row of spicules is reminiscent
of Euangerona and Dectochilus, but in both of those genera, the anellus pro-
cess is directed anteriorly, while in Gugnelve, it is oriented lateroposteriorly.
The uncus and juxta resemble those of Malleco Rindge, but in Gugnelve, the
uncus is glabrous, and the anellus process is trifid, while the gnathos of Malle-
co has multiple spicules on the lateral arms and apex. Finally, the new genus
differs considerably in external characteristics and genitalia from other Ande-
an Nacophorini like Catophoenissa Warren, 1894, Catocalopsis Rindge, 1971,
Talca Rindge, 1971, and Wichanraran Parra, 2018. Gugnelve is distinguished
by the following combination of genital characters: V-shaped gnathos with a
row of 11 spicules at the apex, forked transtilla, halberd-shaped juxta laterally
extended into a pair of tri-spined anellus processes, strongly sclerotized and
glabrous, vesica armed with a series of cornuti, Signum with a long blade-like
ridge on a sclerotized patch.

Description. Antennae thickened and slightly serrated in males, filiform in
females. Robust thorax with yellowish brown piliform scales. Abdomen with
belt of white scales on Al and A2. Forewings yellowish brown, subtriangular,
slightly falcate, with sinuous and inclined antemedial and postmedial bands.
Wing venation (Fig. 10). Two accessory cells, second twice as long as first;
Sc in contact with first accessory cell, R1 originates from middle of second
accessory cell, R2 and R3+4 connate, R3 and R4 pedunculate, R5 terminates at
termen; M2 equidistant from M1 and M3, M3 slightly arched; CuA1 originates
1/8 before end of cell, CuA2 originates near middle of cell. Hindwings paler
than forewings, with yellowish brown piliform scales on the termen. Sc+R1 in
contact with radial trunk up to middle of cell, Rs originates 1/12 before end of
cell, M2 absent. Male genitalia with V-shaped gnathos, expanded apex with
11 small variably-sized denticles towards the center; valvae sub-rhomboidal;
juxta sub-halberd-shaped, laterally extended into a pair of tri-spined, strongly
sclerotized processes. Female genitalia with signum with long blade-like ridge
on sclerotized patch.

Etymology. The generic name is derived from the Mapudungun language
(spoken by the Mapuche, the largest indigenous group in Chile), winyelfe,
meaning bright star or Venus, in reference to its yellowish coloration; its
gender is neuter.

Distribution. As for its only species, G. butleriRamos-Gonzalez & Parra, sp. nov.

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Mario |. Ramos-Gonzdlez et al.: Revision of the genus Digonis (Lepidoptera, Geometridae)

Figure 10. Wing venation of Gugnelve Ramos-Gonzalez & Parra, gen. nov. Scale bar: 10 mm.

Gugnelve butleri Ramos-Gonzalez & Parra, sp. nov.
https://Zzoobank.org/80882365-C942-46DB-AAD3-FC68E5CE32D8
Figs 2Q, R, 11

Erosina cervinaria (Blanchard, 1852); Butler 1882: 347 pl. 16, fig. 4; Bart-
lett-Calvert 1886: 331; Angulo and Casanueva 1981: 12 [misidentification of
Ennomos cervinaria Blanchard].

Type material. Holotype. CHILE * 1 male; Cordillera Prov., Santiago, Guayacan;
XII-1950; “Holotype Gugnelve butleri” [red handwritten label], “Mirg-017” [geni-
talia slide] (MZUC-UCCC). Allotype. Chile + 1 female, Cordillera Prov., Santiago,
Guayacan; I-1951; “Allotype Gugnelve butleri” [red handwritten label], “Mirg-018”
[genitalia slide] (MZUC-UCCC). Paratypes. CHILE — Limari Prov. + 2 males, Oval-
le, Quebrada Seca; 28/29-XI-1997; ZSM. — Cordillera Prov. - 4 females and 1
male, same data as Holotype; * 1 female and 1 male, same data as allotype.

