New World geometrid moths (Lepidoptera: Geometridae): Molecular phylogeny, biogeography, taxonomic updates and description of 11 new tribes

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ISSN 1863-7221 (print)
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DOI: 10.26049/ASP77-3-2019-5
© Senckenberg Gesellschaft für Naturforschung, 2019.
77
(3): 457 – 48 6
2019
New World geometrid moths (Lepidoptera: Geometridae):
Molecular phylogeny, biogeography, taxonomic updates
and description of 11 new tribes
G B
*, 1, L M-R 2, 14, P S 3,
A H 4, B. C S 5, E Õ 6, 7, A M 8,
R M 9, D B 10, F B 11, A L 12,
L E. P 13 & N W 14
1
Institut für Zoologie und Evolutionsbiologie mit Phyletischem Museum, Erbertstr. 1, 07743 Jena, Germany; Gunnar Brehm * [gunnar.brehm @
uni-jena.de] —
2
Departamento de Biología, Universidad de Sucre; Leidys Murillo-Ramos [Leidys.murillo@unisucre.edu.co] —
3
Finnish Mu-
seum of Natural History, Pohjoinen Rautatiekatu 13, 00100 Helsinki, Finland; Pasi Sihvonen [pasi.sihvonen@helsinki.fi] —
4
Staatliche Natur-
wissenschaftliche Sammlungen Bayerns – Zoologische Staatssammlung München, Münchhausenstr. 21, 81247 München, Germany; Axel
Hausmann [axel.hausmann@zsm.mwn.de] —
5
Canadian National Collection of Insects, Arachnids & Nematodes, Agriculture and Agri-Food
Canada, 960 Carling Ave., Ottawa, ON, K1A 0C6, Canada; B. Christian Schmidt [Christian.Schmidt@agr.gc.ca] —
6
Institute of Ecology and
Earth Sciences, University of Tartu, Vanemuise 46, 51014 Tartu, Estonia; Erki Õunap [erki.ounap@ut.ee] —
7
Institute of Agricultural and
Environmental Sciences, Estonian University of Life Sciences, Kreutzwaldi 5, 51006 Tartu, Estonia —
8
UFRGS – Universidade Federal do Rio
Grande do Sul, Porto Alegre, Brazil and UFPR – Universidade Federal do Parana, Curitiba, Brazil; Alfred Moser [a.moser@ensinger.com.br] —
9
Dürerstraße 12, 76709 Kronau, Germany; Rolf Mörtter [rolf.moertter@t-online.de] —
10
Via Crusch 8a, 7013 Domat/Ems, Switzerland; Daniel
Bolt [Daniel.Bolt@anu.gr.ch] —
11
Department of Botany and Biodiversity Research, Division of Tropical Ecology and Animal Biodiversity, Uni-
versity of Vienna, Rennweg 14, 1030 Vienna, Austria; Florian Bodner [florian.bodner@univie.ac.at] —
12
Estonian Museum of Natural History,
Lai St, 29A, 00001 Tallinn, Estonia; Aare Lindt [aare.lindt@gmail.com] —
13
Departamento de Zoología, Universidad de Concepción, casilla
160-C, Concepción, Chile: Luis E. Parra [luparra@udec.cl] —
14
Department of Biology, Lund University, Sweden; Niklas Wahlberg [niklas.
wahlberg@biol.lu.se] — * Corresponding author
Accepted on November 17, 2019.
Published online at www.senckenberg.de/arthropod-systematics on December 06, 2019.
Published in print on December 20, 2019.
Editors in charge: Brian Wiegmann & Klaus-Dieter Klass.
Abstract. We analysed a molecular dataset of 1206 Geometroidea terminal taxa. In this paper we focus on New World taxa, with 102
Nearctic terminal taxa (97 of which have not previously been subject to molecular phylogenetic analysis) and 398 Neotropical terminal
taxa (375 not previously analysed). Up to eleven molecular markers per specimen were included: one mitochondrial (COI) and ten protein-
coding nuclear gene regions (Wingless, ArgK, MDH, RpS5, GAPDH, IDH, Ca-ATPase, Nex9, EF-1alpha, CAD). The data were analysed
using maximum likelihood approach as implemented in IQ-TREE and RAxML. Photographs of almost all voucher specimens are provided
together with relevant type material in illustrated electronic catalogues in order to make identities and taxonomic changes transparent. Our
analysis concentrates on the level of tribes and genera, many of which are shown to be para- or polyphyletic. In an effort towards a natural
system of monophyletic taxa, we propose taxonomic changes: We establish 11 new tribe names (Larentiinae, authors Brehm, Murillo-
Ramos & Õunap): Brabirodini new tribe, Chrismopterygini new tribe, Psaliodini new tribe, Pterocyphini new tribe, Rhinurini new
tribe, Ennadini new tribe, Cophocerotini new tribe, Erebochlorini new tribe; (Ennominae, authors Brehm, Murillo-Ramos & Sihvonen):
Euangeronini new tribe, Oenoptilini new tribe, Pyriniini new tribe. We assign 27 genera for the rst time to a tribe, propose 29 new tribe
assignments and 26 new generic combinations, we synonymize one tribe and seven genera, revive one tribe, and propose to exclude 119
species from non-monophyletic genera (incertae sedis). Our study provides the data and foundation for numerous future taxonomic revi-
sions of New World geometrid moths. We also examine broad-scale biogeographic patterns of New World Geometridae: While Nearctic
species are often nested within the predominantly Neotropical clades, the austral South American fauna forms distinct clades, hinting at a
long isolation from the remaining New World fauna.
Key words. Geometridae, new tribes, molecular phylogeny, paraphyly, polyphyly, systematics.

B et al.: Phylogeny of New World Geometridae
458
1. Introduction
In the family Geometridae, approximately 24,000 valid
species are known (NieukerkeN et al. 2011; AH, un-
published data), but many others are still undescribed.
Increasing evidence shows that many genera are much
more diverse than previously thought, particularly tropi-
cal lineages with small and inconspicuous species, e.g.,
Drepanogynis Guenée, [1858] (krüger 2002), Eois Hüb-
ner, 1818 (Brehm et al. 2011), Prasinocyma Warren, 1897
(hausmaNN et al. 2016) and Oospila Warren, 1897 (LiNdt
et al. 2018). Geometridae show a worldwide distribution,
but the Neotropical region is more species-rich than any
other, with the wet tropical Andes being the global diver-
sity hotspot of the family (Brehm et al. 2016). The group
is well dened by apomorphies such as a tympanal organ
(with an “ansa”) situated at the base of the abdomen in
the adult moth, and the reduction of larval prolegs (e.g.
miNet & scoBLe 1999). The monophyly of the family is
well supported in molecular phylogenies (e.g. heikkiLä
et al. 2015; muriLLo-ramos et al. 2019). The relation-
ships between the large subfamilies have become rather
clear based on molecular phylogenetic studies over the
last two decades (aBraham et al. 2001; Yamamoto &
sota 2007; WahLBerg et al. 2010; strutzeNBerger et
al. 2017; sihvoNeN et al. 2011; ÕuNap et al. 2016), but
the position and denition of the enigmatic subfamilies
Oenochrominae and Desmobathrinae have been a puzzle
until very recently (muriLLo-ramos et al. 2019).
This study is part of a series of papers on the phy-
logeny of Geometridae. Our common dataset comprises
1206 terminal taxa of Geometroidea from all biogeo-
graphic regions (except Antarctica), with a focus on the
species-rich Neotropical and the Nearctic fauna. Since
these biogeographically important regions have been ne-
glected in previous studies, we expected them to hold the
greatest potential with regard to remaining knowledge
gaps in phylogeny and systematics. Our paper deals in
principle with all New World taxa of the large dataset,
and indeed by far most taxonomic changes concern New
World taxa, with a focus on Larentiinae and Ennominae.
Other papers deal with the relationships of the major lin-
eages of Geometridae at the global level, including the
Oenochrominae, Desmobathrinae and the description of
the new subfamily Epidesmiinae (muriLLo-ramos et al.
2019); Sterrhinae (sihvoNeN et al. accepted); Larentiinae
(E. Õunap et al. in prep.), Boarmiini (L. Murillo-Ramos
et al. in prep.), and diversication patterns across the
family (H. Ghanavi et al. in prep.).
There has been substantial progress in the system-
atics of Geometridae during recent decades, including
landmark book series such as the “Moths of Borneo”
(hoLLoWaY 1994, 1996, 1997) and “Geometrid moths of
Europe” (hausmaNN 2001, 2004; miroNov 2003; haus-
maNN & viidaLepp 2012; skou & sihvoNeN 2015; müL-
Ler et al. 2019). No comparable works at such a broad
scale have been published for the Neotropical region,
with the notable exception of papers on genera of Neo-
tropical Geometrinae (pitkiN 1996) and Ennominae (pit-
kiN 2002). Further recent systematic works focused on
certain genera or tribes and / or regions (examples, list
not comprehensive): in Sterrhinae on the Cyllopodini
(LeWis & coveLL 2008); in Larentiinae on Chilean Eu-
pithecia Curtis, 1825 (riNdge 1987, 1991), on Chilean
Trichopterygini (parra 1991, 1996; parra & saNtos-
saLas 1992a,b; parra et al. 2009a, 2017), on Hagnagora
Druce, 1885 (Brehm 2015), and Callipia Guenée, [1858]
(Brehm 2018); in Ennominae on Chilean Diptychini (=
Lithinini, see Discussion) (riNdge 1986; parra 1999a,b;
parra & herNaNdez 2010; parra et al. 2009b, 2010),
on Chilean and Argentinian Nacophorini (riNdge 1971,
1973, 1983), on Palyadini (scoBLe 1994), on Pero Her-
rich-Schäffer, [1855] (pooLe 1987), on Syncirsodes But-
ler, 1882 (Bocaz & parra 2005), on Thysanopyga Her-
rich-Schäffer, [1855] and Perissopteryx Warren, 1897
(krüger & scoBLe 1992), and on Ischnopteris Hübner,
[1823], Stegotheca Warren, 1900 and Rucana Rindge,
1983 (pitkiN 2005). In Geometrinae, viidaLepp (2017)
investigated the phylogeny of Nemoriini; further studied
genera include Chavarriella Pitkin, 1993, Dioscore War-
ren, 1907, Lissochlora Warren, 1900 and Nemoria Hüb-
ner, 1818 (LiNdt et al. 2014a; pitkiN 1993), Haruchlora
Viidalepp & Lindt (viidaLepp & LiNdt 2014a), Oospila
Warren, 1897 (viidaLepp 2002; viidaLepp & LiNdt 2016;
LiNdt et al. 2018), Paromphacodes Warren, 1897 (LiNdt
et al. 2017), Pyrochlora Warren, 1895 (viidaLepp 2009),
Tachyphyle Butler, 1881 (viidaLepp & LiNdt 2017) and
Telotheta Warren, 1895 (LiNdt & viidaLepp 2014b).
Broad-scale authoritative works on the Nearctic fauna
are limited to the Geometrinae (FergusoN 1985) and Ma-
cariini (FergusoN 2008). The Nearctic fauna was treated
in part (Canada) by mcguFFiN (1967, 1972, 1977, 1981,
1987, 1988) and BoLte (1990).
Some New World taxa have been included in previ-
ous molecular phylogenetic works, in particular in those
focusing on Eois (strutzeNBerger et al. 2010, 2017) and
the subfamily Larentiinae (ÕuNap et al. 2016). However,
New World taxa were heavily underrepresented in other
phylogenetic works, for example in studies with a focus
on the Asian taxa of Geometrinae (BaN et al. 2018) and
Boarmiini (JiaNg et al. 2017). In a global phylogeny of
Geometridae (sihvoNeN et al. 2011), New World taxa
were represented with rather few specimens (only 36 out
of 149 samples). For our study, we targeted New World
taxa in order to address this deciency: Of a total 1206
terminal taxa, our study comprises 102 Nearctic terminal
taxa (97 taxa not previously analysed) and 398 Neotropi-
cal terminal taxa (375 taxa not previously analysed).
The primary objective of this paper is to uncover the
phylogenetic relationships of a large number of New
World Geometridae genera in a global context, and to
identify paraphyletic genera and tribes. Many New
World geometrid tribes and genera are currently non-
monophyletic, and many genera are not even assigned to
tribes, despite pitkiN’s (1996, 2002) studies. In species-
rich genera, our sampling often includes two or more
species. Material of the type species of genera or closely

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ARTHROPOD SYSTEMATICS & PHYLOGENY — 77
(3) 2019
related species was preferably included. Striking exam-
ples of polyphyly include Larentia Treitschke, 1825 (at
four different positions in the tree, see Results) and Peri-
zoma Hübner, [1825] (at six positions, see also ÕuNap
et al. in press).
Moreover, we also aim to formally establish taxonom-
ic changes that are required for a system of monophyla,
i.e. we attempt to translate as many results as possible
into an updated taxonomy of Geometridae. This appears
to still be the exception rather than the rule in phylogenet-
ic research, but see e.g. zahiri et al. (2011) ÕuNap et al.
(2016), muriLLo-ramos et al. (2019), dhuNgeL & WahL-
Berg (2018) as examples. Such taxonomic changes will
be useful for all biologists working with these organisms
who require phylogenetic information and a correspond-
ing modern taxonomy including named entities of their
study organisms. Taxonomic changes of our paper focus
on the two most species-rich subfamilies Ennominae and
Larentiinae, while more such changes are performed in
the other papers of the series (see above).
Beyond phylogenetic systematics and taxonomic
changes, we also investigate broad-scale biogeographic
patterns of New World Geometridae in the context of
global phylogeny. We aim to draw preliminary conclu-
sions about the biogeographic origin of certain clades,
i.e. whether New World lineages are nested within Old
World taxa and vice versa. We also aim to investigate the
relationship between Nearctic taxa and Neotropical taxa.
It is well known that the austral South American ora and
fauna differs considerably from the central and northern
parts (morroNe 2015). hoLt et al. (2013), analysing dis-
tribution and phylogenetic relationships of vertebrates,
placed the vertebrate fauna of all of South America into
the Neotropical faunal realm. We therefore analyse the
austral South American geometrid fauna separately from
the fauna of the remaining continent.
2. Material and Methods
2.1. Sampling and documentation
A total of 93 tribe-level taxa (worldwide) are included in
this study following current phylogenetic hypotheses and
classications (pitktiN 1996, 2002; sihvoNeN et al. 2011,
2015; WahLBerg et al. 2010; ÕuNap et al. 2016). In addi-
tion, 14 non-Geometridae species belonging to the super-
family Geometroidea (Sematuridae, Epicopeiidae, Pseu-
dobistonidae and Uraniidae) were included as outgroups
based on the hypothesis proposed by regier et al. (2009).
Where possible, two or more samples were included per
tribe and genus, especially for species-rich groups that are
widely distributed and in cases where genera were sus-
pected to be poly- or paraphyletic (see muriLLo-ramos
et al. 2019 for further details). Identities of species were
investigated by the authors and consulted experts and
in most cases conrmed by comparing the COI region
(‘DNA barcode’) with sequence data available on BOLD
systems (ratNasiNgham & heBert 2007).
Photographs of adult moths were taken of the ex-
amined material as well as of relevant type material and
compiled in three large illustrated pdf catalogues (sup-
plementary material, see Methods). These catalogues
provide an excellent overview of taxa (e.g. kaWahara
et al. 2018), and they allow readers to easily check the
validity of results and critically assess our taxonomic
changes.
2.2. Molecular techniques
DNA was extracted from 1 – 3 legs preserved either in
ethanol or dry. In a few cases, other sources of tissue,
such as parts of larvae, or full abdomens of adults, were
used. The remaining parts of specimens were preserved
as vouchers and deposited in public museum collections.
Genomic DNA was extracted and puried using Nucleo
Spin® Tissue Kit (MACHERY-NAGEL), according to
the manufacturer’s protocol. DNA amplication and se-
quencing were carried out following protocols proposed
by WahLBerg & Wheat (2008) and WahLBerg et al.
(2016). PCR products were visualized on agarose gels.
PCR products were cleaned enzymatically and sent to
Macrogen Europe (Amsterdam) for Sanger sequencing.
One mitochondrial (COI) and 10 protein-coding nuclear
gene regions (Wingless, ArgK, MDH, RpS5, GAPDH,
IDH, Ca-ATPase, Nex9, EF-1alpha, CAD) were se-
quenced.
2.3. Alignment and cleaning sequences
Multiple sequence alignments were done for each gene
based on a reference sequence of Geometridae down-
loaded from the database VoSeq (peña & maLm 2012).
We used MAFFT algorithm as implemented in Geneious
v.11.0.2 (Biomatters, www.geneious.com). Sequences
with bad quality and ambiguities were removed from the
alignments. Finally, aligned sequences were uploaded to
VoSeq (peña & maLm 2012) and then assembled in a
dataset comprising 1206 taxa. The nal dataset had a
concatenated length of 7665 bp with gaps. To check for
potential misidentications, DNA barcode sequences
were compared to those in BOLD (Barcode of Life Data
Systems) (ratNasiNgham & heBert 2007). After clean-
ing, the nal dataset included at least three genes per
taxon except nine samples (see muriLLo-ramos et al.
2019).
2.4. Tree search strategies and model
selection
We ran maximum likelihood analyses with a dataset par-
titioned by codon position. Different substitution models
were determined implementing ModelFinder (kaLYaaN-

