Insect Systematics and Diversity, 7(2), 3; 2023, 1–25
https://doi.org/10.1093/isd/ixad004
Research 
Molecular phylogenetics, phylogenomics, and phylogeography
Phylogenetic systematics, diversification, and 
biogeography of Cerurinae (Lepidoptera: Notodontidae) 
and a description of a new genus
Ryan A. St Laurent,1,2,*,  Paul Z. Goldstein,3,  James S. Miller,† Amanda Markee,2,   
Hermann S. Staude,4,  Akito Y. Kawahara,2,5,6,  Scott E. Miller,1,  Robert K. Robbins1,
1Department of Entomology, Smithsonian Institution, National Museum of Natural History, Washington, DC, USA,2McGuire Center for 
Lepidoptera and Biodiversity, Florida Museum of Natural History, University of Florida, Gainesville, FL, USA,3Systematic Entomology 
Laboratory, USDA, Smithsonian Institution, National Museum of Natural History, Washington, DC, USA,4Lepidopterists’ Society of 
Africa, P.O. Box 2586, Knysna, 6570, South Africa,5Department of Biology, University of Florida, Gainesville, FL, USA,6Entomology 
and Nematology Department, University of Florida, Gainesville, FL, USA *Corresponding author, mail: ryanstlaurent93@gmail.com
†Deceased.
Subject Editor: Marko Mutanen
Received on 15 September 2022; revised on 9 January 2023; accepted on 3 February 2023
We present the first dated molecular phylogeny of the Cerurinae moths (Notodontidae), based on sequence 
data for 666 loci generated by anchored hybrid enrichment. Monophyly of Cerurinae is corroborated, which 
includes the following genera: Pararethona Janse, Pseudorethona Janse, Oreocerura Kiriakoff, stat. rev., 
Cerurella Kiriakoff, Notocerura Kiriakoff, Hampsonita Kiriakoff, Afrocerura Kiriakoff, Cerurina Kiriakoff, 
Neoharpyia Daniel, Furcula Lamarck, Neocerura Matsumura, Americerura St Laurent and Goldstein, gen. nov., 
Cerura Schrank, and Kamalia Koçak & Kemal. The type species of the Neotropical genus Tecmessa Burmeister, 
T. annulipes (Berg), which had been incorrectly assigned to Cerurinae, is recovered in Heterocampinae; and 
Americerura gen. nov. is proposed to receive 17 unambiguously cerurine species transferred from Tecmessa. 
Divergence time estimates recover a crown age of Notodontidae roughly coincident with the K-Pg boundary, 
and a late-Oligocene crown age for Cerurinae. An African origin is inferred for Cerurinae, followed by col-
onization of the Palearctic, the Americas, Indomalaya, and Australasia during the Miocene. At least three 
independent colonizations of the Americas are inferred, one in the mid-Miocene associated with ances-
tral Americerura gen. nov. and two in the Pliocene and Pleistocene within Furcula. We hypothesize that the 
global spread of Cerurinae was enabled by that of its primary caterpillar foodplants in the Salicaceae. State-
dependent diversification analyses suggest that cerurines diversified most rapidly in temperate climates.
Key words: Africa, Salicaceae, temperate, tropical
Introduction (Fabaceae), and grasses (Poaceae) (Schintlmeister 2008, Miller 2009, 
Miller et al. 2018). Caterpillars of some species are regarded as agri-
The Prominent Moths (Lepidoptera: Notodontidae) are one of the 
cultural pests, and others as human food sources (Gschloessl et al. 
most diverse families of large moths and are found on every con-
2018, Mabossy-Mobouna et al. 2022).
tinent except Antarctica in habitats ranging from Arctic tundra to 
Notodontid caterpillars have complex ornaments, varied 
deserts and tropical rainforests (Miller 1991, Kitching and Rawlins 
colors, and otherwise highly irregular structures and forms. They 
1998, Schintlmeister 2008). The family includes over 4,000 rec-
also have complex behaviors and life history strategies, including 
ognized species and more names are being proposed each year 
spectacular displays of crypsis and aposematism, as well as soli-
(Schintlmeister 2013, Becker 2014). Notodontid caterpillars eat a 
tary and gregarious startle displays and silken tent building behav-
wide range of woody and herbaceous plants, with well-documented 
iors (Miller et al. 2018). Most have modified anal prolegs, which 
radiations associated with willows and poplars (Salicaceae), wal-
may be nonfunctional or exceptionally hypertrophied to form 
nuts (Juglandaceae), passion vines (Passifloraceae), Inga Willd. 