Other material examined. CHILE — Cachapoal Prov. - 1 female; Termas de
Cauquenes; 11-Il-1953; “Museo 4516” [Museum ID]; MNNC.

Additional records. CHILE. — Cachapoal Prov. - Mountains of the hacienda
of Cauquenes; January; T. Edmonds leg.; NHMUK (Butler, 1882) — Talca Prov.
- Parque Natural Tricahue; -35.70869903, -71.08727243; 5-l-2020; observed by
Vicente Pantoja and submitted to iNaturalist in: https://inaturalist.mma.gob.
cl/observations/37300513. * Cipreses hydroelectric plant; -35.8099272997,
-70.8359745; 17-XIl-2022; observed by César Picar and submitted to iNaturalist

ZooKeys 1216: 173-200 (2024), DOI: 10.3897/zookeys.1216.129923 193

Mario |. Ramos-Gonzédlez et al.: Revision of the genus Digonis (Lepidoptera, Geometridae)

Figure 11. Genitalia of Gugnelve butleri Ramos-Gonzalez & Parra, sp. nov. (holotype and allotype) A male genitalia in
ventral view B aedeagus in lateral view C female genitalia in ventral view. Scale bars: 1 mm.

in: https://iNaturalist.mma.gob.cl/observations/144651054. + Cipreses hy-
droelectric plant; -35.8096577, -70.8353316; 10-I-2024; observed by César
Picar and submitted to iNaturalist in: https://inaturalist.mma.gob.cl/observa-
tions/196337433.

Diagnosis. As for the genus.

Description. Male (Fig. 2Q). Head: antennae slightly serrated; palpi short,
subequal to the eye diameter, porrect; frons and vertex covered with juxtaposed
brownish scales. Thorax: Patagia covered with elongated scales of same color
as background; tegulae covered with piliform scales of same color as back-
ground; tibial spur formula 0-2-4. Forewings: slightly falcate, subtriangular; fo-
vea absent; background color yellowish brown; antemedial band dark yellowish
brown, externally bordered by pale brown, oblique, extending toward the medial
area, near the end of the discal cell where it curves toward the antemedial sec-
tor to the costa, forming an acute arc; postmedial band dark yellowish brown,
externally bordered by pale brown, slightly sinuous, oblique, and slightly curved,
extending to the subapical sector; discal spot not visible. Hindwings: suboval;
background color pale brown without bands; discal spot not visible. Male gen-
italia (Fig. 11A, B). Uncus conical; socii slightly setose; gnathos V-shaped with
sinuous arms, expanded apex with 11 small denticles of varying sizes toward
the center; valvae sub-rhomboidal, costa and anterior margin sclerotized; trans-
tilla fork-shaped; saccus suboval; juxta sub-halberd-shaped, laterally extended
into a pair of tri-spined strongly sclerotized processes; anellus sclerite weakly
defined. Tubular aedeagus, straight; vesica armed with cornuti, formed by 11
grouped spines of different sizes.

Female (Fig. 2R). Similar to male but with simple filiform antennae, dark
yellowish brown falcate forewings, and slightly crenulated pale brownish gray
subrectangular hindwings. Female genitalia (Fig. 11C). Ductus bursae half the

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Mario |. Ramos-Gonzadlez et al.: Revision of the genus Digonis (Lepidoptera, Geometridae)

length of the corpus bursae; antrum funnel-like; corpus bursae membranous,
subpyriform, signum with a long blade-like ridge on a sclerotized patch; poste-
rior apophyses three times longer than anterior apophyses.

Etymology. The specific name is dedicated to Arthur Gardiner Butler, an orni-
thologist, arachnologist, and entomologist of the 19" century, for his contribu-
tion to the study of Lepidoptera in Chile.

Distribution. This species is found between the provinces of Limari and
Talca. It is distributed in parts of the biogeographic provinces of Coquimbo
and Santiago, Central Chilean subregion; Maule, Subantarctic subregion, in the
Andean region.