B et al.: Phylogeny of New World Geometridae
460
amoorthY et al. 2017). Dataset with different partitions
and models were analysed using IQ-TREE (NguYeN et
al. 2015) with the MFP+MERGE option (see muriLLo-
ramos et al. 2019). Support for nodes were evaluated
with 1000 ultrafast bootstrap (UFBoot2) approximations
(hoaNg et al. 2017) and SH-like approximate likeli-
hood ratio test (guiNdoN et al. 2010) as implemented in
IQ-TREE. Trees were visualized and edited in FigTree
v1.4.3 software (ramBaut 2012). The nal tree was root-
ed with non-Geometridae species (see muriLLo-ramos et
al. 2019 for further details).
2.5. Taxonomic changes
We propose taxonomic changes if our taxon sampling ap-
pears sufcient (including species-richness, and/or mor-
phological diversity and/or geographical extend of the
lineage) and we are condent with the results, i.e. our
conclusions are supported by high branch-support values
(SH > 80 or UFBoot2 > 95) in the molecular phylogeny.
Further requirements are that our classication identies
monophyletic lineages, we have adequate morphological
material available to us, and identity of examined mate-
rial is conrmed and can be tracked. It is thus of cru-
cial importance that material of type species of genera
or morphologically very similar material was used. Our
conclusions are primarily based on molecular results.
It is beyond the scope of our paper to perform supple-
mentary morphological analyses, but where available,
we have used published information on the morphology,
in particular in Ennominae (pitkiN 2002). We neverthe-
less take obvious morphological features into account for
taxonomic decisions (i.e. wing pattern). We explicitly ac-
knowledge the need of an integrative approach combin-
ing morphological and molecular data in the future (padi-
aL et al. 2010). It is obvious that many of our taxonomic
decisions need to be corroborated by (more) morphologi-
cal data. However, we believe that providing an updated
taxonomy (with the possibility that some conclusions
will later be rejected) has by far more advantages than
drawbacks, and hopefully will stimulate more research
on poorly studied taxa. For example, it is more denite
and concise to refer in future works to “Erebochlorini”
than to an “unnamed Larentiinae clade comprising the
genera Erebochlora Warren, 1895, Cirrolygris Warren,
1895, and Deinoptila Warren, 1900”.
Proposed changes to the current classication are ex-
plicitly stated and summarized in a table for three affect-
ed subfamilies. In this paper, we propose: 1) new tribes,
2) new tribe synonymies, 3) new tribe assignments, 4)
new genus-level synonymies, 5) new combinations, and
6) genera listed – ad interim – in quotation marks. The
latter includes the exclusion of a species from its current
combination. For example, many Chilean species were
originally (or later) assigned to Palaearctic or Holarc-
tic genera. We follow the practice used e.g. by scoBLe
(1999) and pitkiN (2002) and put doubtful genus combi-
nations into quotation marks.
Results and Discussion are given at the subfamily
level in the following order: Sterrhinae, Larentiinae, Ar-
chiearinae, Desmobathrinae, Oenochrominae, Geometri-
nae, Ennominae, and within subfamilies, taxa are treated
in the order of the tree derived from IQ-TREE analysis
(Electronic Supplement Files 1 and 2).
3. Results
In this section, we present a short overview of the results,
including all tables and gures. See muriLLo-ramos et
al. (2019) for a more detailed overview. In order to avoid
redundancy, detailed results are presented and discussed
together in the next section. Results of both the IQ-TREE
analyses (Electronic Supplement Files 1 and 2) and the
RAxML analyses (muriLLo-ramos et al. 2019) are very
similar with only a few exceptions. Neotropical taxa are
found throughout the topology, with several larger radia-
tions in South America. Sterrhinae: See Fig. 1 for an
overview at the tribe level and Electronic Supplement
Files 1 and 2. Specimens are illustrated in Electronic
Supplement File 3. Larentiinae: Our analyses show a
large number of new, well-supported, tribe level clades
which are discussed in detail in the Discussion section.
See Fig. 2 for an overview at the tribe level, Fig. 3 for
images of adult moths and Electronic Supplement Files
1 and 2. Specimens are illustrated in Electronic Supple-
ment File 4. Taxonomic changes in Larentiinae are sum-
marized in Table 1. Geometrinae: See Fig. 4 for an over-
view at the tribe level and Electronic Supplement Files 1
and 2. Specimens are illustrated in Fig. 5 and Electronic
Supplement File 3. Taxonomic changes in Geometrinae
are summarized in Table 2. Ennominae: See Fig. 6 for
an overview at the tribe level, Fig. 7 for images of adult
moths and Electronic Supplement Files 1 and 2. All ana-
lysed specimens are illustrated in Electronic Supplement
File 5. Taxonomic changes in Ennominae are summarized
in Table 3. Small subfamilies Archiearinae, Oenochro-
minae, Desmobathrinae, Epidesmiinae: See Elec tronic
Supplement Files 1 and 2. Specimens are illus trated in
Electronic Supplement File 3.
4. Discussion
4.1. Sterrhinae Meyrick, 1892
See Fig. 1 for phylogenetic relationships.
Sterrhinae will be dealt with in detail by sihvoNeN et al.
(accepted), they are not illustated in the text and the dis-
cussion of this subfamily is therefore kept to a minimum.
The genera Almodes Guenée, [1858], Ametris Hübner,

461
ARTHROPOD SYSTEMATICS & PHYLOGENY — 77
(3) 2019
[1822], Ergavia Walker, 1866, and Macrotes Westwood,
1841 will be transferred from Oenochrominae to Ster-
rhinae in the revived tribe Mecoceratini (sihvoNeN et al.
accepted). Existing tribe assignments of New World taxa
are supported in most cases in Cosymbiini, Sterrhini and
Scopulini. Pseudasellodes Warren, 1904 (not assigned
to tribe so far) is sister to Proutoscia Schaus, 1912. Re-
markably, “Lophochorista” porioni Herbulot, 1988 is not
a Geometrinae but belongs to a larger Neotropical clade
within the tribe Sterrhini. The association of “Lopho-
chorista” porioni with Sterrhinae was already suggested
by scoBLe (1999), questioning Herbulot’s generic com-
bination. Our data suggest that two genera currently as-
signed to Cosymbiini (Hemipterodes Warren, 1906 and
Lipotaxia Prout, 1918) also belong to Sterrhini clade.
Furthermore, our data indicate that yellow-black colora-
tion has evolved potentially three times independently
in diurnal Neotropical Sterrhinae. Xanthyris Felder &
Felder, 1862 is not closely related to the other two in-
vestigated Cyllopodini genera Atyria Hübner, 1823 and
Smicropus Warren, 1895. Support for a sister group re-
lationship between Atyria and Smicropus is poor in the
RAxML analysis (muriLLo-ramos et al. 2019), and the
two genera even fall into different parts of the IQ-TREE
analysis.
4.2. Larentiinae Duponchel, 1845
See Fig. 2 for phylogenetic relationships, Fig. 3 for habitus pictures
of exemplary species, and Table 1 for proposed taxonomic changes.
Our study focuses on the tribe composition and formal
taxonomic changes required for a natural system of the
subfamily. Such changes include both New and Old
World taxa. The relationships within the subfamily will
be discussed in more detail by E. Õunap et al. (in prep.).
4.2.1. Dyspteridini. The tribe was revived by viidaLepp
(2011) from synonymy with Trichopterygini. The posi-
tion of Dyspteridini as sister to all other studied Larentii-
nae is conrmed (strutzeNBerger et al. 2010; sihvoNeN
et al. 2011; ÕuNap et al. 2016). sihvoNeN et al. (2011)
found a close relationship of Neotropical Dyspteris Hüb-
ner, 1818 (Fig. 3A), and the New Zealand genus Para-
detis Meyrick, 1885, conrmed by ÕuNap et al. (2016)
and our study. Our analysis conrms a close relationship
of European Celonoptera Lederer, 1862 with Dyspteris
which was already suspected by early authors (cited in
ÕuNap et al. 2016). Celonoptera, Heterophleps Herrich-
Schäffer, [1854] and Chlorotimandra Butler, 1882, are
formally transferred to Dyspteridini (Table 1). Since
apart from the type species, almost all other members of
Heterophleps occur in Asia, the monophyly and tribe as-
signment of Asian species combined to this genus need to
be investigated in future studies.
4.2.2. Brabirodini Brehm, Murillo-Ramos & Õunap,
new tribe. — Type genus. Brabirodes Warren, 1904
(Fig. 3B). — Material examined and phylogeny. Brabi-
rodes cerevia peruviana Warren, 1904. Brabirodes forms
a distinct lineage of Larentiinae which is sister to the rest
of the subfamily, except Dyspteridini. Branch support
values from the IQ-TREE analyses strongly conrm the
sister-relationship to all other Larentiinae with the ex-
ception of Dyspteridini (SH-like = 83.5, UFBoot2 = 83).
— Molecular evidence. The tribe is characterized by
DNA sequence data from the following ve gene regions
(voucher gb-ID-19269, Brabirodes cerevia, from Ecua-
dor, illustrated in Electronic Supplement File 5): ArgK
Scopulini
unnamed clade
Cosymbiini
Timandrini
Rhodometrini / Lythriini
Cyllopodini / Rhodostrophiini
unnamed clade
Sterrhini
unnamed clade
3 1 2
5 1 2
3 1 1
9 5
15 1 12 1
1 1
13 1 9
3 3
25 3
Tribename
1 1 1 1
other region
Nearctic
Neotropical
austral region
Genera
in dataset
Tribe
5
Analysed
specimens
* * *asterisk: known to be
present but not in dataset
0.2
Legend
* *
99.9/100
99.4/99
100/100
100/100
88.7/97
57.9/43
100/98
98.5/98
100/100
99.9/76
100/90
95.8/83
99.2/98
33.9/55
100/100
94.1/86
Fig. 1. Sterrhinae tribe composition. Tribes with New World representatives marked in colour: light green Nearctic, dark green Neotropical,
yellowish green austral region. Support values in blue colour (SH-like and UFBoot values).

B et al.: Phylogeny of New World Geometridae
462
(MK738419), COI (MK739303), EF1a (MK739881),
MDH (MK741089), Nex9 (MK741532). — Morpho-
logy. Brabirodes supercially resembles Eupithecia
species (and is sometimes found in collections among
Eupithecia). Brabirodes can be diagnosed by large and
extremely elongated forewings, hindwings are small with
undulating margin, anal area of male hindwing is with-
out fold, and male antennae are bipectinate. Male geni-
talia are illustrated in viidaLepp (2011). Combination of
these characters differentiates it from Dyspteridini and
Trichopterygini (see viidaLepp 2011 for details).
4.2.3. Trichopterygini. Our results conrm the nd-
ing that Trichopterygini and Chesiadini are not sister
taxa (ÕuNap et al. 2016). However, the position of these
tribes has switched in our analysis compared to previous
studies, meaning that Trichopterygini have branched off
from the main lineage Larentiinae earlier than Chesia-
dini. While tribe assignment is conrmed in most cas-
es, there are four formal new tribe assignments of New
World genera: Aloba Warren, 1895, Anomozela Fletcher,
1979 (Fig. 3C), Isosauris Warren, 1894, and Synpelurga
Butler, 1882 are transferred to Trichopterygini (Table
1). Lobidiopteryx Warren, 1902 was treated by prout
(1929 – 1935) as “one of the few African representatives
of the Lobophora group of genera”, with the Old World
genus Episteira Warren, 1899 being listed almost imme-
diately after it. The former “Lobophora group” has sub-
sequently been changed to Lobophorini, and then syn-
onymized with Trichopterygini (ÕuNap et al. 2016). We
formally transfer the two genera as well as New Zealand-
ian Tatosoma Butler, 1874 to Trichopterygini (Table 1),
as earlier suggested by dugdaLe (1980) and ÕuNap et al.
(2016). Moreover, an undescribed genus (voucher PS225
from South Africa) also belongs to this tribe.
4.2.4. Chesiadini. Analysed specimens currently com-
prise three Palaearctic samples (genera Aplocera Ste-
phens, 1827, and Chesias Treitschke, 1825), but none of
our New World samples falls into this tribe. Currently,
several New World species are assigned to Lithostege
Hübner, [1825]. None of the Nearctic species belongs
to Lithostege (B.C.S., unpubl. data), and it needs to be
established whether any of the Neotropical species are
actually congeneric with the Palaearctic type species of
the genus.
4.2.5. Chrismopterygini Brehm, Murillo-Ramos &
Õunap, new tribe. — Type genus. Chrismopteryx Prout,
1910 (Fig. 3D). — Material examined and phylogeny.
The clade comprises Chrismopteryx politata Fletcher,
1953, an unidentied Chrismopteryx species, “Nebula”
pseudohalia (Butler, 1882), and “Anticlea” oculisigna
Prout, 1923. Branch support values from the IQ-TREE
analyses clearly conrm the monophyly of this clade
(SH-like = 99.9, UFBoot2 = 100). — Molecular evi-
dence. The tribe is characterized by DNA sequence data
from the following seven gene regions (exemplar C.
politata, voucher bo_chi_120 from Chile, illustrated in
Fig. 3D): ArgK (MK738169), CAD (MK738909), COI
(MK739064), EF1a (MK739699), GADPH (MK740314),
Nex9 (MK741346), Wingless (MK742140). — Mor-
phology. Delicately built species. Forewings wide, post-
medial line often undulating, medial area often weakly
darkened. Hindwings with weak markings or markings
absent. External features of analysed species are illustrat-
ed in Fig. 3D and Electronic Supplement File 4. — Re-
marks and taxonomic changes. Psaliodes pseudohalia
Butler, 1882 is transferred from Nebula to Chrismopteryx
(comb.n.) (Table 1). All other Chilean “Nebula” species
are excluded from the genus (Table 1). Since “Anticlea”
oculisigna Prout, 1923 [1855] is misplaced, the genus
should be listed – ad interim – with quotation marks (Ta-
ble 1); the type species of Anticlea Stephens, 1831 (Lar-
entiini) is Palaearctic. Immature stages of Chrismopteryx
undularia (Blanchard, 1852) are described in vargas et
al. (2010).
4.2.6. Eudulini. Our results confirm the phylogenetic
position of the tribe as presented by ÕuNap et al. (2016).
They showed that the New World genera Eubaphe Hüb-
ner, 1823, and Eudulophasia Warren, 1897, form a well
supported clade. Our analysis now also includes Eud-
ule Hübner, 1823, and it shows that the three genera are
closely related. Our results also suggest that the Neo-
tropical genera Graphidipus Herrich-Schäffer, [1855]
(Fig. 3E) and Crocypus Herrich-Schäffer, [1855] form a
lineage sister to this clade, and can thus be formally inte-
grated into the Eudulini (Table 1, Electronic Supplement
File 4).
4.2.7. Asthenini. This tribe is represented only by rela-
tively few taxa in the New World, namely the Holarctic
Hydrelia Hübner, [1825] and Venusia Curtis, 1839 (with
Palaearctic species in the analysis). The only known ge-
nus of this tribe occurring in the Neotropical region is
Eois (Fig. 3F) – but with more than 200 described and
many more undescribed species (Brehm et al. 2011)
probably outnumbering all other taxa of this tribe in
terms of species richness. Phylogenetic relationships
within the Asthenini were already reported by sihvoNeN
et al. (2011) and are supported by further analyses (e.g.
ÕuNap et al. 2016).
4.2.8. Perizomini. Our analysis only comprises mate-
rial sampled in Europe: the type species of Perizoma
Hübner, [1825], P. albulata ([Denis & Schiffermüller],
1775); three more species of Perizoma, and one species
of Mesotype Hübner, [1825]. Probably none of the na-
tive North American “Perizoma” species is congeneric
with true Perizoma – P. alchemillata (Linnaeus, 1758)
has been introduced to North America from Europe – and
it is possible that the tribe is naturally not present at all in
the New World (B.C.S., unpublished). All other sampled
“Perizoma” species belong to other tribes (for details
see 4.2.14. Psaliodini, 4.2.18. Scotopterygini, 4.2.20.2.
Larentiini, 4.2.20.4 Ennadini). Perizoma has thus been
a Larentiinae “trash bin”, and it seems likely that even

463
ARTHROPOD SYSTEMATICS & PHYLOGENY — 77
(3) 2019
more lineages were assigned to the genus, e.g. species
around P. fallax Warren, 1905. Similar results have also
been shown by ÕuNap et al. (in press). We conclude that
it is very likely that all New World species assigned to
Perizoma are probably misplaced, and we therefore sug-
gest to list the genus – ad interim – with quotation marks
for all its Neotropical species (Table 1). An integrative
revision of the genus is required to provide new nomen-
clatorial combinations – and to clean this “Larentiinae
trash bin”.
7 1 6
Dyspteridini
Brabirodini, new tribe
Trichopterygini
Chesiadini
Chrismopterygini, new tribe
Eudulini
Asthenini
Perizomini
Melanthiini
Eupitheciini
Unnamed clade
Trip+Phil+Rheu
Psaliodini, new tribe
Unnamed clade
Unnamed clade
Cidariini
Scotopterygini
Unnamed clade
Euphyiini
Pterocyphini, new tribe
Rhinurini, new tribe
Larentiini
Unnamed clade
Ennadini, new tribe
Hydriomenini
Heterusiini
Cophoceratini, new tribe
Erateinini
Erebochlorini, new tribe
Stamnodini
2 1 1 1
6
11
28 7 3 14
3 2
10 7 1
5 2
6 2
33 5
1 1
10 3 1 1
14 8 1
5 2
3 2
14 7 2
5 1
2 1 1
11 1 3
4 2 2
2 1 1
4 4
4 2 1
10 3 4
5 1 1 1
3 2
3 2
5 1
5 3
11 1 1 5 1
*
*
* *
7 2 2
7 3 1 1
4 2 1
6 3 1
Xanthorhoini
Cataclysmini
Epirhoini, stat rev.
Operophterini
*
0.2
4 3
99.9/100
100/100
100/100
97.8/95
100/100
83.5/83
95.8/77
99.9/100
100/100
100/100
100/99
52.8/37
5.6/45
97.8/89
98.8/70
100/100
96/93 100/100
100/100
9.6/44
100/100
99.9/96
86.3/68
87.4/47
100/100
99.1/84
98.7/80
93.3/97
87/79
96.7/76
99.9/100
100/99
89.3/55
94/76
99.9/100
91.9/66
100/99
88.4/72
96.9/97
100/100
86.1/62
88.9/74
82.6/65
100/100
99.9/100
99.7/99
78.4/54
100/100
100/100
100/100
92.1/82
28.1/71
99.8/100
100/100
97.5/71
18.2/24
100/100
99.9/99
95.9/91
98.2/98
95.5/84
99.5/98
100/100
51.1/68
100/98
Larentiini complex Euphyiini-Xanthorhoini complex
Fig. 2. Larentiinae tribe composition. Tribes with New World representatives marked in colour: light green Nearctic, dark green Neotropi-
cal, yellowish green austral region; see also explanatory box in Fig. 1. Trip+Phil+Rheu: Triphosini + Phileremini + Rheumapterini clade.
Support values in blue colour (SH-like and UFBoot values).