© The Author(s) 2023. Published by Oxford University Press on behalf of Entomological Society of America. 1
This is an Open Access article distributed under the terms of the Creative Commons Attribution License 
(https://creativecommons.org/licenses/by/4.0/), which permits unrestricted reuse, distribution, and 
reproduction in any medium, provided the original work is properly cited.
Version of Record, first published online April 05, 2023 with fixed content and layout in compliance with Art. 8.1.3.2 ICZN.
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2 Insect Systematics and Diversity, 2023, Vol. 7, No. 2
extendable whip-like projections called stemapods, which are per- these subfamilies are endemic to the Americas, and this precludes 
haps best exemplified by the Cerurinae (Fig. 1). Considering their their utility for understanding global biogeographic questions, for 
diversity, widespread distribution, and fascinating life histories, example. Since many currently recognized subfamilies occur glo-
Notodontidae have been the focus of much research since the dawn bally (Miller et al. 2018), their systematics are highly provisional. 
of lepidopterology. In the past four decades, a particular focus has Fortunately, however, the described global diversity of taxa has been 
been understanding their evolutionary history and phylogenetic cataloged (Schintlmeister 2013, Becker 2014).
placement within the Noctuoidea, the most diverse Lepidoptera Thus far, no study has attempted to infer a molecular phyl-
superfamily. ogeny of an entire subfamily or any other major higher-level clade 
Notodontidae have long been recognized as belonging to in the Notodontidae. Because of high notodontid species rich-
Noctuoidea on the basis of morphology (Packard 1895, Forbes ness and the lack of a family-wide monograph or well-supported 
1923), and recent phylogenetic efforts have robustly supported phylogeny, it is difficult to work towards a sound understanding 
this placement (Zahiri et al. 2011, Kawahara and Breinholt 2014, of the biogeography, evolution, or diversity of Notodontidae. 
Timmermans et al. 2014, Heikkilä et al. 2015, Regier et al. 2017, While no study has yet tackled the global biogeography of the 
Kawahara et al. 2019). Notodontid classification has largely fol- Notodontidae, Schintlmeister (1989, 2008) studied the biogeog-
lowed Miller’s (1991, 1992) phylogenetic analysis of adult, pupal, raphy of Notodontidae in the Old World, principally the Palearctic 
and larval morphological characters, and recent molecular efforts fauna, using the zoogeological categories paradigm of de Lattin 
have either had insufficient taxon sampling (Regier et al. 2017) (1967). So far, contemporary statistical frameworks using com-
or deficient molecular coverage (Kobayashi and Nonaka 2016) to parative phylogenetics have not been used to study notodontid 
resolve relationships within the group. No single notodontid clas- biogeography.
sification assigns all genera to established subfamilies. Ongoing To understand global patterns of Notodontidae biogeography, 
phylogenomic efforts, including the present study, use more expan- we focus on the Cerurinae, a charismatic group of notodontids, 
sive genome-wide molecular sampling to address key issues in the best known for their photogenic caterpillars, which have whip-like 
higher classification of this family. ‘tails’ (modified anal prolegs) and false thoracic ‘faces’ used to startle 
A particular impediment to Notodontidae research, apart from predators (Fig. 1). Caterpillars of at least one species are even cap-
the lack of a robust higher classification, is the lack of studies able of spraying formic acid from their thorax as a defense mech-
with worldwide coverage. Most monographs are regionally fo- anism (Poulton 1887). Adult cerurines are also intriguing, with 
cused, albeit comprehensive in their respective geographic areas their densely scaled bodies earning them the European common 
(Schintlmeister 2008, 2019, 2020, 2022; Miller et al. 2018, 2021). names ‘puss moths’ (Cerura spp.) and ‘kittens’ (Furcula spp.). The 
The only subfamily to be reviewed and fully revised on a global subfamily Cerurinae (96 species and 51 subspecies) is globally dis-
scale is the New World endemic Dioptinae (Miller 2008, 2009). The tributed and is perhaps the most taxonomically stable group among 
subfamily Nystaleinae has been reviewed (Weller 1992). Both of Notodontidae at the species level. The Eurasian genera, including the 
Fig. 1. Examples of Cerurinae larvae from nine of the 14 genera: A, Pseudorethona albicans (photo H. Staude); B, Notocerura spiritalis (photo S. Woodhall); 
C, Hampsonita esmeralda (photo H. Staude); D, Cerurina marshalli (photo L. Mulvaney); E, Furcula furcula (photo W. Bacher); F, Neocerura liturata (photo R. 