Flight period. Specimens were captured in November, December, and Janu-
ary. There are no records for other months.

Discussion

The genus Digonis, described by Butler in 1882, initially included D. asper-
sa, D. cuprea, D. punctifera, and D. alba. Additionally, Butler (1882) described
various varieties for cuprea and punctifera, differentiating them based on the
coloration of the forewings. However, the analysis of the genital structures of
D. punctifera indicates that these differences in coloration are not sufficient to
separate them into distinct species or subspecies within the genus, which is
why they are synonymized.

The inclusion of the species D. gungnir Ramos-Gonzalez & Parra, sp. nov. and
D. apocrypha Ramos-Gonzalez & Parra, sp. nov. in the genus Digonis is justified
by the congruence in their external characteristics and genitalia. The distinction
from their sister species is primarily focused on male genitalia characteristics.
Therefore, the genus Digonis is composed of the following five species: D. as-
persa, D. cervinaria, D. punctifera, D. gungnir Ramos-Gonzalez & Parra, sp. nov.,
and D. apocrypha Ramos-Gonzalez & Parra, sp. nov. Its validity as a taxon within
the tribe Ennomini is supported by the presence of a membranous and weakly
developed saccus, valvae with well-sclerotized outstanding costa, and well-de-
veloped furca (Beljaev 2008).

Based on the analysis of genital structures and wing venation, a strong in-
congruence is recognized between the species D. a/ba Butler and the rest of
the species in Digonis. This aligns with what Pitkin (2002) indicated, suggesting
that D. alba should be excluded from the genus due to certain incongruences
with genital characters. The main differences between both genera lie in the
shape of the valvae, furca, and wing venation configuration, a combination of
characters not present in any other Ennomini genus in the Andean region (e.g.,
Syncirsodes Butler, Gonogala Butler, Microclysia Butler, Perusia Herrich-Schaffer,
Eusarca Hiibner), which supports the proposal of Phasmadigonis as a new ge-
nus. Additionally, we report the first record of this taxon for Argentina, specifi-
cally for the province of Rio Negro.

The different species of the genus Digonis have distributions that cover
most of the vegetation formations (sensu Luebert and Pliscoff 2018) in cen-
tral-southern Chile. However, the highest species richness appears to be as-
sociated with the high-Andean deciduous forest of the Cordillera de Chillan
(sensu Gajardo 1994). The lack of records in other areas may be due to a lack
of comprehensive sampling, particularly in the central-northern zone of Chile

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Mario |. Ramos-Gonzédlez et al.: Revision of the genus Digonis (Lepidoptera, Geometridae)

(north of Santiago). Future studies should address the rearing of different spe-
cies to describe their host plants and immature stages, currently unknown.

Blanchard (1852) described the species Ennomos cervinaria based on a spec-
imen from Coquimbo, providing a brief and imprecise description, and without
including figures. Later, Guenée (1858) redescribed it in more detail, character-
izing it by the elongated whitish marks on the wing veins, visible in both dorsal
and ventral views. Furthermore, Guenée proposed a new name for this species,
arguing that “cervinaria” was already used for a species of Eubolia Duponchel,
1829 (current Larentia clavaria Duponchel, 1845). Subsequently, Butler (1882)
incorporated this species into the genus Erosina Guenée, 1858, arguing that,
if his determination of the specimen used is correct, it has a second oblique
but indistinct angular line from the costa to the inner margin, almost parallel
to the line described by Blanchard (1852) in his description. Additionally, Butler
(1882) added a sketch of the species for easier future determination. Finally,
Scoble (1999) included it in the genus Digonis. However, the analysis of the
type material reveals that the specimens determined as Ennomos cervinaria
by Butler (1882) and subsequent authors are not congruent with the holotype
of Ennomos cervinaria Blanchard. The external characteristics and genitalia of
Ennomos cervinaria correspond to those of Digonis cuprea and its varieties,
so these are synonymized.