B et al.: Phylogeny of New World Geometridae
464
4.2.9. Melanthiini. Our data only comprises material
sampled in Europe and Africa (several species of Horisme
Hübner, [1825], and the type species of Melanthia Du-
ponchel, 1829). Only a few Horisme species occur in the
New World, and their assignment needs to be checked in
future studies.
4.2.10. Eupitheciini. Our phylogeny currently comprises
no New World samples. However, the globally most spe-
cies-rich genus Eupithecia Curtis, 1825, is also one of the
most species-rich genera in North America (BoLte 1990),
and the Andes are possibly even the most species-rich
region in the world for Eupithecia (Brehm et al. 2016)
where it shows remarkable morphological diversication
(herBuLot 2001).
4.2.11. Operophterini. Our analysis comprises the Hol-
arctic genera Operophtera Hübner, [1825], Malacodea
Tengström, 1869, and Epirrita Hübner, 1822. We are not
aware of the presence of Operophterini in the Neotropi-
cal region.
4.2.12. Solitaneini. Baptria Hübner, [1825] is an enig-
matic genus currently assigned to the tribe Solitaneini
based on morphology (hausmaNN & viidaLepp 2012).
The phylogenetic position of Solitanea Djakonov, 1924
and Solitaneini needs to be tested in future studies (Õu-
Nap et al. 2016).
4.2.13. Clade Triphosini + Phileremini + Rheumapte-
rini. A close relationship of these tribes has been recog-
nized before (e.g., ÕuNap et al. 2016; schmidt 2017) –
and a synonymization could be considered in future
works. Our representatives of New World “Triphosa”,
and the type species of Coryphista Hulst, 1896 fall into
the Rheumapterini, very close to Rheumaptera Hübner,
1822. We regard monotypic Coryphista as a junior syno-
nym of Rheumaptera (Table 1). Scotosia pallidividata
Snellen, 1874 is transferred from Triphosa Stephens,
1829 to Rheumaptera (comb.n.) (Fig. 3G), and Scotosia
afrmata Guenée, [1858] is transferred from Triphosa to
Rheumaptera (comb.n.) (Table 1, illustrated in Electron-
ic Supplement File 4). The generic placement of further
New World species currently assigned to Triphosa need
to be investigated in future studies.
4.2.14. Psaliodini Brehm, Murillo-Ramos & Õunap,
new tribe. — Type genus. Psaliodes Guenée, [1858]
(Fig. 3H). — Material examined and phylogeny. Psali-
odes near planiplaga Warren, 1904 clusters with P. pru-
nicolor (Warren, 1904), and the two taxa are sister to Dis-
toneura pastaza (Prout, 1934). Only Psaliodini s.str. has
high support values from the IQ-TREE analyses (SH-like
= 99.9, UFBoot2 = 100) whereas branch support values of
Psaliodini s.l. are weaker (SH-like = 78.4, UFBoot2 = 54),
The assignment of the genera Anthalma Warren, 1901,
Plemyriopsis Warren, 1895, and Smileuma Prout, 1910 to
Psaliodini therefore requires further scrutiny. — Molecu-
lar evidence. The tribe is characterized by DNA sequence
data from the following six gene regions (exemplar Psali-
odes near planiplaga, voucher gb-CR-S-1708 from Costa
Rica, illustrated in Electronic Supplement File 5) CAD
(JF785161), COI (JF784674), EF1a (JF785299), IDH
(JF785474), MDH (JF784818), Wingless (JF785049).
Our analysis includes Psaliodes near planiplaga which
is – judged from wing morphology – closely related with
P. avagata Guenée [1858], the type species of Psali-
odes (Electronic Supplement File 4). Distoneura Fletcher,
1979, is the second genus that can safely be assigned to
Psaliodini. Further phylogenetic studies should estab-
lish whether this genus is sister to or nested within the
species-rich genus Psaliodes. — Morphology. External
features of analysed species are illustrated in Fig. 3H and
Electronic Supplement File 4. Further detailed morpho-
logical analysis is required to identify potential diagnos-
tic features. — Remarks and taxonomic changes. One
clade comprises “Monarcha” (scoBLe 1999: no published
reference found; apparently preoccupied in Aves: Mo-
narcha Vigors & Horseld, 1827) and “Psaliodes” picta
Warren, 1904. Another clade comprises three unidenti-
ed species of Anthalma Warren, 1901 and “Euphyia”
balteata (Warren, 1905) (wrong generic placement: see
below). Another clade includes Plemyriopsis and Smi-
leuma Prout, 1910, another includes “Nebula” cynthia
(Butler, 1882), “Nebula” near emilia (Butler, 1882) and
“Euphyia” psyroides Warren, 1897 stat.rev. (from Peru)
which we revive from synonymy with “Euphyia” psyra
Druce, 1883 (from Guatemala) (Table 1). A strange co-
incidence is that Herbulot described Epirrhoe psyroides
Herbulot, 1988 from Bolivia which appears to be the
same species as psyroides Warren (all taxa illustrated in
Electronic Supplement File 4). Herbulot’s taxon would be
a junior homonym of Warren’s taxon only once the two
taxa are combined with a nomenclaturally available (new)
genus name in future studies. For erroneous placement
of Chilean “Nebula”, see also Chrismopterygini and En-
nadini. Since “Epirrhoe” psyroides Herbulot, 1988 is not
related to true Epirrhoe (see Epirrhoini), the genus should
be listed – ad interim – with quotation marks (Table 1).
All species are illustrated in Electronic Supplement File
5. Because of their wing morphology, the species Cidar-
ia bogotata Walker, 1862 and Plerocymia rhombifascia
Warren, 1905 are transferred from Perizoma to Smileuma
(comb.n.) (Table 1, illustrated in Electronic Supplement
File 4). “Psaliodes” picta should be listed – ad interim –
with quotation marks (see true Psaliodes in Psaliodini)
because it is not in the same subclade as Psaliodes near
planiplaga. P. Strutzenberger et al. will revise the Psali-
odes group (including “Monarcha” magicaria Felder &
Rogenhofer, 1875) and will revive Alydda Walker, 1861
with subsequent new nomenclatorial combinations. Orth-
oprora balteata Warren, 1905 is transferred from Euphyia
Hübner, [1825] to Anthalma (comb.n.) and Rhopalodes
parecida Dognin, 1892 is transferred from Rhopalodes
to Anthalma (comb.n.) (Table 1). Because of their wing
morphology, twelve further species are transferred to An-
thalma, either from Euphyia or Perizoma (Table 1, illus-
trated in Electronic Supplement File 4).

465
ARTHROPOD SYSTEMATICS & PHYLOGENY — 77
(3) 2019
4.2.15. Unnamed clade. This entirely Neotropical clade
comprises a species that is apparently undescribed and
that cannot be assigned to a genus, and four species of
Perizoma that are hereby excluded from Perizoma, i.e.
the taxa vireonaria Maassen, 1890, cretinotata Bastel-
berger, 1907, versatilis Dognin, 1911 and cyrtozona
Prout, 1922 (Table 1). As judged by wing morphology,
“Perizoma” amplata Warren, 1904 and “Perizoma” miri-
ca Warren, 1904 also belong to this clade (illustrated in
Electronic Supplement File 4). This amplata-group prob-
ably requires the establishment of a new genus which is
beyond the scope of this study.
4.2.16. Unnamed clade. This well supported clade (SH-
like = 100, UFBoot2 = 100) comprises Lampropteryx Ste-
phens, 1831 and Coenotephria Prout, 1914. Both gen era
are mainly distributed in the Old World, although Lam-
pro pteryx suffumata ([Denis & Schiffermüller], 1775) is
also naturally Holarctic (deWaard et al. 2008). In the
analysis by ÕuNap et al. (2016), this clade was the basal-
A C DB
E G HF
I J K L
M N O P
Q R S T
Fig. 3. Illustrations of selected Neotropical Larentiinae taxa. A: Dyspteridini, Dyspteris sp. (Pe-Geo-0205). B: Brabirodini, new tribe,
Brabirodes cerevia peruviana Warren, 1904 (Pe-Geo-0495). C: Trichopterygini, Anomozela cirrhiata (Felder & Rogenhofer, 1875) (Ch-
Geo-0009). D: Chrismopterygini, new tribe, Chrismopteryx politata Fletcher, 1953 (bo_chi_120). E: Eudulini, Graphidipus pilifera (Do-
gnin, 1912) (Pe-Geo-0621). F: Asthenini, Eois near golosata (Dognin, 1893) (Pe-Geo-0119). G: Rheumapterini, Rheumaptera pallidivittata
(Snellen, 1874) comb.n. (Pe-Geo-0039). H: Psaliodini, new tribe, Psaliodes near tripartita (Warren, 1904) (Pe-Geo-0199). I: Euphyiini,
Oligopleura malachitaria (Herrich-Schäffer, [1855]) (Pe-Geo-0012). J: Pterocyphini, new tribe, Pterocypha gibbosaria Herrich-Schäffer,
[1855] (Br-Geo-059). K: Xanthorhoini, Orthonama near plemyrata (Felder & Rogenhofer, 1875) (GB_Geo_068). L: Epirrhoini, revived
tribe, “Euphyia” sturnularia Herrich-Schäffer, [1855] (EO1180). M: Rhinurini, new tribe, Rhinura variegata (Warren, 1901), synonym
of R. populonia (Druce, 1893) (type specimen in NHM, London). N: Ennadini, new tribe, Ennada pellicata (Felder & Rogenhofer, 1875)
(Ch-Geo-0010). O: Hydriomenini, Ersephila prema Druce, 1893 (Gu_Geo_006). P: Heterusiini, Heterusia quadruplicaria (Geyer, 1832)
(AH7173). Q: Cophoceratini, new tribe, Cophocerotis costinotata (Warren, 1908) (gb-ID-19302). R: Erateinini, Erateina drucei (Thierry-
Mieg, 1893) (Pe-Geo-0534). S: Erebochlorini, new tribe, Erebochlora near tesserulata Felder & Rogenhofer, 1875 (gb-CR-S-1218). T:
Stamnodini: Callipia anthocharidaria (Oberthür, 1881) comb.n. (Pe-Geo-0804).

B et al.: Phylogeny of New World Geometridae
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Table 1. Taxonomic changes in Larentiinae at the level of tribes, genera and species, in alphabetical order. *Old World taxa.
Revived tribes Included taxa
Epirrhoini stat. rev. Epirrhoe Hübner, [1825], Catarhoe Herbulot, 1951, Mimoclystia Warren,
1901, Euphyia sturnularia Herrich-Schäer, [1855]
New tribes Included taxa
Brabirodini new tribe Brabirodes Warren, 1904
Chrismopterygini new tribe Chrismopteryx Prout, 1910
Cophoceratini new tribe Cophocerotis Warren, 1895, Hagnagora Druce, 1885
Ennadini new tribe Ennada Blanchard, 1852, Spargania Guenée, [1858], several unnamed
genera
Erebochlorini new tribe Erebochlora Warren, Cirrolygris Warren, 1895, Deinoptila Warren, 1900
Pterocyphini new tribe Pterocypha Herrich-Schäer, [1855], Obila Walker, 1869, Archirhoe
Herbulot, 1951
Psaliodini new tribe Psaliodes Guenée, [1858], Distoneura Fletcher, 1979
Rhinurini new tribe Rhinura Warren, 1904, Haplolabida Fletcher, 1958 *, Urocalpe Warren,
1904
Revived genera Was in synonymy with; included species
Synneuria Mabille, 1885 stat. rev. Stamnodes Guenée, [1858]; Synneuria camposi Orfila & Schajovski,
1964, Synneuria carcavalloi Orfila & Schajovski, 1962, Synneuria ditis-
sima Thierry-Mieg, 1904
Revived species Was in synonymy with
Euphyia psyroides Warren, 1897 stat. rev. Euphyia psyra Druce, 1883
Synonymized genera Valid genus
Anemplocia Warren, 1905 syn.n. Erateina Doubleday, 1848
Coryphista Hulst, 1896 syn.n. Rheumaptera Hübner, 1822
Cyclica Grote, 1882 syn.n. Hydriomena Hübner, [1825]
Priapodes Warren, 1895 syn.n. Erebochlora Warren, 1895
„Trocherateina“ Prout, ‘no published reference’ (F 1979) Erateina Doubleday, 1848
New generic combinations Originally described in genus, transferred from genus, decision
based on
Anthalma alboscripta (Dognin, 1892) comb.n. Cidaria, Euphyia, external morphology
Anthalma apicesignata (Dognin, 1913) comb.n. Perizoma, external morphology
Anthalma arcillata (Dognin, 1893) comb.n. Cidaria, Perizoma, external morphology
Anthalma artemas (Schaus, 1912) comb.n. Anapalta, Euphyia, external morphology
Anthalma balteata (Warren, 1905) comb.n. Orthoprora, Euphyia, molecular data and external morphology
Anthalma cortada (Dognin, 1893) comb.n. Cidaria, Euphyia, external morphology
Anthalma cortatoides (Dognin, 1893) comb.n. Cidaria, Euphyia, external morphology
Anthalma parecida (Dognin, 1892) comb.n. Lobophora?, Rhopalodes, external morphology
Anthalma plumbeipennis (Dognin, 1914) comb.n. Orthoprora, Euphyia, external morphology
Anthalma curviviata (Dognin, 1914) Euphyia, external morphology
Anthalma rojiza (Dognin, 1893) comb.n. Cidaria, Euphyia, external morphology
Anthalma terminisecta (Dognin, 1914) comb.n. Anapalta, Euphyia, external morphology
Anthalma zara (Thierry-Mieg, 1893) comb.n. Cidaria, Euphyia, external morphology
Callipia anthocharidaria (Oberthür, 1881) comb.n. Larentia, Stamnodes, molecular data and external morphology
Chrismopteryx pseudohalia (Butler, 1882) comb.n. Psaliodes, Nebula, molecular data and external morphology
Euphyia tricolorata (Dognin, 1902) comb.n. Ochyria, Xanthorhoe, molecular data and external morphology
Smileuma bogotata (Walker, 1862) comb.n. Cidaria, Perizoma, external morphology
Smileuma rhombifascia (Warren, 1905) comb.n. Plerocymia?, Perizoma, external morphology
Rheumaptera pallidivittata (Snellen, 1874) comb.n. Scotosia, Triphosa, molecular data and external morphology
Rheumaptera armata (Guenée, [1858]) comb.n. Scotosia, Triphosa, molecular data and external morphology
Scotopteryx bitrita (Felder & Rogenhofer, 1875)* comb.n. Ortholitha, Larentia, molecular data
Scotopteryx epipercna (Wehrli, 1931)* comb.n. Onychia, Perizoma, molecular data
Orthonama inflexa (Dognin, 1914) comb.n. Coenocalpe, Scotopteryx, external morphology
Spargania coeruleopicta Warren, 1908 comb.n. Perizoma, external morphology
Spargania emmelesiata (Snellen, 1874) comb.n. Cidaria, Perizoma, external morphology
Spargania zenobia (Thierry-Mieg, 1893) comb.n. Cidaria, Perizoma, molecular data and external morphology
Tribe changes Genus
Hydriomenini to Cidariini Ceratodalia Packard, 1876