Tawade); G, Americerura scitiscripta (photo R. St Laurent); H, Cerura vinula (photo L. Jonaitis); I, Kamalia australis (photo C. & T. Deane).
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Insect Systematics and Diversity, 2023, Vol. 7, No. 2 3
most diverse Old World genera Furcula Lamarck, 1816 and Cerura lepidopterans that specialize on these plants, such as Cerurinae (de 
Schrank, 1802 were treated in Schintlmeister (2008). The most di- Mestier et al. 2022, Wang et al. 2022). As with Salicaceae, cerurines 
verse tropical Asian genus, Kamalia Koçak and Kemal, 2006, was display a distributional pattern of species richness that does not con-
treated in Schintlmeister (2002, 2008, 2020). The majority of African form to a common pattern predicted by the Latitudinal Diversity 
taxa were reviewed in Schintlmeister and Witt (2015). Finally, the Gradient (LDG) (Fischer 1960, Pianka 1966), in that the majority of 
North American fauna was fully revised by Miller et al. (2018, named species are found in the Northern Hemisphere in temperate 
2021). Only Neotropical taxa have yet to be revised, although they climates (Miller et al. 2018). Because of the close relationship of 
have been cataloged (Schintlmeister 2013, Becker 2014). Although Cerurinae with Salicaceae, the highlighted chemical defenses of these 
Cerurinae are relatively well understood, no single taxonomic effort plants, and their global distribution that contradicts the LDG, we 
has treated all species together or conclusively presents evidence that consider Cerurinae relevant in the continued effort to understand 
all genera currently placed in Cerurinae belong to this subfamily. the elements governing the distribution of biodiversity on the planet 
One possible reason for the lack of a uniform taxonomic treat- and how Lepidoptera became one of the most successful radiations 
ment of Cerurinae may be the fact that taxonomy of the group has of herbivores.
traditionally been based on adult morphology, with larval morph- The primary aims of the present study are (i) to provide the first 
ology—which contains the single most defining suite of charac- phylogenetic hypothesis for Cerurinae, (ii) establish its composition, 
ters for the subfamily—largely absent from taxonomic treatments. (iii) test its monophyly and that of its component genera, and (iv) 
Despite this, the original description of Ceruridae (=Cerurinae) by infer a dated biogeographic hypothesis to account for the group’s 
Butler (1881) was based on caterpillars, which Butler stated ‘…are distribution. We also explore the impacts of latitude on the diversifi-
broad in front, with a distinct angle or hump at the fourth segment, cation of Cerurinae and the potential role of Salicaceae in permitting 
fourteen legs and a forked pair of projecting tails, from which, when the global spread of cerurines. Finally, we consolidate the long taxo-
annoyed, bright-colored filaments are exserted.’ Since the time of nomic history of cerurines in a checklist.
Butler, the treatment of Cerurinae as a family group taxon has dif-
fered. Members of Cerurinae have either been included in their own Material and Methods
subfamily (Schintlmeister 2008, Becker 2014, Regier et al. 2017, 
Miller et al. 2018) or recognized as a closely related group of genera Taxon Sampling and Terminology
that belonged within a more broadly conceptualized Notodontinae: Sequences in this study are published/publicly available (Wellcome 
Dicranurini, a concept that includes many non-cerurine genera Sanger Institute 2022) or generated de novo from specimens in the 
(Miller 1991, Kobayashi and Nonaka 2016). Some authors have following collections: American Museum of Natural History, New 
treated this tribal name as a subfamily, Dicranurinae (Schintlmeister York, NY, USA (AMNH); African Natural History Research Trust, 
2008, 2020, Schintlmeister and Witt 2015). Leominster, UK (ANHRT); Carnegie Museum of Natural History, 
Recent phylogenetic efforts that either support the status of Pittsburgh, PA, USA (CMNH); collection of Hermann Staude, South 
Cerurinae as a valid subfamily (Regier et al. 2017) or as nested Africa (HS); McGuire Center for Lepidoptera and Biodiversity, 
within Notodontinae (Kobayashi and Nonaka 2016) differ greatly Florida Museum of Natural History, University of Florida, 
in the amount of genetic data used. Regier et al.’s (2017) analysis Gainesville, FL, USA (MGCL); National Museum of Natural 
was more inclusive, using 5–19 genes, whereas Kobayashi and History, Smithsonian Institution, Washington D.C., USA (USNM).