The analysis of specimens incorrectly determined as E. cervinaria by But-
ler reflects a morphological correspondence with the concept of “Nacophori-
ni” (sensu Rindge 1973, 1983), which brought together the Ennominae with
a broad and “hairy” thorax that now form parts of the tribes Euangeronini,
Odontoperini, and Nacophorini in part (Brehm et al. 2019). The most char-
acteristic feature in the male genitalia corresponds to the row of spicules in
the sagittal plane of the gnathos and the trifid, strongly sclerotized postero-
laterally directed process of the anellus. This form and configuration of the
gnathos are present in the genera Euangerona Butler and Dectochilus Butler;
however, the process of the anellus is not congruent with both genera or
any other Euangeronini, which show a process of the anellus pointing an-
teriorly and a highly developed setose socius (possible synapomorphies of
the tribe). On the other hand, it shares with Mallomus Blanchard the spine-
shaped form of the anellus process; however, in Mallomus, it appears as a
single sclerotized spine, and the valvae have various modifications absent
in Gugnelve gen. nov. such as a membranous harpe and digitiform process-
es on the cucullus. It was also not possible to assign this species to any of
the Nacophorini genera in southern South America treated by Rindge (1973,
1983), Pitkin (2002), Parra and Pascual-Toca (2003), or Parra (2018). For this
reason, Gugnelve butleri Ramos-Gonzalez & Parra, gen. et sp. nov. is provi-
sionally included in the Nacophorini.

Finally, this revision will serve as the basis for future molecular phylogenetic
analyses that will help clarify the evolutionary relationships among species and
related taxa. With this work, the number of Ennomini species present in Chile
increases to 36 (Parra et al. 2009; Bocaz et al. 2016; Brehm et al. 2019), and the
Nacophorini to 20 (Pitkin 2002; Brehm et al. 2019). These numbers are likely
to increase in the future, considering that there have been few taxonomic revi-
sions of Ennominae in the Andean region so far.

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Mario |. Ramos-Gonzadlez et al.: Revision of the genus Digonis (Lepidoptera, Geometridae)

Acknowledgements

We would like to express our gratitude to M.J. Scoble and L. Pitkin (NHMUk)
for their assistance during L.E. Parra’s visit to the Natural History Museum.
We also thank the staff at MHNC for their help during M.R.-G.’s visits. Our
thanks extend to Joél Minet (MNHN) and Mario Elgueta (MNNC) for their
cooperation in allowing the study of specimens deposited in their institu-
tions. We are grateful to Axel Hausmann (ZSM) for sharing data and per-
mitting the use of photographs of some specimens from the collection. We
also acknowledge the support of the staff at the Museum of Zoology at the
Universidad de Concepcion, Chile (MZUC-UCCC) during the study of spec-
imens. Additionally, we extend our sincere gratitude to Claudio Maureira,
Flor Susana, Luis Chavarriga, Fpizarro, Antonio Maureira Navarrete, Ricardo
Huenuanca, Michael Weymann, and Waldo Moyano for their contributions to
the iNaturalist platform, particularly in observations of Digonis species. We
extend our gratitude to Fundacién Mongen (Chile) for sponsoring and sup-
porting the field campaign in Las Cascadas (Los Lagos, Chile) during May
2023, which enabled the recording of a Digonis species. Thanks to Carolina
Rose and Mario Ramos San Martin who collaborated in the field collection
of specimens in Chiloé. M.R.-G. and C.Z.-M. acknowledge financial support
from the “Beca Doctorado Nacional” ANID No. 21210517 and CONICYT No.
21161423, respectively.

Additional information

Conflict of interest

The authors have declared that no competing interests exist.

Ethical statement

No ethical statement was reported.

Funding

This research was supported by the Vicerectoria de Investigaci6n y Desarrollo at the
Universidad de Concepcion (VRID 214.113.087-1.0).