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ARTHROPOD SYSTEMATICS & PHYLOGENY — 77
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Trichopterygini to Dyspteridini Celonoptera Lederer, 1862 *
Trichopterygini to Dyspteridini Heterophleps Herrich-Schäer, [1854]
unassigned to Dyspteridini Chlorotimandra Butler, 1882
unassigned to Trichopterygini Aloba Warren, 1895
unassigned to Ennadini “Hagnagora” mesenata Felder & Rogenhofer, 1875
unassigned to Eudulini Graphidipus Herrich-Schäer, [1855]
unassigned to Eudulini Crocypus Herrich-Schäer, [1855]
unassigned to Heterusiini Spiloctenia Warren, 1897
unassigned to Trichopterygini Anomozela Fletcher, 1979
unassigned to Trichopterygini Isosauris Warren, 1894
unassigned to Trichopterygini Synpelurga Butler, 1882
unassigned to Trichopterygini Tatosoma Butler, 1874*
unassigned to Trichopterygini Lobidiopteryx Warren, 1902*
unassigned to Trichopterygini Episteira Warren, 1899*
unassigned to Stamnodini Pseudopsodos Thierry-Mieg, 1903
unassigned to Stamnodini Scordyliodes Thierry-Mieg, 1903
Xanthorhoini to Cataclysmini Zenophleps Hulst, 1896
Species proposed to be excluded from genera = incertae sedis Tribe assignment, country, originally described in
“Anticlea” oculisigna (Prout, 1923) (analysed) Chrismopterygini, Argentina, Larentia
“Anticlea” badiiplaga (Fletcher, 1953) (not analysed) unknown, Argentina, Earophila
“Anticlea” chillanensis (Butler, 1882) (type image checked) unknown, Chile, Larentia
“Anticlea” crepusculata (Fletcher, 1953) (not analysed) unknown, Argentina, Earophila
”Epirrhoe” psyroides Herbulot, 1988 (analysed) Plemyriopsini, Bolivia, Epirrhoe
“Heterusia” picata Dognin, 1904 (analysed) unnamed tribe, Ecuador, Heterusia
“Heterusia” plagia (Druce, 1893) (analysed) unnamed tribe, Ecuador, Trochiodes
“Heterusia” adventa Prout, 1934 (type image checked) Ennadini, Argentina, Scordylia
“Heterusia” barrioso Ureta, 1956 (type image checked) Ennadini, Chile, Heterusia
“Nebula” adela (Butler, 1893) (type image checked) unknown, Chile, Cidaria
“Nebula” aleucidia (Butler, 1882) (type image checked) Plemyriopsini, Chile, Cheimatobia
“Nebula” bellissima (Butler, 1893) (not analysed) unknown, Chile, Spargania
“Nebula” corticalis (Butler, 1882) (type image checked) unknown, Chile, Anticlea
“Nebula” ceres (Butler, 1882) (type image checked) unknown, Chile, Cidaria
“Nebula” cylon (Druce, 1893) (type image checked) unnamed lineage, Mexico, Hammaptera
“Nebula” cynthia (Butler, 1882) (analysed) unnamed lineage, Chile, Cidaria
“Nebula” decipiens (Butler, 1882) (type image checked) unnamed lineage, see emilia
“Nebula” detritaria (Staudinger, 1899) (not analysed) unknown, Chile, Coremia
“Nebula” diana (Butler, 1882) (type image checked) unnamed lineage, Chile, Cidaria
“Nebula” dubia (Butler, 1882) (type image checked) unknown, Chile, Camptogramma
“Nebula” emilia (Butler, 1882) (analysed) unnamed lineage, Chile, Cidaria
“Nebula” flexuosa (Dognin, 1914) (type image checked) unknown, Colombia, Anticlea
“Nebula” ignipennis (Butler, 1882) (type image checked) Ennadini, Chile, Ochyria
“Nebula” macidata (Felder & Rogenhofer, 1875) (type image checked) unknown, Chile, Cidaria
“Nebula” mathewi (Butler, 1883) (type image checked) unknown, Chile, Psaliodes
“Nebula” misera (Butler, 1882) (type image checked) unknown, Chile, Cidaria
“Nebula” mutabilis (Mabille, 1885) (not analysed) unknown, Chile, Cidaria
“Nebula” pusilla (Butler, 1882) (type image checked) unknown, Chile, Chalastra?
“Perizoma” ablata (Hulst, 1896) (not analysed) unknown, USA, Hydriomena
“Perizoma” actuata (Pearsall, 1909) (not analysed) unknown, USA, Mesoleuca
“Perizoma” alaskae (Hulst, 1896) (not analysed) unknown, USA, Coenocalpe
“Perizoma” amplata Warren, 1904 (type image checked) Plemyriopsini, amplata-group, Peru, Perizoma
“Perizoma” alumna (Prout, 1925) (analysed)* Larentiini, South Africa, Ortholitha
“Perizoma” anguliferata (Maassen, 1890) (not analysed) unknown, Bolivia, Cidaria
“Perizoma” apiceflava (Prout, 1910) (not analysed) unknown, Peru, Perizoma?
“Perizoma” aspersa Dognin, 1904 (type image checked) possibly Ennadini, Ecuador, Perizoma
“Perizoma” aurantaria (Jones, 1921) (not analysed) unknown, Brazil, Psaliodes
Table 1 continued.

B et al.: Phylogeny of New World Geometridae
468
“Perizoma” aureoviridis Warren, 1904 (type image checked) unknown, Peru, Perizoma
“Perizoma” baptopennis (Dyar, 1916) (type image checked) unknown, Mexico, Anapalta
“Perizoma” bogotata (Walker, 1862) (type image checked) unknown, Colombia, Cidaria
“Perizoma” brunneopicta Dognin, 1913 (type image checked) unknown, Colombia, Perizoma
“Perizoma” caeruleosecta (Prout, 1916) (type image checked) unknown, Peru, Hammaptera
“Perizoma” carnepicta Warren, 1905 (type image checked) unknown, Peru, Perizoma
“Perizoma” egena (Bastelberger, 1911) (not analysed) unknown, Peru, Anapalta
“Perizoma” carnetincta Dognin, 1911 (type image checked) unknown, Colombia, Perizoma
“Perizoma” cinereolimitata (Thierry-Mieg, 1892) (type image checked) unknown, Colombia, Cidaria
“Perizoma” complicata Dognin, 1911 (type image checked) unknown, Colombia, Perizoma
“Perizoma” constellata Dognin, 1913 (type image checked) unknown, Colombia, Perizoma
“Perizoma” costiguttata (Hulst, 1896) (not analysed) unknown, USA, Hydriomena
“Perizoma” cretinotata Bastelberger, 1907 (analysed) Plemyriopsini, amplata-group, Peru, Perizoma
“Perizoma” curvisignata Warren, 1909 (type image checked) Plemyriopsini, amplata-group, Peru, Perizoma
“Perizoma” curvilinea curvilinea (Hulst, 1896) (not analysed) unknown, Canada, Hydriomena
“Perizoma” occidens (Hulst, 1898) (not analysed) unknown, USA, Hydriomena
“Perizoma” curvilinea foxi (Wright, 1924) (not analysed) unknown, USA, Venusia
“Perizoma” custodiata (Guenee, [1858]) (not analysed) unknown, USA, Eubolia
“Perizoma” carnata (Packard, 1874) (not analysed) unknown, USA, Phibalapteryx
“Perizoma” carneata (Packard, 1876) (not analysed) unknown, USA, Ochyria
“Perizoma” gueneeata (Packard, 1876) (not analysed) unknown, USA, Ochyria
“Perizoma” polygrammata (Hulst, 1896) (not analysed) unknown, USA, Coenocalpe
“Perizoma” cyrtozona Prout, 1922 (not analysed) Plemyriopsini, amplata-group, Colombia, Perizoma
“Perizoma” diltilla (Dyar, 1913) (type image checked) unknown, Peru, Epirrhoe
“Perizoma” discors (Warren, 1901) (type image checked) unknown, Peru, Epirrhoe?
“Perizoma” epictata Barnes & McDunnough, 1916 (not analysed) unknown, USA, Perizoma
“Perizoma” eudoxia Prout, 1934 (type image checked) unknown, Colombia, Perizoma
“Perizoma” fallax Warren, 1905 (type image checked) unknown, Peru, Perizoma
“Perizoma” fractifascia Dognin, 1911 (type image checked) unknown, Colombia, Perizoma
“Perizoma” grandis (Hulst, 1896) (not analysed) unknown, USA, Eucymatoge
“Perizoma” grandis saawichata (Swett, 1915) (not analysed) unknown, Canada, Hydriomena
“Perizoma” herrichiata (Snellen, 1874) (not analysed) unknown, Colombia, Opisogonia
“Perizoma” iduna Prout, 1910 (not analysed) unknown, Argentina, Perizoma?
“Perizoma” illimitata Prout, 1922 (type image checked) unknown, Peru, Perizoma
“Perizoma” impromissata (Walker, 1862) (type image checked) unknown, Uruguay, Ypsipetes?
“Perizoma” corticeata (Walker, [1863]) (type image checked) unknown, Uruguay, Camptogramma
“Perizoma” fasciolata Warren, 1897 (type image checked) unknown, Paraguay, Perizoma
“Perizoma” muscosata Warren, 1900 (type image checked) unknown, Argentina, Perizoma
“Perizoma” ochritincta Warren, 1905 (type image checked) unknown, Mexico, Perizoma
“Perizoma” puella Prout, 1910 (not analysed) unknown, unknown, Perizoma
“Perizoma” interlauta Warren, 1907 (not analysed) unknown, Peru, Perizoma
“Perizoma” mirifica Warren, 1904 (not analysed) Plemyriopsini, amplata-group, Peru, Perizoma
“Perizoma” mixticolor Dognin, 1913 (type image checked) possibly Euphyiini, Colombia, Perizoma
“Perizoma” mollis Dognin, 1913 (type image checked) unknown, Eupithecia?; Colombia, Perizoma
“Perizoma” nigrostipata Dognin, 1913 (type image checked) unknown, Colombia, Perizoma
“Perizoma” obtusa (Warren, 1907) (type image checked) unknown, Peru, Opisogonia
“Perizoma” ochreata (Grossbeck, 1910) (not analysed) unknown, USA, Mesoleuca
“Perizoma” oxygramma (Hulst, 1896) (not analysed) unknown, USA, Coenocalpe
“Perizoma” tahoensis Barnes & McDunnough, 1916 (not analysed) unknown, USA, Perizoma
“Perizoma” pastoralis (Butler, 1882) (type image checked) unknown, Chile, Ypsipetes
“Perizoma” pecata (Dognin, 1893) (type image checked) unknown, Ecuador, Cidaria
“Perizoma” perryi Rindge, 1973 (not analysed) unknown, Ecuador, Perizoma?
“Perizoma” persectata (Maassen, 1890) (type image checked) unknown, Ecuador, Cidaria
“Perizoma” plumbinotata (Warren, 1904) (type image checked) unknown, Peru, Gagitodes
“Perizoma” pravata (Dognin, 1900) (type image checked) possibly Euphyiini, Bolivia, Eucosmia
Table 1 continued.

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ARTHROPOD SYSTEMATICS & PHYLOGENY — 77
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most lineage of Cidariini, although with poor support. A
formal description will be given by ÕuNap et al. (in prep).
4.2.17. Cidariini. The Nearctic taxa Ceratodalia gue-
nea ta Packard, 1876 and Trichodezia albovittata Guenée,
[1858] clearly belong to this clade. Both are type species
of their genera, respectively. We transfer Ceratodalia
Packard, 1876 from Hydriomenini to Cidariini (Table 1).
Trichodezia Warren, 1895 was assigned to Cidariini by
viidaLepp (1996, 2011) already. Our results support this
view but not the assignment to Asthenini by FergusoN
(1983) and pohL et al. (2015, 2018).
4.2.18. Scotopterygini. We are not aware of any New
World members of this tribe. We transfer the South Af-
rican taxon bitrita Felder & Rogenhofer, 1875 from La-
rentia Treitschke, 1825 to Scotopteryx Hübner, [1825],
and the South African taxon epipercna Wehrli, 1931 from
Perizoma to Scotopteryx (Table 1).
4.2.19. Euphyiini-Xanthorhoini complex. The follow-
ing six clades form a well supported lineage, and we
considered other systematic options, i.e. either the fusion
into a single large tribe Euphyiini, or a split into Euphyii-
ni + unnamed clade + Xanthorhoini as well. We opted for
a solution of ve named clades and one unnamed clade,
as Cataclysmini are a clearly distinct group according to
the morphology of both male and female genitalia (for
details see hausmaNN & viidaLepp 2012).
4.2.19.1. Unnamed clade. The node supporting this
clade as sister to Euphyiini is not strongly supported
(SH-like = 28.1, UFBoot2 = 71). Its phylogenetic posi-
tion and taxonomic rank thus requires further study.
The clade comprises Disclisioprocta Wallengren, 1861
(assigned to Xanthorhoini by Pohl et al. 2018) and un-
assigned Ptychorrhoe Warren, 1900. Morphology (e.g.
genitalia) of the widespread New World species D. stel-
lata (Guenée, [1858]) clearly shows that it is congeneric
with the two valid Old World taxa (our sample is D. na-
talata Walker, 1862) (A.H., unpublished data). It seems,
however, uncertain whether the type species of Ptychor-
rhoe, P. rayada Dognin, 1893, is actually congeneric
with P. blosyrata (Guenée, [1858]) because the wing pat-
tern of these two species differ substantially (see photos
in Electronic Supplement File 4).
“Perizoma” camptogrammaria Warren, 1907 (not analysed) unknown, Peru, Perizoma
“Perizoma” quadriplaga Dognin, 1911 (type image checked) unknown, Colombia, Perizoma
“Perizoma” renitens Prout, 1910 (type image checked) unknown, Peru, Perizoma?
“Perizoma” rostrinotata Dognin, 1913 (type image checked) unknown, Colombia, Perizoma
“Perizoma” sordescens Dognin, 1908 (type image checked) unknown, Peru, Perizoma
“Perizoma” spilophylla Prout, 1934 (type image checked) unknown, Argentina, Perizoma
“Perizoma” strictifascia Warren, 1907 (type image checked) unknown, Peru, Perizoma
“Perizoma” tenuisecta Prout, 1934 (type image checked) unknown, Argentina, Perizoma
“Perizoma” vacillans (Warren, 1905) (type image checked) unknown, Bolivia, Antepirrhoe
“Perizoma” vacillans tolimensis Prout, 1922 (type image checked) unknown, Colombia, Perizoma
“Perizoma” venisticta (Dognin, 1912) (type image checked) probably Plemyriopsini, Alydda; Colombia, Psaliodes
“Perizoma” versatilis Dognin, 1911 (type image checked) Plemyriopsini, amplata-group; Colombia, Perizoma
“Perizoma” vireonaria (Maassen, 1890) (analysed) Plemyriopsini, Ecuador, Cidaria
“Perizoma” virescentaria (Maassen, 1890) (not analysed) unknown, Bolivia, Thalassodes
“Psaliodes” picta Warren, 1904 (analysed) Plemyriopsini, Peru, Psaliodes
“Larentia” irma Prout, 1923 (analysed) unknown, Argentina, Larentia
“Larentia” macerata (Felder & Rogenhofer, 1875) (analysed) Ennadini, Chile, Heterusia?
“Larentia” albifilata Walker, [1863] (not analysed) unknown, Venezuela, Larentia
“Larentia” baliata Herrich-Schäer, 1870 (not analysed) unknown, Cuba, Larentia
“Larentia” danae (Druce, 1893) (type image checked) unknown, Mexico, Eubolia
“Larentia” horismeata Fletcher, 1953 (type image checked) unknown, Argentina, Larentia
“Larentia” lineolaria Blanchard, 1852 (not analysed) unknown, Chile, Larentia
“Larentia” omphacina Dognin, 1901 (not analysed) likely Ennadini, Brazil, Larentia
“Larentia” scarata (Felder & Rogenhofer, 1875) (type image checked) likely Ennadini, Chile, Fidonia
“Larentia” subgaliata Herrich-Schäer, 1870 (not analysed) unknown, Cuba, Larentia
“Scopteryx” ferridotata Walker, [1863]* (analysed) Larentiini, South Africa, Eubolia
“Stamnodes” eludens (Warren, 1908) (type image checked) Stamnodini, Peru, Marmopteryx
“Stamnodes” instar instar (Dognin, 1904) (type image checked) Stamnodini, Peru, Cophocerotis
“Stamnodes” instar casta (Dognin, 1904) (not analysed) Stamnodini, Peru, Cophocerotis
“Stamnodes” uniformata (Warren, 1877) (type image checked) Stamnodini, Argentina, Carisa
“Stamnodes” unilineata (Walker, 1867) (not analysed) unknown, Colombia, Tora
Table 1 continued.

B et al.: Phylogeny of New World Geometridae
470
4.2.19.2. Euphyiini. Results show that Euphyia Hüb-
ner, [1825] is present both in the Neotropical region and
in the Holarctic region. We transfer the species tricol-
orata Dognin, 1902 from Xanthorhoe Hübner, [1825] to
Euphyia (Table 1). Our analysis also includes Oligopleu-
ra Herrich-Schäffer, [1855] (Fig. 3I) and Hammaptera
Herrich-Schäffer, [1855], the latter for the rst time in a
molecular phylogenetic analysis.
4.2.19.3. Pterocyphini Brehm, Murillo-Ramos &
Õunap, new tribe. — Type genus. Pterocypha Herrich-
Schäffer, [1855] (Fig. 3J). — Material examined and
phylogeny. This clade comprises Pterocypha, Obila
Walker, 1869, and Archirhoe Herbulot, 1951. Our cur-
rent knowledge suggests that Pterocyphini are possibly
restricted to the New World. We analysed the type spe-
cies of Pterocypha, gibbosaria Herrich-Schäffer, [1855].
Branch support values from the IQ-TREE analyses
clearly conrm the monophyly of this clade (SH-like =
100, UFBoot2 = 99). — Molecular evidence. The tribe
is characterized by DNA sequence data from the follow-
ing seven gene regions (exemplar P. gibbosaria, vouch-
er Br-Geo-0059 from Brazil, illustrated in Electronic
Supplement File 5): ArgK (MK738221), Ca-ATPase
(MK738618), COI (MK739110), EF1a (MK739723),
Nex9 (MK741384), RPS5 (MK741726). Wingless
(MK742188). — Morphology. External features of an-
alysed species are illustrated in Fig. 3J and Electronic
Supplement File 4. Further detailed morphological anal-
ysis is required to identify potential diagnostic features.
— Remarks and taxonomic changes. We transfer Ar-
chirhoe from Hydriomenini and Obila and Pterocypha
from unassigned to Pterocyphini (Table 1).
4.2.19.4. Xanthorhoini. Our analysis include Hela-
stia Guenée, 1868, Orthonama Hübner, [1825] (Fig. 3K),
and Xanthorhoe Hübner, [1825], the latter genus in-
cluding representatives from both the New and the Old
World. Judged by wing morphology, we transfer the tax-
on inexa Dognin, 1914 from Scotopteryx to Orthonama
(illustrated in Electronic Supplement File 4, Table 1).
4.2.19.5. Cataclysmini. Our analysis includes Cata-
clysme Hübner, [1825], Phibalapteryx Stephens, 1829,
and Zenophleps Hulst, 1896, the latter being transferred
to Cataclysmini from Xanthorhoini (Table 1). Zeno-
phleps is an exclusively Nearctic genus.
4.2.19.6. Epirrhoini, stat.rev. We revive pierce’s
(1914) Epirrhoinae (which comprised both Epirrhoe
Hübner, [1825] and Catarhoe Herbulot, 1951 in his
treatment) at the tribe level as Epirrhoini. Since “Euphy-
ia” sturnularia Herrich-Schäffer, [1855] is misplaced,
the genus should be listed – ad interim – with quotation
marks (illustrated in Fig. 3L). Herewith, we include stur-
nularia in Epirrhoini as well as the African Mimoclystia
Warren, 1901 (Table 1). Further study must reveal the
relationship between Neotropical sturnularia and the
Old World genera Catarhoe, Mimoclystia and Epirrhoe
Hübner, [1825].
4.2.20. Larentiini complex. All following Larentiinae
taxa form a large, rather well supported clade (SH-like
= 87, UFBoot2 = 79) with a dominance of Neotropical
taxa. Palaearctic species are represented in our dataset
with one or a few species in Larentiini, Ennadini, Hydri-
omenini and Stamnodini – but more sampling in the Old
World is required. Genetic divergences between the line-
ages proposed as tribes Heterusiini, Cophocerotini, Er-
ateinini, Erebochlorini and Stamnodini are rather small,
and all these tribes could potentially be synonymized
with Hydriomenini. However, many of the currently
recognized tribes are rather species-rich (Hydriomenini,
Heterusiini, Erateinini, Stamnodini), and there is consid-
erable diversity in the external morphology of the moths
(Fig. 3M – T), possibly related to several switches to
diurnal lifestyle (in particular the genera Hagnagora,
Heterusia Hübner, [1825] and Erateina Doubleday,
1848), see Brehm & suLLivaN (2005).
4.2.20.1. Rhinurini Brehm, Murillo-Ramos & Õu-
nap, new tribe. — Type genus. Rhinura Warren, 1904
(Fig. 3M). — Material examined and phylogeny. Rhin-
ura near populonia (Druce, 1893) is sister to Haplolabi-
da inaequata (Prout, 1935). Rhinurini are sister to a large
as semblage including e.g. Larentiini and Stamnodini
(Fig. 2). Branch support values from the IQ-TREE anal-
yses strongly conrm the monophyly of this clade (SH-
like = 99.9, UFBoot2 = 100). — Molecular evidence.
The tribe is characterized by DNA sequence data from
the following seven gene regions (exemplar Rhinura near
populonia, voucher EO1166 from Ecuador, illustrated
in Electronic Supplement File 5): CAD (MK738977),
COI (MK739207), EF1a (MK739789), GAPDH
(MK740423), IDH (MK740793), RPS5 (MK741786),
Wingless (MK742294). — Morphology. External fea-
tures of analysed species are illustrated in Fig. 3M and
Electronic Supplement File 4. Further detailed morpho-
logical analysis is required to identify potential diagnos-
tic features. — Remarks and taxonomic changes. This
tribe currently comprises only two genera from differ-
ent continents. More taxon sampling is required to show
whether more African genera might belong to Rhinurini.
We also transfer the monotypic Neotropical genus Uro-
calpe Warren, 1904 to Rhinurini, based on the wing pat-
tern that is very similar to that of Rhinura (illustrated in
Electronic Supplement File 4, Table 1). Comprehensive
further morphological and molecular study is required.
4.2.20.2. Larentiini. Our analysis comprises no spe-
cies from the New World in this tribe. Since the follow-
ing species are misplaced, the respective genus should
be listed – ad interim – with quotation marks: Old World
“Perizoma” alumna (Prout, 1925) (see Perizomini) and
“Scotopteryx” ferridotata (Walker, [1863]) (see Scoto-
pterygini) (Table 1). An integrative revision of the afore-
mentioned taxa is required to provide new nomenclato-
rial combinations.
4.2.20.3. Unnamed clade. This clade comprises four
species: One is unidentied, one is “Larentia” near irma