Author contributions

Conceptualization: LER MFVG, MIRG. Data curation: MIRG, MFVG, CZM. Funding ac-
quisition: LEP. Investigation: MIRG. Methodology: MIRG. Project administration: MIRG.
Resources: MFVG. Supervision: LEP. Visualization: CZM, MIRG. Writing - original draft:
MIRG, CZM, MFVG, LEP. Writing - review and editing: CZM, MFVG, MIRG, LEP.

Author ORCIDs

Mario |. Ramos-Gonzalez © https://orcid.org/0000-0002-5385-6836
Carlos Zamora-Manzur ® https://orcid.org/0000-0001-7755-4077
Luis E. Parra © https://orcid.org/0000-0002-3271-0037

Data availability

All of the data that support the findings of this study are available in the main text.

ZooKeys 1216: 173-200 (2024), DOI: 10.3897/zookeys.1216.129923 197

Mario |. Ramos-Gonzédlez et al.: Revision of the genus Digonis (Lepidoptera, Geometridae)

References

Angulo AO, Casanueva ME (1981) Catalogo de los lepiddpteros geométridos de Chile
(Lepidoptera: Geometridae). Boletin de la Sociedad de Biologia de Concepcion
(Chile) 51: 7-34.

Bartlett-Calvert W (1886) Catalogo de los lepiddpteros Rhopaléceros i Heterdéceros de
Chile. Anales de la Universidad de Chile 69: 314-352.

Beljaev EA (2006) A morphological approach to the Ennominae phylogeny (Lepidoptera,
Geometridae). Spixiana 29(3): 215-216.

Beljaev EA (2008) A new concept of the generic composition of the geometrid moth
tribe Ennomini (Lepidoptera, Geometridae) based on functional morphology of
the male genitalia. Entomological Review 88(1): 50-60. https://doi.org/10.1134/
S0013873808010089

Blanchard E (1852) Fauna Chilena. Insectos. Orden VI. Lepiddpteros. In: Gay C (Ed.) His-
toria fisica y politica de Chile. Imprenta de Maulde et Renou, Paris, 1-112.

Bocaz P Parra LE (2005) Revision y bionomia del género Syncirsodes Butler, 1882 (Lep-
idoptera: Geometridae). Revista Chilena de Historia Natural 78: 89-111. https://doi.
org/10.4067/S071 6-078X2005000100007

Bocaz P, Sepulveda-Zuniga E, Zamora-Manzur C, Parra LE (2016) Filogenia del género
Syncirsodes (Lepidoptera: Geometridae). Revista Colombiana de Entomologia 42(1):
75-80. https://doi.org/10.25100/socolen.v42i1.6673

Brehm G, Murillo-Ramos L, Sihvonen P Hausmann A, Schmidt BC, Ounap E, Moser A,
Mortter R, Bolt D, Bodner F, Lindt A, Parra LE, Wahlberg N (2019) New World geo-
metrid moths (Lepidoptera: Geometridae): Molecular phylogeny, biogeography, taxo-
nomic updates and description of 11 new tribes. Arthropod Systematics & Phylogeny
77(3): 457-486. https://doi.org/10.26049/ASP77-3-2019-5

Butler AG (1882) Heterocerous Lepidoptera collected in Chili by Tomas Edmonds, Esq.
Part IlI-Geometrites. Transactions of the Entomological Society of London 30(3):
339-427. https://doi.org/10.1111/j.1365-2311.1882.tb01580.x

Chalup A (2020) The biodiversity of the family Geometridae for Argentina. In: Jufent
SA, Claps LE, Morrone JJ (Eds) Biodiversidad de artrépodos argentinos. Vol. 4.
INSUE-UNT Ediciones, San Miguel de Tucuman, Argentina, 47-62.

Gajardo R (1994) La vegetacion natural de Chile: Clasificacion y distribucién geografica.
Editorial Universitaria, Santiago, Chile, 165 pp.