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ARTHROPOD SYSTEMATICS & PHYLOGENY — 77
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Prout, 1923 from Chile (illustrated in Electronic Supple-
ment File 4), and two are wrongly assigned to Heterusia
Hübner, [1825]. Since “Heterusia” picata Dognin, 1904
and “Heterusia” plagia Druce, 1893 are misplaced, the
genus should be listed – ad interim – with quotation
marks (illustrated in Electronic Supplement File 5, Table
1). Since Heterusia comprises more species with plagia-
like habitus (e.g. polymela Druce, 1893 and prusa Druce,
1893) it is likely to be non-monophyletic and requires
revision. A formal description of a new tribe is not per-
formed because the taxonomy of the examined material
is still unclear.
4.2.20.4. Ennadini Brehm, Murillo-Ramos & Õu-
nap, new tribe. — Type genus. Ennada Blanchard,
1852 (Fig. 3N). — Material examined and phylogeny.
The clade comprises mostly species misplaced in the
genera Hagnagora, Larentia, Perizoma, and Nebula. It
also comprises Ennada and three species of Spargania
Guenée, [1858], a genus previously assigned to Larentii-
ni (hausmaNN & viidaLepp 2012). The type species of the
genus, S. magnoliata Guenée [1858] from North Ameri-
ca, is not included in the analysis. However, S. magnolia-
ta and Palaearctic S. luctuata ([Denis & Schiffermüller],
1775) are actually congeners (E.Õ. and Andro Truuverk,
unpublished). Branch support values from the IQ-TREE
analyses conrm the monophyly of this clade (SH-like
= 97.5, UFBoot2 = 71). — Molecular evidence. The
tribe is characterized by DNA sequence data from the
following ve gene regions (exemplar Ennada pellicata
Felder & Rogenhofer, 1875, voucher Ch-Geo-0010 from
Chile, illustrated in Electronic Supplement File 5): COI
(MK739121), EF1a (MK739731), MDH (MK740958),
RPS5 (MK741737), Wingless (MK742200). — Mor-
phology. External features of all analysed species are il-
lustrated in Fig. 3N and Electronic Supplement File 4.
Further detailed morphological analysis is required to
identify potential diagnostic features. — Remarks and
taxonomic changes. Ennada species were revised by
parra & aLvear (2009). All Chilean species of the gen-
era Hagnagora, Larentia and Nebula are misplaced to the
respective genera, which therefore should be listed – ad
interim – with quotation marks (Table 1). “Hagnagora”
mesenata Felder & Rogenhofer, 1875 was already pro-
posed to be excluded from Hagnagora by Brehm (2015).
An integrative revision of the mentioned taxa is required
to provide new nomenclatorial combinations. We for-
mally transfer the taxa coeruleopicta Warren, 1908 and
emmelesiata Snellen, 1874 to Spargania because they are
apparently closely related to S. zenobia (Table 1).
4.2.20.5. Hydriomenini. This lineage includes ve
analysed taxa, among them two European and one Neo-
tropical species of Hydriomena Hübner, [1825]. Mono-
typic Cyclica Grote, 1882, is nested within Hydriom-
ena and is therefore synonymized (Table 1). Ersephila
prema Druce, 1893 (Fig. 3O) is sister to the other four
analysed species. Ersephila Hulst, 1896 is retained but it
remains to be shown in further studies whether Ersephila
is monophyletic.
4.2.20.6. Heterusiini. The clade includes the Neo-
tropical genera Heterusia Hübner, [1831] and Spilocte-
nia Warren, 1897, both represented by their type species
in the analysis (Fig. 3P: H. quadruplicaria Geyer, 1832).
Spiloctenia is transferred from unassigned to Heterusiini
which is plausible also from wing morphology (illustrat-
ed in Electronic Supplement File 4, Table 1).
4.2.20.7. Cophocerotini Brehm, Murillo-Ramos &
Õunap, new tribe. — Type genus. Cophocerotis War-
ren, 1895 (Fig. 3Q). — Material examined and phylo-
geny. The clade includes Cophocerotis and Hagnagora.
We chose Cophocerotis for naming the tribe because
there are morphological differences between the ana-
lysed Hagnagora species and the type species H. buck-
leyi Druce, 1885 (Brehm 2015), questioning their status
as congeners. Branch support values from the IQ-TREE
analyses conrm the monophyly of Cophocerotini (SH-
like = 100, UFBoot2 = 100). — Molecular evidence.
The tribe is characterized by DNA sequence data from
the following seven gene regions (exemplar Cophoce-
rotis costinotata Warren, 1908, voucher gb-ID-19302
from Ecuador, illustrated in Electronic Supplement File
5): COI (MK739304), EF1a (MK739882), GADPH
(MK740547), MDH (MK741090), Nex9 (MK741533),
RpS5 (MK741896), Wingless (MK742433). — Mor-
phology. External features of analysed species are il-
lustrated in Fig. 3Q and Electronic Supplement File 4.
Further detailed morphological analysis is required to
identify potential diagnostic features. — Remarks and
taxonomic changes. The analysis of the phylogenetic
placement of Hagnagora buckleyi urgently requires to be
studied. If it turns out not to be congeneric with other
species currently assigned to Hagnagora, those will need
to be transferred to another genus.
4.2.20.8. Erateinini. The clade includes the genera
Erateina Doubleday, 1848 (Fig. 3R) and Anemplocia
Warren, 1905. It also includes the taxon “Trocherateina”
Prout, but according to scoBLe (1999), this name had not
been published before. As the most straight-forward way
towards a system of monophyla, we synomynize Anem-
plocia, and transfer all species currently assigned to una-
vailable “Trocherateina” to Erateina (Table 1).
4.2.20.9. Erebochlorini Brehm, Murillo-Ramos &
Õunap, new tribe. — Type genus. Erebochlora Warren,
1895 (Fig. 3S). — Material examined and phylogeny.
This clade includes the three Neotropical genera Erebo-
chlora, Cirrolygris Warren, 1895, and Deinoptila Warren,
1900. Branch support values from the IQ-TREE analy-
ses clearly support the monophyly of this clade (SH-like
= 100, UFBoot2 = 100). — Molecular evidence. The
tribe is characterized by DNA sequence data from the
following eight gene regions (exemplar Erebochlora
near tesserulata Felder & Rogenhofer, 1875, voucher
GB-CR-1218, from Costa Rica, illustrated in Electron-
ic Supplement File 5): ArgK (MK738311), Ca-ATPase
(MK738705), COI (MK739228), EF1a (MK739809),
GAPDH (MK740443), MDH (MK741007), Nex9

B et al.: Phylogeny of New World Geometridae
472
(MK741444), Wingless (MK742314). — Morphology.
External features of all analysed species are illustrated
in Fig. 3S and Electronic Supplement File 4. Further de-
tailed morphological analysis is required to identify po-
tential diagnostic features. — Remarks and taxonomic
changes. Priapodes Warren, 1895 was described by War-
ren only because of prolonged palpi; he stated that “oth-
erwise the types of the two genera [Priapodes and Ere-
bochlora] are supercially wonderfully alike“. Since the
two genera indeed have a very similar habitus, size and
wing pattern (illustrations in Electronic Supplement File
5), we synonymize Priapodes with Erebochlora (Table
1). We suggest to place the genus of “Erebochlora” api-
ciava Dognin, 1892 – ad interim – in quotation marks,
as it was recovered apart form its congeners and sister to
a clade comprising Cirrolygris and Deinoptila (Table 1).
Further study of Erebochlora is required because it cur-
rently is a paraphyletic assemblage.
4.2.20.10. Stamnodini. Our analysis includes the Pa-
laearctic type species of Stamnodes Guenée, [1858], viz.
S. pauperaria Eversmann, 1848 and the Nearctic S. to-
pazata Strecker, 1899 (ÕuNap et al. 2016). We also ana-
lysed the taxon triangularia Bartlett-Calvert, 1891. As a
step towards a natural system, we revive Synneuria Ma-
bille, 1885, from synonymy with Stamnodes and trans-
fer three more species that were originally described in
Synneuria back to this genus (Table 1). The Neotropical
species anthocharidaria Oberthür, 1881 (Fig. 3T) is sister
to Callipia Guenée, [1858]. We therefore transfer it from
Stamnodes to Callipia (Table 1). C. anthocharidaria has a
similar general habitus and wing shape as Callipia species
(Brehm 2018), although its wing pattern is largely reduced
and it is considerably smaller than all previously known
Callipia species (illustrations in Electronic Supplement
File 4). Four South American “Stamnodes” species are
misplaced, and their genus name should be listed – ad in-
terim – with quotation marks. Moreover, we transfer the
genera Pseudopsodos Thierry-Mieg, 1903, and Scordyli-
odes Thierry-Mieg, 1903 to Stamnodini (Table 1).
4.3. Archiearinae Fletcher, 1953
Archiearinae are represented, in our analysis, by four
species, including two Nearctic taxa. Nearctic Archiearis
infans Möschler, 1862 clusters together with Palaearc-
tic A. parthenias (Linnaeus, 1761) while Leucobrephos
brephoides (Walker, 1857) is sister to Archiearis Hüb-
ner, [1823] + Boudinotiana Hübner, [1803] clade. The
sister relationship of Leucobrephos Grote, 1874 with
Archiearis + Boudinotiana is plausible and well con-
rmed by morphology (müLLer et al. 2019). The Aus-
tralian genera Dirce Prout, 1910 and Acalyphes Turner,
1926 were transferred from Archiearinae to Ennominae
earlier (YouNg 2006; muriLLo-ramos et al. 2019). Rep-
resentatives from Central and South America await fur-
ther study, i.e. Caenosynteles Dyar, 1912 (one species),
Archiearides Fletcher, 1953 (two species), and Lachno-
cephala Fletcher, 1953 (one species). There is evidence
that at least Archiearides indeed belongs to Archiearinae
because of a “Archiearinae-like” tympanum (FLetcher
1953; cook & scoBLe 1992). On the other hand, the very
isolated distribution of the austral South American taxa
suggests possible convergence with Holarctic taxa due to
similar (diurnal) behaviour and resulting similar colour
patterns (illustrations in Electronic Supplement File 4).
4.4. Desmobathrinae Meyrick, 1886,
Oenochrominae Guenée, [1858],
Epidesmiinae Murillo-Ramos, Sihvonen
& Brehm, 2019
These subfamilies were treated in detail by muriLLo-ra-
mos et al. (2019). Six Neotropical genera in two separate
lineages belong to the Desmobathrinae: Zanclopteryx
Herrich-Schäffer, [1855] clusters together with Ozola
Walker, 1861. The second clade comprises Neotropical
Racasta Walker, 1861, Leptoctenopsis Warren, 1895,
Ophiogramma Hübner, [1831], Pycnoneura Warren,
1894 and Dolichoneura Warren, 1894 as sister to the
Indopacic genus Noreia Walker, 1861. There are no
representatives of the subfamilies Oenochrominae, Epi-
desmiinae and Orthostixinae from the New World in our
analysis. We could not study two monotypic New World
genera currently assigned to Oenochrominae, viz. Car-
mala Walker, [1863] and Cortixa Schaus, 1901. Carma-
la is unknown to us, and Cortixa comprises small and
slender-bodied moths that are more likely to belong to
Desmobathrinae than to Oenochrominae.
4.5. Geometrinae Stephens, 1829
See Fig. 4 for phylogenetic relationships, Fig. 5 for habitus pictures
of exemplary species, and Table 2 for proposed taxonomic changes.
By far most sampled New World Geometrinae taxa are
concentrated in the New World tribe Nemoriini, a group
recently studied in detail by viidaLepp (2017). Apart from
Nemoria Hübner, with its type species bistriaria Hüb-
ner, 1818 (Fig. 5A), our data conrm the assignment to
Nemoriini of the genera Assachlora Viidalepp & Lindt,
2012, Chavariella Pitkin, 1993, Dichorda Warren, 1900,
Hyalochlora Prout, 1912, Lissochlora Warren, 1900,
Neagathia Warren, 1897, Phrudocentra Warren, 1895,
Pyrochlora Warren, 1895, Rhodochlora Warren, 1894,
Tachyphyle Butler, 1881 and Tachychlora Prout, 1912. In
addition, our data suggest that the currently unassigned
genus Hydata Walker [1863] also needs to be transferred
to Nemoriini: (Table 2). viidaLepp (2017) discussed the
absence of a midrib of the last abdominal sternite of the
male as a basic nemoriine characteristic of Hydata and
Methydata Prout, 1933, but he also found possible other
synapomorphies linking them with Nemoriini.
The Synchlorini genus Synchlora Guenée, [1858] –
represented with its type species aerata (Fabricius, 1798)

473
ARTHROPOD SYSTEMATICS & PHYLOGENY — 77
(3) 2019
(Fig. 5B) – is nested within Nemoriini. muriLLo-ramos
et al. (2019) therefore synonymized Synchlorini with
Nemoriini. Our data show that Nemoria itself is not
monophyletic and requires revision which, however, is
beyond the scope of our study. Possible steps towards
a system of natural entities could include the transfer
of (presumably many) species around N. nigrisquama
(Dognin, 1904) (Fig. 5C) to Lissochlora (Fig. 5D) and
reviving one or more generic synonyms of Nemoria in
future studies. In our study, we recognize the studied spe-
cies nigrisquama and erina (Dognin, 1896) (Fig. 5E) as
misplaced in Nemoria. For these cases, we suggest that
the genus is listed – ad interim – with quotation marks
(Table 2). An integrative revision of the mentioned taxa
is required to provide new nomenclatorial combinations.
Only a few studied New World Geometrinae taxa be-
long to tribes other than Nemoriini. In the Hemitheini,
Lophochorista Warren, 1904 (Fig. 5F) denes the Lopho-
choristina (Lophochoristini in pitkiN 1996, Lophochoris-
titi in BaN et al. 2018), but its closest relatives are the
unassigned African genera Rhadinomphax Prout, 1912
and Adicocrita Prout, 1930 which we formally assign to
the subtribe Lophochoristina (Table 2). Two other New
World genera form a monophylum: Anomphax War-
ren, 1909 (Fig. 5G) and Oospila Warren, 1897. They
are not closely related to Lophochorista, and therefore
are not part of Lophochoristina (Table 2). The position
of Chloropteryx Hulst, 1896 (Fig. 5H) and Xerochlora
Ferguson, 1969 in Hemitheini/Hemitheina conrms pre-
vious results (pitkiN 1996). The North American genus
Dichordophora Prout, 1913 needs to be investigated in
future studies because it is representing the tribe Dichor-
dophorini (FergusoN 1969).
4.6. Ennominae Duponchel, 1845
See Fig. 6 for phylogenetic relationships, Fig. 7 for habitus pictures
of exemplary species, and Table 3 for proposed taxonomic changes.
Species from the New World are present in the large ma-
jority of Ennominae tribes (Fig. 6), and the subfamily is
doubtlessly the most species-rich subfamily in this region
(piktiN 2002). In addition to Palyadini and probably Na-
cophorini, three new tribes (see below: Euangeronini,
Oenoptilini, Pyriniini) comprise exclusively Neotropi-
cal genera. It remains to be shown whether further taxon
sampling in the Old World and Australasia will add taxa
from these regions.
4.6.1. Euangeronini Brehm, Murillo-Ramos & Sihvo-
nen, new tribe. — Type genus. Euangerona Butler,
1882 (Fig. 7A). — Material examined and phylogeny.
The tribe exclusively comprises taxa from austral South
America and is related to a clade comprising only Idial-
cis Warren, 1906, Gonodontini and Gnophini. None of
those are morphologically similar to Euangeronini (see
illustrations in pitkiN 2002 and müLLer et al. 2019). Fur-
ther analysed genera included in the tribe: Dectochilus
Butler, 1882, Malleco Rindge, 1971, and Odontothera
Nemoriini
20 3 16
Ornithospilini
Chlorodontoperini
Aracimini
unnamed clade
Timandromorphini
Geometrini
Comibaenini
Hemitheini
Crypsiphona
Pseudoterpnini
Xenozancla
Archaeobalbini
unnamed clade
6 1
2 1
5 3
5 3
2 1
2 1
18 5
12 5
2 1
17 10
1 1
2 1
5 3
12 5
0.2
2 1
Agathiini
Neohipparchini
Dysphanini
64 33 1 3 1
19.6/47
94.6/83
100/100
99.2/97
100/100
100/100
88.3/64
94.1/55
100/100
99/94
66.8/86
100/100
100/100
0/14
98/90 59.6/33
94.2/86
100/100
100/100
76.3/21
99.2/99
100/100
100/100
100/100
98.3/91
100/100
83.9/75
93.6/58
11.7/65
75.4/37
62.8/50
Fig. 4. Geometrinae tribe composition. Tribes with New World representatives marked in colour: light green Nearctic, dark green Neo-
tropical, yellowish green austral region; see also explanatory box in Fig. 1. Support values in blue colour (SH-like and UFBoot values).