Gaston KJ, Scoble MJ, Crook A (1995) Patterns in species description: a case study
using the Geometridae (Lepidoptera). Biological Journal of the Linnean Society 55:
225-237. https://doi.org/10.1111/j.1095-8312.1995.tb01061.x

Guenée MA (1858) Histoire naturelle des insectes; spécies généraldes lépidoptéres.
Vol. 10 Uranides et Phalénites. Librairie Encyclopédique de Roret, Paris, 584 pp.

Hausmann A, Parra LE (2009) An unexpected hotspot of moth biodiversity in Chilean
northern Patagonia (Lepidoptera, Geometridae). Zootaxa 1989: 23-38. https://doi.
org/10.11646/zootaxa.1989.1

Holloway J (1993) The moths of Borneo, part 11: Family Geometridae, subfamily
Ennominae. Malayan Nature Journal 47: 1-309.

Klots AB (1970) Lepidoptera. In: Tuxen SL (Ed.) Taxonomist’s glossary of genitalia in
insects, 115-130. Second Enlarged Edition. Munksgaard, Copenhagen, 359 pp.

Luebert F, Pliscoff P (2018) Sinopsis bioclimatica y vegetacional de Chile. 2" edn.
Editorial Universitaria, Santiago, Chile.

ZooKeys 1216: 173-200 (2024), DOI: 10.3897/zookeys.1216.129923 198

Mario |. Ramos-Gonzédlez et al.: Revision of the genus Digonis (Lepidoptera, Geometridae)

Morrone JJ (2015) Biogeographical regionalisation of the Andean region. Zootaxa
3936(2): 207-236. https://doi.org/10.11646/zootaxa.3936.2.3

Ounap E, Javois J, Viidalepp J, Tammaru T (2011) Phylogenetic relationships of select-
ed European Ennominae (Lepidoptera: Geometridae). European Journal of Entomol-
ogy 108(2): 267-273. https://doi.org/10.14411/eje.2011.036

Parra LE (1991) Revision y filogenia del género Pachrophylla Blanchard, 1852 (sensu
auctorum) (Geometridae: Larentiinae: Trichopterygini). Gayana Zoologia (Chile)
55(2): 145-199.

Parra LE (1999a) Revision of the Neotropical genus Psilaspilates (Lepidoptera: Geo-
metridae). Annals of the Entomological Society of America 92: 460-472. https://doi.
org/10.1093/aesa/92.4.460

Parra LE (1999b) Revision del género Euclidiodes Warren, 1895 (Lepidoptera: Geometri-
dae). Revista Chilena de Historia Natural 72: 643-659.

Parra LE (2018) A new genus of Nacophorini (Geometridae) from Chile. Journal of the
Lepidopterologist Society 72(4): 265-269. https://doi.org/10.18473/lepi.72i4.a3

Parra LE, Henriquez-Rodriguez JL (1993) Aportes al conocimiento de las polillas del
género Mallomus Blanchard, 1852 (Geometridae, Nacophorini). Boletin de la Socie-
dad de Biologia de Concepcion (Chile) 64: 171-187.

Parra LE, Hernandez CE (2010) Estudio filogenético de los géneros de Lithinini de
Sudamérica Austral (Lepidoptera, Geometridae): una nueva clasificacién. Re-
vista Brasileira de Entomologia 54(1): 1-27. https://doi.org/10.1590/S0085-
56262010000100001

Parra LE, Pascual-Toca M (2003) Revision taxonémica de los géneros Oratha Walker,
1863 y Hasodima Butler, 1882 (Lepidoptera: Geometridae). Revista Chilena de Histo-
ria Natural 76: 117-128. https://doi.org/10.4067/S0716-078X2003000100011

Parra LE, Vargas HA (2000) Revision del género Neorumia Bartlett-Calvert, 1893 (Lepi-
doptera: Geometridae). Revista Chilena de Entomologia 27: 91-98.