B et al.: Phylogeny of New World Geometridae
474
Butler, 1882. We also examined “Opisogonia” difssata
Felder & Rogenhofer, 1875 and “Chlorochlydon” rino-
daria Fel der & Rogenhofer, 1875. Both are not conge-
neric with the type species of the respective genera (pit-
kiN 2002), see photos in Electronic Supplement File 1.
Chloroclydon Warren, 1894, is a junior synonym of
Herochroma Swinhoe, 1893, an Old World Geometrinae
genus. Branch support values from the IQ-TREE analy-
ses clearly conrm the monophyly of Euangeronini (SH-
like = 99.5, UFBoot2 = 96). — Molecular evidence.
The tribe is characterized by DNA sequence data from
the following ve gene regions (exemplar Euangerona
valdiviae Butler, 1882, voucher bo-chi-109 from Chile,
illustrated in Fig. 7A): Ca-ATPase (MK738586), COI
(MK739063), EF1a (MK739698), Nex9 (MK741345),
RPS5 (MK741700). — Morphology. External features
of analysed species are illustrated in Fig. 7A and Elec-
tronic Supplement File 5. Further detailed morphologi-
cal analysis is required to identify potential diagnostic
features. — Remarks and taxonomic changes. We pro-
visionally also assign Omaguacua Rindge, 1983 to Eu-
an ge ronini because it is similar to Dectochilus accord-
ing to its external morphology, however without dentate
forewing margins (illustrated in Electronic Supplement
File 5). See pitkiN (2002) for more information on the
included genera and species.
4.6.2. Unnamed clade. Idialcis Warren, 1906 (Fig. 7B),
is part of the Euangeronini-Gonodontini-Gnophini clade.
It is an independent lineage which might represent a
separate tribe and requires further study. Idialcis is trans-
ferred from Ennomini to unassigned (Table 3).
4.6.3. Gonodontini. This tribe is represented by two Old
World genera in our phylogeny and it is unlikely that
Gonodontini are represented in the New World. The type
genus of the tribe, Gonodontis Hübner, [1823], was not
included in the analysis.
4.6.4. Gnophini. This tribe comprises only a few New
World taxa in our analysis, namely Nearctic Euchlaena
Hübner, [1823], and Lytrosis Hulst, 1896. These were as-
signed to Angeronini by FergusoN (1983) but we follow
recent literature (e.g. skou & sihvoNeN 2015; BeLJaev
2016; müLLer et al. 2019; muriLLo ramos et al. 2019)
who considered Angeronini a junior synonym of Gno-
phini. The Chilean genus Neorumia Bartlett-Calvert,
1893 (see parra & vargas 2000) can be assigned to
Table 2. Taxonomic changes in Geometrinae at the level of tribes, genera and species, in alphabetical order. *Old World taxa.
From tribe x to tribe yGenus
unassigned to Nemoriini Hydata Walker, 1895
Genus From subtribe x to subtribe y
Rhadinomphax Prout, 1912* unassigned to Lophochoristina
Adicocrita Prout, 1930* unassigned to Lophochoristina
Anomphax Warren, 1909 Lophochoristina to unassigned
Oospila Warren, 1897 Lophochoristina to unassigned
Species proposed to be excluded from genera = incertae sedis Tribe assignment, country, originally described in genus
“Nemoria” nigrisquama Dognin, 1904 Nemoriini, Peru, Miantonota
“Nemoria” erina Dognin, 1896 Nemoriini, Ecuador, Achlora
A C DB
E G HF
Fig. 5. Illustrations of selected Neotropical Geometrinae taxa. A: Nemoriini, Nemoria bistriaria Hübner, 1818 (CNC580945). B: Nemori-
ini, Synchlora aerata (Fabricius, 1798) (CNC541241). C: Nemoriini, “Nemoria” nigrisquama (Dognin, 1904) (Pe-Geo-3142) D: Nemo-
riini, Lissochlora latuta (Dognin, 1898) (ID 18194). E: Nemoriini, “Nemoria” erina (Dognin, 1896) (AH7057). F: Hemitheini, Lopho-
chorista near curtifascia Prout, 1933 (GB-Geo-083). G: Hemitheini, Anomphax gnoma (bo_chi_433). H: Hemitheini, Chloropteryx sp.
(Pe-Geo-0614).

475
ARTHROPOD SYSTEMATICS & PHYLOGENY — 77
(3) 2019
Gnophini (Fig. 7C, Table 3). However, “Neorumia” gra-
cilis Bartlett-Calvert, 1893 was already excluded from
Neorumia by pitkiN (2002) and clusters in Ennomini.
4.6.5. Odontoperini. This clade comprises the type spe-
cies of Odontopera Stephens, 1831, Palaearctic O. biden-
tata Clerck, 1759, Henicovalva Krüger, 2017 from South
Africa, Nemeris Rindge, 1981 from North America and
the austral South American genera Dentinalia Heimlich,
1960, Macrolyrcea Butler, 1882, Mallomus Blanchard,
1852, Praeantarctia Heimlich, 1956, and Talca Rindge,
1971. All these genera (except for Odontopera) are trans-
ferred to Odontoperini (Table 3). The Azelinini are nest-
ed within Odontoperini and are therefore synonymized
with it (Table 3). Members of the tribe Odontoperini have
recently been classied as Ennomini of uncertain associ-
ation (skou & sihvoNeN 2015) or as Odontoperini (BeL-
Jaev 2016). Nepitia Walker, 1866, is nested within Pero
Herrich-Schäffer, 1855, and is therefore synonymized
with it (Fig. 7D, Table 3). A close relationship of Odonto-
perini, Azelinini and Nacophorini was already suggested
by hoLLoWaY (1994), with a possible synapomorphy of
16 setae on the proleg of the caterpillar on A6, and our
data provide strong support for this hypothesis.
4.6.6. Unnamed clade. Bryoptera Guenée, [1858] forms
a lineage of its own that cannot clearly be assigned to
either Odontoperini or Nacophorini. Bryoptera has previ-
ously been assigned to Boarmiini because of its Boarmii-
ni-like wing pattern (illustrated in Electronic Supplement
File 5), but pitkiN (2002) already noted that its genitalia
characters were unusual for that tribe. Bryoptera is trans-
ferred from Boarmiini to unassigned (Table 3), but it is
certainly part of the Odontoperini-Nacophorini clade.
Tephrosia vaga Dognin, 1895 is transferred from “Ectro-
pis” to Bryoptera (Fig. 7E, Table 3). Three Chilean spe-
cies previously assigned to Bryoptera were transferred to
Leucolithodes by parra & hormazáBaL (1993).
4.6.7. Nacophorini. Many species from all around the
world have been assigned to this tribe, but our results
clearly show that Nacophorini are a New World clade,
as previously stated by YouNg (2003). Nacophora Hulst,
1896 is a junior synonym of Phaeoura Hulst, 1896. The
Diptychini
Baptini+Theriini clade
unnamed clade
Plutodini+Palyadini
Apei+Epio+Anag+Hypo clade
Drepanogynini
Pyriniini, new tribe
Caberini
Cass+Abra+Euto+Maca clade
Boarmiini
44 22 2 2 6
3 1 2
2 1 1
8 1 6
10 7 3
19 4
6 4
15 1 6 4
23 19 6 8
296 115 26 16
Euangeronini, new tribe
Idialcis
Gonodontini
Gnophini
Odontoperini
Bryoptera
Nacophorini
Ennomini
Camp+Also+Wile+Colo clade
Declana
7 7
1 1
4 3
20 15 2 1
13 2 1 2 5
3 1
29 4 20 1
151 5 19 100 6
6 5
3 1
Oenoptilini, new tribe
3 3
0.2
97.3/60
87.4/62
92/88
87.2/86
100/100
100/100
94.2/84
100/99
87.6/80
97.3/90
39.4/39
92.2/81
100/100
100/100
99.9/94
92.6/95
54.1/56
100/100
100/99
100/100
93.8/61
99.9/100
100/97
100/99
100/100
73.1/36
26.6/52
100/98
96.5/65
100/99
90.3/87
98.1/88
100/100
0/12
42.3/39
81.8/72
100/100
99.6/100
99.5/96
48.8/51
Fig. 6. Ennominae tribe composition. Tribes with New World representatives marked in colour: light green Nearctic, dark green Neotropi-
cal, yellowish green austral region; see also explanatory box in Fig. 1. Camp+Also+Wile+Colo: Campaeini + Alsophilini + Wilemaniini
+ Prosopolophini clade. Ther+Bapt+Plut+Paly: Theriini + Baptini + Plutodini + Palyadini clade. Apei+Epio+Anag+Hypo: Apeirini +
Epionini + Anagogini + Hypochrosini clade. Cass+Abra+Euto+Maca: Cassymini + Abraxini + Eutoeini + Macariini clade. Support values
in blue colour (SH-like and UFBoot values).

B et al.: Phylogeny of New World Geometridae
476
type species of Nacophora is Phaeoura quernaria Smith,
1797 (Fig. 7F), and it is represented in our ana lysis. Na-
cophorini s.str. form a well supported clade, including (in
addition to P. quernaria) the genera Aethaloida McDun-
nough, 1920, Betulodes Thierry-Mieg, 1904, Gabriola
Taylor, 1904, Holochroa Hulst, 1896, and Thyrinteina
Möschler, 1890. We suggest a concept of Nacophorini
s.l., which at the moment includes New World gen-
era only, with several well supported clades. One clade
comprises the Neotropical genera Charca Rindge, 1983,
Chrysomima Warren, 1894, Cundinamarca Rindge, 1983,
Ischnopteris Hübner, [1823], Paradoxodes Warren, 1904,
Quillaca Rindge, 1983, Rucana Rindge, 1983, Stego-
theca Warren, 1900, an unnamed genus, and the Nearc-
tic Ceratonyx Guenée, [1858] – all already assigned to
Nacophorini. Another mostly Neotropical clade com-
prises Achagua Rindge, 1983, Cargolia Schaus, 1901,
Cidariophanes Warren, 1895, Eustenophasma Warren,
1897, Leucochesias Mabille, 1889, Nazca Rindge, 1983,
Oratha Walker, 1863, and Postazuayia Rindge, 1986.
Eustenophasma and Leucochesias are transferred from
unassigned to Nacophorini (Table 3). The colourful ge-
nus Catophoenissa Warren, 1894 and probably also the
similar unsampled genus Catocalopsis Rindge, 1971 – il-
A C DB
E G HF
I J K L
M N O P
Q R S T
Fig. 7. Illustrations of selected Neotropical Ennominae taxa. A: Euangeronini, new tribe, Euangerona valdiviae Butler, 1882 (bo_chi_109).
B: unnamed clade, Idialcis jacintha (Butler, 1882) (bo_chi_648). C: Gnophini, Neorumia gigantea Bartlett-Calvert, 1893 (bo_chi_167).
D: Odontoperini, Pero detractaria (Walker, 1866) comb.n. (Pe-Geo-0659). E: unnamed clade, Bryoptera vaga (Dognin, 1895) comb.n.
(gb-ID-22872). F: Nacophorini, Phaeoura quernaria (Smith, 1797) (CNC583542). G: Ennomini, “Nephodia” panacea Thierry-Mieg,
1892 (AH7126). H: Ennomini, “Perusia” viridis Warren, 1907 (Pe-Geo-0680). I: Prosopolophini, Himeromima aulis Druce, 1892 (Gu-
Geo-005). J: Diptychini, “Loxaspilates” torcida Dognin, 1900 (ID 19263). K: Oenoptilini, new tribe, Oenoptila mixtata Guenée, [1858]
(Br-Geo-0006). L: Palyadini, Ophthalmoblysis cinerea (Warren, 1909) (Vz-Geo-014). M: unassigned to tribe, Sericosema juturnaria
Guenée, [1858] (CNC533584). N: Pyriniini, new tribe, Pyrinia abditaria (Warren, 1905) (gb-ID-16080). O: Caberini, Aplogompha lafayi
(Dognin, 1899) (Pe-Geo-0545). P: unassigned to tribe, Hypometalla scintillans Warren, 1906 (Pe-Geo-0503). Q: Cassymini s.l., Leuciris
beneciliata Prout, 1910 (Pe-Geo-0545). R: Macariini, Macaria cardinea (Druce, 1893) (gb-ID-17469). S: Boarmiini, Perigramma famu-
lata (Felder & Rogenhofer, 1875) (Pe-Geo-3039). T: Boarmiini, “Synnomos” near apistrigata Warren, 1895 (Br-Geo-0008).

477
ARTHROPOD SYSTEMATICS & PHYLOGENY — 77
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lustrated in Electronic Supplement File 5 – does not have
a stable position in the tree. Australian “Nacophorini”
(see for instance YouNg 2006, 2008) are mostly nested
in Diptychini (see 4.6.11. below), and most African “Na-
cophorini” also belong to Diptychini or group together in
Drepanogynini (muriLLo-ramos et al. 2019).
4.6.8. Ennomini. Ennomini are sister to a lineage com-
prising all aforementioned Ennominae clades (Fig. 6). An
Old World lineage comprises both Ennomos Treitschke,
1825, and Ourapteryx Leach, 1814 (known from previ-
ous analyses, e.g. sihvoNeN et al. 2011). Both taxa are
closely related and we therefore agree with the synonymi-
zation of Ennomini and Ourapterygini by BeLJaev (2008),
as results are also consistent with earlier hypotheses (e.g.
sihvoNeN et al. 2011). All austral South American taxa,
i.e. Atopodes Warren, 1906, monotypic Gonogala Butler,
1882, Microclysia Butler, 1882, Syncirsodes Butler, 1882,
and “Tetracis” edmondsii Butler, 1882 (not congeneric
with true Tetracis Guenée, [1858]) are found in a single,
well supported clade. In contrast, Nearctic species are
widely scattered between 14 independent lineages within
Ennomini (see discussion of biogeography below). In our
analysis, Ennomini comprise the highest number of Neo-
tropical genera (more than 100, including undescribed
taxa) in a single tribe. Most genera were already assigned
to this tribe by pitkiN (2002) (as Ourapterygini), but many
also to the related informal Cratoptera-group, to the re-
lated Nephodiini, and a few to Caberini and Nacoph-
orini, while more than 60 were left unassigned by pitkiN
(2002). BeLJaev (2008) synonymized Nephodiini and the
Cratoptera group (misspelled as Catoptera group) with
the Ennomini, and assigned many previously unassigned
taxa to Ennomini. Our data clearly conrm the synonymy
of Nephodiini with Ennomini, but the broad-scale assign-
ment of taxa by BeLJaev (2008) requires additions and
adjustment in several cases (Table 3). Three species are
transferred from unassigned to Ennomini, but genus com-
binations are incertae sedis: “Acrotomia” mucia Druce,
1892, “Bassania” crocallinaria Oberthür, 1883 and “Cy-
phoedma” transvolutata Walker, 1860 (Table 3). In ad-
dition to the list provided by BeLJaev (2008), the genera
Acrosemia Herrich-Schäffer, [1855], Eutomopepla War-
ren, 1894, Microgonia Herrich-Schäffer, [1855], Polla
Herrich-Schäffer, [1855], and Tarma Rindge, 1983, are
transferred to Ennomini (Table 3). Brachyctenistis War-
ren, 1904 is transferred from Nacophorini to Ennomini
(Table 3).
Ennomini comprise several non-monophyletic gen-
era. For these cases, we suggest that the respective ge-
neric names should be listed – ad interim – with quota-
tion marks (pitkiN 2002; scoBLe 1999) and integrative
revisions of the mentioned taxa are required to provide
new nomenclatorial combinations. “Mychonia” gala-
nata Dognin, 1895 is not congeneric with type species-
related members of Mychonia Herrich-Schäffer, [1855]
(Table 3) and is also morphologically rather distinct (il-
lustrated in Electronic Supplement File 5). The same ap-
plies for “Isochromodes” analiplaga Warren, 1907 and
“Isochromodes” sabularia Dognin, 1900 which are not
congeneric with true Isochromodes Warren, 1894 – al-
ready suspected by pitkiN (2002) (Table 4). Nephodia
nubilaria Hübner, 1823 is the type species of the genus,
whereas many other taxa currently assigned to Nephodia
Hübner, 1823 most likely need to be transferred to other
genera in future studies. One option is the re-erection
of Nipteria Guenée, [1858] that would include the sam-
pled species panacea Thierry-Mieg, 1892 (Fig. 7G)
and presumably many other species currently assigned
to Nephodia. Sabulodes Guenée, [1858] is represented
by the type species of the genus, S. caberata Guenée,
[1858] which does not cluster together with “Sabulodes”
thermidora Thierry-Mieg, 1894. Enypia Hulst, 1896 is
nested within Sabulodes and is therefore synonymized
(Table 3). Nematocampa Guenée, [1858] is represented
by its North American type species N. resistaria Herrich-
Schäffer, [1856] (lamentaria Guenée, [1858] is a jun-
ior synonym). Neotropical N. angulifera Oberthür, 1883
clusters with N. resistaria. “Nematocampa” confusa
Warren, 1904 was already excluded from this genus by
pitkiN (2002) and “Nematocampa” falsa Warren, 1906
was suspected to be misplaced by pitkiN (2002) (Table
3). True Melinodes Herrich-Schäffer, [1855] are closely
related to Nematocampa and these genera share the ten-
tacle-like structures of the larvae (Brehm 2003), a poten-
tial synapomorphy of the lineage. Two species analysed
in this study, “Melinodes” fulvitincta Warren, 1905 and
“Melinodes” ignea Warren, 1907, were previously ex-
cluded from the genus by pitkiN (2002), and our results
support this view. Perusia Herrich-Schäffer, [1855] is
represented in our tree by three species, all of which re-
present independent lineages. “Perusia” zoma (Dognin,
1896) and “Perusia” viridis Warren, 1907 appear to be
misplaced (Fig. 7H, Table 3). The latter species shares
green pigmentation with the closely related genus Phyle
Herrich-Schäffer, [1855], a potential synapomorphy of
this lineage. “Eusarca” bogotata Snellen, 1874 does
not cluster together with Eusarca nemora Druce, 1892,
a species that closely resembles the type species of the
genus (Electronic Supplement File 5, Table 3). A similar
case are “Anisoperas” near tessellata Walker, [1863] and
Anisoperas proxima Dognin, 1914 (Electronic Supple-
ment File 5, Table 3).
4.6.9. Unnamed clade. A well supported clade is formed
by Declana Walker, 1858, from New Zealand, probably
representing an undescribed tribe.
4.6.10. Campaeini + Alsophilini + Wilemaniini + Pros-
opolophini clade. Each tribe is represented by only one
or two species in our analysis. The Central American spe-
cies Himeromima aulis Druce, 1892 (Fig. 7I) could not
be assigned to a tribe by pitkiN (2002), and our results
suggest that it either forms an own lineage or is part of
the Prosopolophini. We here assign monotypic Himero-
mima Warren, 1904 to Prosopolophini (Table 3). The
clade certainly requires a more comprehensive taxon
sampling in future studies.

B et al.: Phylogeny of New World Geometridae
478
Table 3. Taxonomic changes in Ennominae at the level of tribes, genera and species, in alphabetical order. *Old World taxa.
Synonymized tribes Valid tribes
Azelinini Forbes, 1948 syn.n. Odontoperini Tutt, 1896
New tribes Included taxa
Euangeronini new tribe Euangerona Butler, 1882, “Chloroclydon” rinodaria Felder & Rogenhofer, 1875,
Dectochilus Butler, 1882, Odontothera Butler, 1882, Mal leco Rindge, 1971,
“Opisogonia” dissata Felder & Rogenhofer, 1875
Oenoptilini, new tribe Neobapta Warren, 1905, Oenoptila Warren, 1895
Pyriniini new tribe Pyrinia Hübner, 1818, Acrotomia Herrich-Schäer, Acrotomodes Warren, 1895,
Falculopsis Dognin, 1913, Trotogonia Warren, 1905
Synonymized genera Valid genus
Nepitia Walker, 1866 syn.n. Pero Herrich-Schäer, 1855
Enypia Hulst, 1896 syn.n. Sabulodes Guenée, [1858]
Species proposed to be excluded from genera = incertae
sedis
Tribe assignment, country, originally described in genus
“Anisoperas” tessellata (Walker, [1863]) (analysed) Ennomini, Brazil, Hyperetis?
“Anisoperas” albimorsa Warren, 1905 Ennomini, Peru, Anisoperas
“Eusarca” bogotata (Snellen, 1874) (analysed) Ennomini, Colombia, Epione
“Hypomecis“ ectropodes (Prout, 1913) (analysed)* unassigned, South Africa, Boarmia
“Isochromodes” analiplaga Warren, 1907 (analysed) Ennomini, Peru, Paracomistis
“Isochromodes” sabularia Dognin, 1900 (analysed) Ennomini, Ecuador, Organopoda?
“Mychonia” galanata Dognin, 1895 (analysed) Ennomini, Ecuador, Mychonia
“Nematocampa” falsa Warren, 1906 (analysed) Ennomini, French Guyana, Nematocampa
“Sabulodes” thermidora (Thierry-Mieg, 1894) (analysed) Ennomini, Bolivia, Epione?
“Perusia” zoma (Dognin, 1896) (analysed) Ennomini, Ecuador, Acidalia
“Perusia” viridis Warren, 1907 (analysed) Ennomini, Peru, Perusia
Tribe transfer Taxon
Boarmiini to Macariini Dasyfidonia Packard, 1876
Boarmiini to unassigned Bryoptera Guenée, [1858]
Caberini to Ennomini Acrosemia Herrich-Schäer, [1855]
Caberini to Ennomini Microgonia Herrich-Schäer, [1855]
Caberini to unassigned Erastria Hübner, [1813]
Caberini to unassigned Sericosema Warren, 1895
Cassymini to unassigned Ballantiophora Butler, 1881
Cassymini to unassigned Berberodes Guenée, [1858]
Cassymini to unassigned Cirrhosoma Warren, 1905
Cassymini to unassigned Hemiphricta Warren, 1906
Cassymini to unassigned Hypometalla Warren, 1904
Cassymini to unassigned Phaludia Schaus, 1901
Ennomini to Boarmiini Mnesipenthe Warren, 1895
Ennomini to Gnophini Neorumia Bartlett-Calvert, 1893
Ennomini to Odontoperini Henicovalva Krüger, 2017
Ennomini to Palyadini Pityeja Walker, 1861
Ennomini to unassigned Idialcis Warren, 1906
Hypochrosini to Epionini Metanema Guenée, [1858]
Lithinini to Odontoperini Talca Rindge, 1971
Nacophorini to Ennomini Tarma Rindge, 1983
Nacophorini to Ennomini Brachyctenistis Warren, 1904
Nacophorini to Odontoperini Dentinalia Heimlich, 1960
Nacophorini to Odontoperini Macrolyrcea Butler, 1882
Nacophorini to Odontoperini Mallomus Blanchard, 1852
Nacophorini to Odontoperini Praeantarctia Heimlich, 1956
unassigned to Cassymini Orbamia Herbulot, 1966*
unassigned to Cassymini Pycnostega Warren, 1905*
unassigned to Boarmiini “Synnomos” apicistrigata Warren, 1895
unassigned to Ennomini “Acrotomia” mucia Druce, 1892
unassigned to Ennomini “Bassania” crocallinaria Oberthür, 1883
unassigned to Ennomini “Cyphoedma” transvolutata Walker, 1860

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ARTHROPOD SYSTEMATICS & PHYLOGENY — 77
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4.6.11. Diptychini. muriLLo-ramos et al. (2019) syno-
nymized Lithinini with Diptychini. A well supported
clade comprises former Lithinini including the Holarc-
tic name-bearing genus Petrophora Hübner, [1811] with
samples both from North America and Europe. Apart
from the Palaearctic genera, the former core Lithinini
also comprises Neotropical “Loxaspilates” (Fig. 7J), (not
congeneric with the Asian Gnophini genus Loxaspilates
Warren, 1893) and Neazata Warren, 1906. Neazata was
transferred from Caberini to Diptychini by muriLLo-ra-
mos et al. (2019). Three Chilean genera were previously
assigned to Lithinini: Tacparia Walker, 1860, Martindoe-
lloia Orla & Schajovski, 1963, and Tanagridia Butler,
1882 (pitkiN 2002). Another clade, exclusively compris-
ing austral South American taxa comprises Euclidiodes
Warren, 1895, Franciscoia Orla & Schajovski, 1963,
Psilaspilates Butler, 1893, and Rhinoligia Warren, 1895.
Further Diptychini include two large clades. Genera be-
longing to a clade of Australian taxa previously assigned
to Nacophorini were transferred to Diptychini by muriL-
Lo-ramos et al. (2019). Another clade comprises African
taxa previously assigned to Lithinini, to Nacophorini, to
Diptychini or that were unassigned.
4.6.12. Oenoptilini Brehm, Murillo-Ramos & Sihvo-
nen, new tribe.
—
Type genus. Oenoptila Warren, 1895
(Fig. 7K).
—
Material examined and phylo geny. Our
analysis comprises O. mixtata Guenée, [1858], the type
species of Oenoptila. Neobapta Warren, 1904 is also
included in the tribe. Branch support values from the
IQ-TREE analyses clearly conrm the monophyly of
Oe no ptilini (SH-like = 99.6, UFBoot2 = 100). How-
ever, the deeper phylogenic relationships of the tribe
are unresolved. — Molecular evidence. The tribe is
characterized by DNA sequence data from the fol-
lowing nine gene regions (exemplar Oenoptila mix-
tata, voucher Br-Geo-0006 from Brazil, illustrated in
Electronic Supplement File 5): ArgK (MK738208),
Ca-ATPase (MK738605), CAD (MK738934), COI
(MK739100), EF1a (MK740300), MDH (MK740945),
Nex9 (MK741374), RPS5 (MK741714), Wingless
(MK742175). — Morphology. Oenoptila and Neobapta
are structurally dissimilar: Oenoptila are characterised
by a pair of sclerotised, nger-like processes of the juxta,
which are tipped with a pouch. Neobapta has multiple
pairs of coremata and leaf-shaped process of the anallus.
These genera share the densely setose valva (illustrated
in pitkiN 2002). External features of analysed species are
illustrated in Fig. 7K and Electronic Supplement File 5.
Further detailed morphological analysis is required to
identify potential diagnostic features. — Remarks and
taxonomic changes. Neobapta and Oenoptila were pre-
viously assigned to Caberini by pitkiN (2002). However,
true Caberini (comprising Old and New World Cabera
Treitschke, 1825) form a separate clade phylogenetically
distant from Oenoptilini, see 4.6.19 below.
4.6.13. Baptini + Theriini clade. Baptini (Lomographa
Hübner, [1825]) are not closely related to Caberini. Bap-
tini and Theriini form a well supported clade and Lo-
mographa is represented with species from Europe (L.
bimaculata Fabricius, 1775) and Ecuador (unidentied
species).
4.6.14. Unnamed clade. This well supported lineage
com prises New World Erastria decrepitaria Hübner,
[1823], Madagascan and Afrotropical E. madecassaria
Boisduval, 1833 – and nested among these species, Ne-
arctic Metarranthis obrmaria Hübner, [1823]. We ex-
clude Erastria from Caberini (Table 3) and suggest more
study on Erastria and its relationship (and possible syn-
onymy) with Metarranthis, but current taxon sampling is
too limited for formal changes.
4.6.15. Plutodini + Palyadini clade. Old World Pluto-
des Guenée, [1858] (Plutodini) is sister to Neotropical
Palyadini that were revised by scoBLe (1994) and pitkiN
(2002). However, in the RAxML tree, Plutodes is sister
to Eumelea, and both are sister to Pyrinini (muriLLo-ra-
mos et al. 2019). Our analysis of Palyadini comprises Ar-
gyrotome Warren, 1894, Ophthalmoblysis Scoble, 1995
(Fig. 7L), Opisthoxia Hübner, [1825], Palyas Guenée,
[1858], Pityeja Walker, 1861 and Phrygionis Hübner,
[1825]. Without obvious reason, BeLJaev (2008) trans-
ferred Pityeja to Ennomini; we hereby reverse this trans-
fer (Table 3).
4.6.16. Apeirini + Epionini + Anagonini + Hypochro-
sini clade. Because the four tribes are represented only
by between one and three species, they are treated here
together. The clade comprises representatives of four
mostly Old World tribes: Apeirini (Apeira Gistl, 1848),
unassigned to Ennomini Eutomopepla Warren, 1894
unassigned to Ennomini “Neorumia” gracilis Bartlett-Calvert, 1893
unassigned to Ennomini Polla Herrich-Schäer, [1855]
unassigned to Nacophorini Eustenophasma Warren, 1897
unassigned to Nacophorini Leucochesias Mabille, 1899
unassigned to Odontoperini Nemeris Rindge, 1981
unassigned to Prosopolophini Himeromima Warren, 1904
New generic combinations Originally described in genus, transferred from genus, decision based on
Bryoptera vaga (Dognin, 1895) comb.n. Tephrosia, “Ectropis“, molecular data and external morphology
Table 3 continued.

B et al.: Phylogeny of New World Geometridae
480
Epionini (Epione Duponchel, 1829), Anagonini (Probole
Herrich-Schäffer, [1855] and Plagodis Hübner, [1823],
and Hypochrosini (Hypochrosis Guenée, [1858]). Three
Nearctic taxa are present in this lineage, and we propose
to change the tribe assignment of two taxa: Metanema
Guenée, [1858] is transferred from Hypochrosini to Epi-
onini. Sericosema Warren, 1895 (Fig. 7M) is excluded
from Caberini, but remains unassociated with tribe (Ta-
ble 3).
4.6.17. Drepanogynini. This tribe was described by
muriLLo-ramos et al. (2019). Sister to the aforemen-
tioned clade, this well supported tribe comprises four
African genera that previously were assigned to Naco-
phorini.
4.6.18. Pyriniini Brehm, Murillo-Ramos & Sihvonen,
new tribe. — Type genus. Pyrinia Hübner, 1818 (Fig.
7N). — Material examined and phylogeny. In addi-
tion to Pyrinia, this clade comprises Acrotomia Herrich-
Schäffer, [1855], Acrotomodes Warren, 1895, and Tro-
togonia Warren, 1905. Branch support values from the
IQ-TREE analyses clearly conrm the monophyly of Py-
riniini (SH-like = 100, UFBoot2 = 100). The tribe forms
a well suppported clade with Caberini, and tribes around
Cassymini (4.6.19. – 4.6.20.) but it is uncertain which is
sister to Pyriniini. — Molecular evidence. The tribe is
characterized by DNA sequence data from the following
ve gene regions (exemplar Pyrinia abditaria, Warren,
1905, voucher gb-ID-17449 from Ecuador, illustrated in
Electronic Supplement File 5): CAD (JF785178), COI
(JF784698), EF1a (JF785322), MDH (JF784839), RPS5
(JF784951). — Morphology. pitkiN (2002) already rec-
ognized the group and suggested the pair of setose pro-
cesses adjoining the juxta postero-laterally in the male
genitalia as a synapomorphy (Acrotomodes, Falculopsis
Dognin, 1913, and Pyrinia). They also share the divided
valva (not present in all species), and extended chae-
tosemata on dorsal side of head. Externally the species
are very diverse, see Fig. 7N and Electronic Supplement
File 5. — Remarks and taxonomic changes. Because
of pitkiN’s (2002) study, we also include Falculopsis in
Pyriniini (Table 3).
4.6.19. Caberini. Our data (and those of previous mo-
lecular analyses) do not support a close relationship of
Baptini and Caberini, e.g. as suggested by pitkiN (2002).
Many New World taxa assigned to Caberini by pitkiN
(2002) are not part of this clade (e.g. Paragonia Hübner,
[1823], Neazata, Oenoptila), but belong to Ennomini,
Diptychini and Oenoptilini, respectively (see above). On
the other hand, the Neotropical genera Aplogompha War-
ren, 1897 (Fig. 7O), Lobopola Warren, 1900, Sphacelodes
Guenée, [1858], and Thysanopyga Herrich-Schäffer,
1855, are Caberini indeed, as well as the Nearctic genera
Apodrepanulatrix Rindge, 1949, Chloraspilates Packard,
1876, Eudrepanulatrix Rindge, 1949, and Ixala Hulst,
1896. Cabera Treitschke, 1825 in its current limits is
polyphyletic, obviously requiring revision.
4.6.20. Cassymini + Abraxini + Eutoeini + Macariini
clade. This well supported clade (SH-like = 99.9, UF-
Boot2 = 100) comprises Cassymini, Abraxini, Eutoeini
and Macariini. The majority of the recovered phyloge-
netic relationships between the clades is, however, poorly
supported. The clade is strongly supported as sister group
to Boarmiini, and all four tribes have previously been
proposed to form a monophylum based on shared mor-
phological traits, viz. reduction of the pupal cremaster to
a pair of strong spines and the presence of a forewing
fovea (hoLLoWaY 1994; hoLLoWaY et al. 2001). While
most of the investigated taxa can safely be assigned to
one of the four tribes, eight genera can currently not be
assigned to tribe, namely African Dorsifulcrum Herbu-
lot, 1979, Palaearctic Odontognophos Wehrli, 1951, and
New World Ballantiophora Butler, 1881, Berberodes
Guenée, [1858], Cirrhosoma Warren, 1905, Hemiphricta
Warren, 1906, Hypometalla Warren, 1904 (Fig. 7P, Table
3), and Phaludia Schaus, 1901. Up to four clades might
represent tribes of their own, but further investigation
and broader taxon sampling is required.
Cassymini s.l. has modest support (SH-like = 65.7,
UFBoot2 = 66), whereas Cassymini s.str. is well support-
ed (SH-like = 100, UFBoot2 = 97). Cassymini s.str. com-
prises several Old World genera (including the species-
rich genus Zamarada Moore, [1887]) as well as Nearctic
Protitame McDunnough, 1939. FergusoN (2008) also
included Nematocampa, which is hereby transferred to
Ennomini (see above), and Taeniogramma Dognin, 1913
(not sampled in this study). A clade that comprises Neo-
tropical Leuciris Warren, 1894 (Fig. 7Q), as well as Af-
rican Orbamia Herbulot, 1966, and Pycnostega Warren,
1905 is sister to Cassymini s.str.. We transfer Pycnostega
and Orbamia from unassigned to Cassymini (Table 3).
Abraxini is well supported (SH-like = 99.7, UFBoot2 =
94). Abraxini has no representatives in the Neotropical
region (pitkiN 2002). FergusoN (2008) considered Ligdia
wagneri Ferguson & Adams, 2008 to be the sole repre-
sentative of Abraxini in North America while all other
species of Ligdia Guenée, [1858] occur in the Palaearc-
tic region (Scoble 1999). Eutoeini is also well supported
(SH-like = 98.2, UFBoot2 = 99) and appears to be absent
from the New World.
Macariini is perfectly supported (SH-like = 100, UF-
Boot2 = 100) and is divided between two diverse line-
ages based around Macaria Curtis, 1826 (Old World
and New World) (Fig. 7R) and Chiasmia Hübner, [1823]
(Old World), respectively. Both genera were resolved
as monophyletic, although our taxon sampling was lim-
ited. Macariini assignment is conrmed for Digrammia
Gumppenberg, 1887, Eumacaria Packard, 1873, Isturgia
Hübner, [1823], Heliomata Grote & Robinson, 1866,
and Narraga Walker, 1861. Dasydonia Packard, 1876
is transferred from Boarmiini to Macariini (Table 3).
mcguFFiN (1977) considered Dasydonia as being re-
lated to Hypagyrtis Hübner, 1818, based on similarities
in genitalic morphology, wing venation and the presence
of a forewing fovea; FergusoN (2008) did not mention
Dasydonia in relation to the North American Macari-