Parra LE, Villagran-Mella R, Marquet PA (2009) Phylogeny of the genera Euclidiodes
and Hasodima (Lepidoptera: Geometridae) and description of two new species
from the Fray Jorge relict forest in northern Chile. Zootaxa 2273: 59-68. https://doi.
org/10.11646/zootaxa.2273.1.3

Parra LE, Villagran-Mella R, Hernandez EA, Hernandez CE (2010) Filogenia del géne-
ro Psilaspilates (Butler 1893) (Lepidoptera: Geometridae) con la descripcién de
una nueva especie del bosque relicto Fray Jorge, Chile. Gayana 74(2): 94-101.
https://doi.org/10.4067/S071 7-65382010000200003

Pitkin LM (2002) Neotropical ennomine moths: a Review of the genera (Lepidop-
tera: Geometridae). Zoological Journal of the Linnean Society 135(2-3): 121-401.
https://doi.org/10.1046/j.1096-3642.2002.00012.x

Rajaei H, Hausmann A, Scoble MJ, Wanke D, Plotkin D, Brehm G, Murillo-Ramos L, Sih-
vonen P (2022) An online taxonomic facility of Geometridae (Lepidoptera), with an
overview of global species richness and systematics. Integrative Systematics 5(2):
145-192. https://doi.org/10.18476/2022.577933

Rindge FH (1973) Notes on and descriptions of South American Nacophorini (Lepidop-
tera, Geometridae). American Museum Novitates 2531: 1-42.

Rindge FH (1983) A generic revision of the new world Nacophorini (Lepidoptera, Geo-
metridae). Bulletin of the American Museum of Natural History 175(2): 147-262.
Rindge FH (1986) Generic descriptions of new world Lithinini (Lepidoptera, Geometri-

dae). American Museum Novitates 2838: 1-68.

ZooKeys 1216: 173-200 (2024), DOI: 10.3897/zookeys.1216.129923 199

Mario |. Ramos-Gonzélez et al.: Revision of the genus Digonis (Lepidoptera, Geometridae)

Scoble MJ (1995) The Lepidoptera: Form, Function and Diversity. The Natural History
Museum & Oxford University Press, London, 416 pp.

Scoble MJ (1999) Geometrid moths of the world: a catalogue (Lepidoptera, Geomet-
ridae). CSIRO Publishing. Natural History Museum, London, 1016 pp. https://doi.
org/10.1071/9780643101050

Scoble MJ, Hausmann A (2007) Online list of valid and available names of the Geo-
metridae of the World. Lepidoptera Barcode of Life. https://www.lepbarcoding.org/
geometridae/species_checklists.php [accesed on 15 Feb 2022]

Vargas HA (2007) Dos nuevas especies de /ridopsis Warren (Lepidoptera, Geometri-
dae) del norte de Chile. Revista Brasileira de Entomologia 51(2): 138-141. https://doi.
org/10.1590/S0085-56262007000200003

Vargas HA (2021) Iridopsis socoromaensis sp. n., a geometrid moth (Lepidoptera, Geo-
metridae) from the Andes of northern Chile. Biodiversity Data Journal 9: e61592.
https://doi.org/10.3897/BDJ.9.e61592

Vargas HA, Parra LE, Hausmann A (2005) Macaria mirthae: una nueva especie de
Ennominae (Lepidoptera: Geometridae) de Chile. Neotropical Entomology 34(4):
571-576. https://doi.org/10.1590/S1 519-566X2005000400006

Vargas HA, Hausmann A, Parra LE (2020) A new species of Macaria Curtis (Lepidoptera:
Geometridae: Ennominae) from the Andes of northern Chile. Revista Brasileira de En-
tomologia 64(2): e20200016. https://doi.org/10.1590/1806-9665-RBENT-2020-0016

ZooKeys 1216: 173-200 (2024), DOI: 10.3897/zookeys.1216.129923 200