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ARTHROPOD SYSTEMATICS & PHYLOGENY — 77
(3) 2019
ini. Dasydonia differs considerably from most Nearctic
Boarmiini in genitalic structure, but is surprisingly simi-
lar to Isturgia, Eumacaria, Trigrammia Herrich-Schäffer,
[1855] and Mellilla Grote, 1873, sharing strongly and
smoothly emarginate male valve and the presence of a
prominent, scoop-like gnathos. Wing shape and pattern
elements of Dasydonia are also more similar to Macari-
ini, sharing the prominent three transverse forewing lines
and absence of discal spots.
4.6.21. Boarmiini. This extremely large clade is prob-
ably the best-sampled tribe of Ennominae and will be
treated in a separate paper by L. Murillo-Ramos et al. (in
prep.). JiaNg et al. (2017) concentrated their sampling on
the Oriental and Palaearctic region, and our study com-
plements this with material from the New World, Africa
and other regions. The clade does not only include “typi-
cal” well camouaged boarmiines but also conspicuous-
ly coloured genera such as Perigramma Guenée, [1858]
(Fig. 7S). We transfer Mnesipenthe Warren, 1895 from
Ennomini (BeLJaev 2008) to Boarmiini (Table 3). “Syn-
nomos” near apicistrigata Warren, 1895 is transferred
from unassigned to Boarmiini (Fig. 7T, Table 3).
4.7. Biogeographic patterns
We make some remarks with regard to New World taxa
here, but a global biogeographic analysis of the family
Geometridae is planned (H. Ghanavi et al. in prep.). Fig-
ures 1, 2, 4, 6 provide an overview of some major biogeo-
graphic patterns with regard to the New World, shown in
detail in Electronic Supplement File 2 (all New World
taxa are marked with colour). New World taxa are not
randomly distributed in the tree, but usually form more or
less large clusters, indicating colonizing events followed
by local diversication. While the taxon sampling is now
very good in the New World, much more sampling is re-
quired in other regions. Despite these principal limita-
tions, the results allow us some preliminary conclusions.
While Nearctic and Neotropic taxa are often intermixed,
the austral South American taxa tend to be more isolated.
This pattern is consistent with long recognized biogeo-
graphic patterns showing a separation of South America
into a tropical northern/central region on the one hand,
and a southern temperate region on the other (morroNe
2015). The southern region still has biogeographic con-
nections to other southern temperate regions due to their
peleogeographic connection via Antarctica (saNmartíN
& roNquist 2004). morroNe (2006) placed the entomo-
fauna of southern South America in the “Austral King-
dom”, together with Australia, South Africa and New
Zealand, and separate from the Neotropics. On the other
hand, hoLt et al. (2013), analysing distribution and phy-
logenetic relationships of vertebrates, placed all of South
America in the Neotropical faunal realm, though sepa-
rated into rather distinct northern and southern regions.
Currently, our data do not show any strong connections
that indicates an “Austral Kingdom” of geometrid moths,
but a clear isolation of the austral South American fauna
is evident (Andean Region). Further taxon sampling in
southern Africa, Australia and New Zealand is required
for a more complete view.
Figures 1 – 2 and 5 – 6 demonstrate that many lineages
are dominated by New World taxa or are even restricted
to this region. Many exclusively New World lineages are
described as new tribes in this paper, highlighting the
previous systematic bias, pointed out by e.g. ÕuNap et al.
(2016). On the other hand, many lineages are restricted
to other parts of the World. However, even our sampling
in the New World is still incomplete and some species-
rich genera were not sampled although they form diverse
groups in the Neotropical region also (for example, the
genera Scopula and Eupithecia).
In Sterrhinae, Neotropical species are present in
most major clades. Idaea Treitschke, 1825 is nested
deeply within a Neotropical clade, suggesting that the
ancestors of Idaea were Neotropical. Dithecodes War-
ren, 1900 is distributed in Asia and in the Neotropics – a
rather exceptional biogeographic pattern that calls for
more study. In Larentiinae-Trichopterygini, a clear nu-
cleus of southern South American taxa is visible (Fig. 1)
whereas only two samples are from the tropical parts of
the continent (Rhopalodes concinna Dognin, 1911 and
an unidentied genus + species). An Old World clade
comprises Trichopteryx Hübner, [1825] as well as the
Nearctic genus Cladara Hulst, 1896. In the Asthenini,
Eois represents the single (and extremely large) radia-
tion of the tribe in the Neotropics, and only two other
Asthenini genera are known to have a Holarctic dis-
tribution. Psaliodini comprise both species from both
the Neotropical region and austral South America. The
clade with the highest number of New World taxa is
formed of an unnamed lineage in the Larentiini complex
(4.2.20.2. – 4.2.20.10.). This clade is currently mostly
Neotropical with some nested austral South American
and Nearctic taxa. Remarkably, Stamnodes, the type ge-
nus of Stamnodini is represented with a Nearctic and
a Palaearctic species and nested deep within the clade.
A Neotropical origin can also be assumed for Sparga-
nia: It is very diverse in the Neotropical region, less di-
verse in the Nearctic region, and only one species occurs
also in the Palaearctic region.
In Geometrinae, New World representatives are re-
stricted to only four distinct lineages in our dataset. This
conrms previous ndings (pitkiN 1996; pohL et al.
2016). A large number of tribes are obviously restricted to
the Old World. By far, the largest radiation is Nemoriini
with Nearctic species nested within a large Neotropical
clade. Due to rather good taxon sampling of Geometrinae
(BaN et al. 2018), it can be concluded that these moths
are likely to have an origin in the Old World, and that the
New World was successfully colonized only a few times.
Ennominae is dominated by two large tribes, the En-
nomini and the Boarmiini. These tribes show very differ-
ent biogeographic patterns. Ennomini comprise mostly
Neotropical taxa with many nested Nearctic taxa. Cur-
rently, ve Old World Ennomini genera are all part of

B et al.: Phylogeny of New World Geometridae
482
a single clade that also comprises Nearctic taxa. Old
World lineages of Ennomini are likely to increase with
better taxon sampling, but currently available data sug-
gests a Neotropical origin of Ennomini. On the contrary,
Boarmiini show a completely different pattern: In this
tribe, the New World was probably colonized by many
independent lineages from the Old World. Since taxon
sampling in the New World is good, it is unlikely that
the current view will be challenged with denser taxon
sampling. In Boarmiini, colonization has probably pri-
marily occurred from the Palaearctic to the Nearctic re-
gion. New World taxa are mostly concentrated in only
two larger radiations, one around Physocleora Warren,
1897, Glena Hulst, 1896 and Iridopsis Warren, 1894,
and one around Prionomelia Warren, 1895, Melanolo-
phia Hulst, 1896 and Carphoides McDunnough, 1920.
The latter three genera, in addition to six other genera,
formed the former Melanolophiini (mcguFFiN 1987). In
addition, a large number of distinct lineages with one or
more representatives occurring in the New World are
widely scattered in the tree: Aethalura McDunnough,
1920, Epimecis Hübner, [1825], Hesperumia Packard,
1873, Hypagyrtis Hübner, 1818, Orthodonia Packard,
1876, Paleacrita Riley, 1876, Protoboarmia McDun-
nough, 1920 and “Synnomos” near apicistrigata Warren,
1895. Most of these genera are phylogenetically isolated
in the New World, but have close relatives in Eurasia.
For example, Orthodonia is closely related to the Eura-
sian genera Arichanna Moore, 1868, and Bupalus Leach,
[1815], in addition to the genera that are primarily Eura-
sian with one or few Nearctic representatives: Biston
Leach, [1815], Hypomecis Hübner, 1821, Lycia Hübner,
[1825], and Phigalia Duponchel, 1829. These Nearctic
boarmiine genera form a considerable portion of the ge-
ometrid fauna of the deciduous forest regions of eastern
North America, and their evolutionary links to Eurasia
hint at similarities to that of Tertiary relict plant distribu-
tions (miLNe & aBBott 2002). Genera, or even species
with clearly Holarctic distributions are concentrated in
the boreal forest region of the northern Nearctic, where
genera such as Dysstroma, Thera, Lampropteryx, Epir-
rita, Operophthera, Epirrhoe, Scopula, and Xanthorhoe
comprise a signicant portion of the total geometrid di-
versity. In contrast, the arid and semi-arid regions of the
southwestern Nearctic is dominated by lineages with Ne-
otropic origins, particularly the Boarmiini, Nacophorini,
Psaliodini, and Pterocyphini.
5. Conclusions
Our study comprises hundreds of New World Geo-
metridae taxa that have not been included in a phyloge-
netic study before. It signicantly pushes the New World
geometrid fauna from one of the phylogenetically least
studied to one of the best studied lepidopteran taxa, along
with a series of related papers (see Introduction). It was
our goal not “only” to provide a phylogenetic hypothesis,
but also to translate many of the results into taxonomy.
We are well aware that this was a balancing act: On the
one hand, we did not want to produce another phyloge-
netic study suggesting required changes but not perform-
ing them. On the other hand, it was beyond the scope of
our study to deeply examine the morphology of a broad
range of taxa. One might argue that the description of
nearly a dozen new tribes requires a detailed morpho-
logical study of each taxon. We agree that morphological
studies are indeed needed and data should be analysed in
a future integrative approach. However, we think that our
data offer a sufcient basis for many taxonomic changes,
and we only performed them in “clear” cases in terms
of branch support and available generic names – and in
agreement with ICZN regulations on the establishment of
new family group names. Should some of our hypotheses
be falsied in future studies, it is well possible that some
names will be synonymized. We regard this as a normal
process when more, both morphological and molecu-
lar data will become available, particularly for African
and Australian taxa. However, until we will know bet-
ter in the future, providing names for otherwise unnamed
clades in Geometridae signicantly eases communica-
tion in the community. Our paper, including illustrated
catalogues of nearly all sampled New World taxa, assigns
many taxa for the rst time to tribe. Moreover, it is a
basis for future taxonomic work, and we believe it will
ease the description and assignment of a large number
of taxa, including many new generic names and new ge-
neric combinations. We hope that our paper stimulates
further research on New World geometrids, particularly
in taxonomy and ecology.
6. Acknowledgements
We are indebted to colleagues who supported us with additional
material, with access to collections under their care and for pho-
tographs of (type) specimens that allowed us to validate identi-
cations and produce the electronic illustrated catalogues. Patricia
Gentili-Poole kindly allowed access to photographs of type speci-
mens deposited at the USNM (Washington, D.C.). We thank John
Chainey, Geoff Martin and Linda Pitkin at the NHM (London) for
providing access to the collections and photographs of Neotropi-
cal Ennominae moths.Visits by GB to the NHM in 2011 and 2017
were funded by grants from the SYNTHESYS programme (GB
TAF1048 and 6817). Charlie Covell (Gainesville, USA) and Wolf-
ram Mey (MfN, Berlin, Germany) provided photographs of several
specimens. Cathy Byrne (Hobart, Australia), Andreas Kopp (St.
Margarethen, Switzerland), Stefan Naumann (Berlin, Germany),
Dominik Rabl (Vienna, Austria), Hermann Staude (Magaliesburg,
South Africa), Toomas Tammaru (Tartu, Estonia) and Jaan Viida-
lepp (Tartu, Estonia), thankfully provided further specimens for the
molecular analyses. Support of DFG for eldwork in Costa Rica,
Ecuador and Peru for GB is acknowledged (Fi 547/10-1 and 10-
2, FOR 816, FOR 402, Br 2280/1-1, Br 2280/6-1). EÕ received
nancial support by institutional research funding (IUT 20-33) of
the Estonian Ministry of Education and Research. LM-R acknowl-
edges funding from Colciencias, 756-2016 and Universidad de
Sucre, Colombia. NW acknowledges funding from the Academy
of Finland (Grant No. 265511) and the Swedish Research Council
(Grant No. 2015-04441).

483
ARTHROPOD SYSTEMATICS & PHYLOGENY — 77
(3) 2019
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Electronic Supplement Files
at http://www.senckenberg.de/arthropod-systematics
File 1: brehm&al-geometridaenewworld-asp2019-electronicsup
plement-1.pdf — IQ tree including all taxonomic changes, tribes
colour-coded.
File 2: brehm&al-geometridaenewworld-asp2019-electronicsup
plement-2.pdf — IQ tree including all taxonomic changes, regions
colour-coded.
File 3: brehm&al-geometridaenewworld-asp2019-electronicsup
plement-3.pdf — Illustrated pdf catalogue of analysed New World
small subfamilies (Sterrhinae, Geometrinae, Archiearinae, Desmo-
bathrinae).
File 4: brehm&al-geometridaenewworld-asp2019-electronicsup
plement-4.pdf — Illustrated pdf catalogue of analysed New World
Larentiinae.
File 5: brehm&al-geometridaenewworld-asp2019-electronicsup
plement-5.pdf — Illustrated pdf catalogue of analysed New World
Ennominae.
Authors’ contributions
The idea for a series of contributions to Geometridae phylogeny
was by G.B., P.S. and N.W. G.B. has coordinated the taxon sam-
pling for New World taxa. G.B. and all other authors have writ-
ten the manuscript. Material was sampled by G.B., L.M.-R., A.H.,
B.C.S., E.Õ., A.M., D.B., F.B., R.M., and A.L. L.M.-R. has per-
formed most of the laboratory work and the data analysis. G.B. has
prepared gures, tables and the supplement les.
Zoobank Registrations
at http://zoobank.org
Present article: http://zoobank.org/urn:lsid:zoobank.org:pub:
CDFC8D5E-451F-4A40-B024-84720AAC1FA4
Brabirodini Brehm, Murillo-Ramos & Õunap, 2019:
http://zoobank.org/urn:lsid:zoobank.org:act:A758E6BC-184A-
47D1-8BD9-287026B57FED
Chrismopterygini Brehm, Murillo-Ramos & Õunap, 2019:
http://zoobank.org/urn:lsid:zoobank.org:act:CC03F324-6A37-
4BC2-B92C-B4D4D0C6A245
Cophoceratini Brehm, Murillo-Ramos & Õunap, 2019:
http://zoobank.org/urn:lsid:zoobank.org:act:2A577723-0060-
452E-AA37-66639529F6B6
Ennadini Brehm, Murillo-Ramos & Õunap, 2019:
http://zoobank.org/urn:lsid:zoobank.org:act:037D2FE7-B603-
4BAB-B99C-88B35FD6BB8A
Erebochlorini Brehm, Murillo-Ramos & Õunap, 2019:
http://zoobank.org/urn:lsid:zoobank.org:act:C0949D5F-E557-
4A89-B58F-CC345F6643E7
Psaliodini Brehm, Murillo-Ramos & Õunap, 2019:
http://zoobank.org/urn:lsid:zoobank.org:act:E1619B9F-
E96A-48C8-A68E-8FC2ED39D87E
Pterocyphini Brehm, Murillo-Ramos & Õunap, 2019:
http://zoobank.org/urn:lsid:zoobank.org:act:5051AC71-
34AC-4C17-8B1A-6B70DD4425A0
Rhinurini Brehm, Murillo-Ramos & Õunap, 2019:
http://zoobank.org/urn:lsid:zoobank.org:act:DB69B514-38C6-
4D8A-AB9B-BB79040F2D55
Euangeronini Brehm, Murillo-Ramos & Sihvonen, 2019:
http://zoobank.org/urn:lsid:zoobank.org:act:5372C36C-BACD-
4206-94AC-2CDAB45AEECD
Oenoptilini Brehm, Murillo-Ramos & Sihvonen, 2019:
http://zoobank.org/urn:lsid:zoobank.org:act:77A2C077-
0BDE-454F-8B68-1EB746FB7ABE
Pyriniini Brehm, Murillo-Ramos & Sihvonen, 2019:
http://zoobank.org/urn:lsid:zoobank.org:act:D115CF79-450F-
4C5A-9911-66990F606A4F