Nonaka (2016) only used one (short) ribosomal gene and one mito- Identifications were made using the literature (Schintlmeister 
chondrial gene. Most recently however, the concept of Cerurinae as a 2002, 2008, 2016, 2020; Schintlmeister and Witt 2015; Miller et 
distinct lineage within the Notodontidae has gained traction, mainly al. 2018) and expertly curated collections in MGCL and USNM. 
following the subfamily arrangement of Schintlmeister (2008, 2013) Morphological terminology follows Miller (1991) and Miller et al. 
and a subfamily diagnosis and treatment later given in Miller et al. (2018). Cerurinae taxonomy follows Becker (2014), Miller et al. 
(2018). Despite these works, monophyly of Cerurinae has not been (2018), Schintlmeister (2013), and Schintlmeister and Witt (2015), 
adequately tested in a phylogenetic context. as well as some taxonomic changes made in the present study. For 
Life histories of Cerurinae have received a fair amount of atten- outgroups we broadly follow the aforementioned references and 
tion in the literature, with caterpillar food plant information docu- some changes proposed by Kobayashi and Nonaka (2016). The 
mented for all but one genus. Nearly all Cerurinae feed on Salicaceae, identifications of caterpillars of Tecmessa annulipes (Berg, 1878) 
and studying these moths presents an opportunity to examine the and T. elegans Schaus, 1901, which we figure for the first time, with 
evolution and biogeography of a globally distributed lepidopteran color photos are based on inflated caterpillars and images curated in 
clade with conserved food plant associations. The relationship of the Natural History Museum, London, U.K. (NHMUK), which were 
Cerurinae with Salicaceae is noteworthy because these plants are examined by the first author.
known to be chemically defended by phenolic glycosides, which Sampled taxa include all 14 Cerurinae genera and 53 of the species/
have been shown to affect the fitness of generalist insect herbivores subspecies that we recognize based on a combination of taxonomic 
and act as feeding deterrents (Boeckler et al. 2011). Cerura vinula sources (including some unnamed taxa) (Table S1). To root the tree and 
(Linnaeus, 1758), perhaps the most well-known of the cerurines, de- determine the placement of Cerurinae, all other Notodontidae sub-
toxifies these compounds (Feistel et al. 2018). Ehrlich and Raven families sensu Miller et al. (2018), except Scranciinae, are sequenced, 
(1964) introduced coevolutionary theory based on Lepidoptera and using type genera whenever possible. Complete taxon samplings of 
their food plants, and fostered the idea that food plant specialization the ingroup and outgroups are listed in Table S2.
and shifts have frequently acted as mediators of the diversification of In addition to some taxonomic changes that we make in the 
Lepidoptera (Powell et al. 1998). Less attention, however, has been Results below, our treatments of Cerurina Kiriakoff, 1963 and the 
given to the impact of monophagy and oligophagy on diversifica- subspecies of Cerura erminea (Esper, 1783) differ from those of 
tion. Salicaceae are believed to have undergone a number of radi- Mulvaney (2021) and Schintlmeister (2008) respectively. We treat 
ations and, importantly, to have spread globally since the Paleocene Cerurina as monotypic with two subspecies C. marshalli marshalli 
and Eocene, thereby forming a pathway for the global dispersal of (Hampson, 1910) and C. marshalli argentata (Gaede, 1934) based 
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4 Insect Systematics and Diversity, 2023, Vol. 7, No. 2
on ongoing research (Mulvaney et al., unpubl. data). The East Phylogenetics
Palearctic/Indomalayan Cerura menciana Moore, 1877 is treated We base our results on a maximum likelihood (ML) phylogeny in-
as a valid species with subspecies C. e. formosana (Matsumura, ferred in IQ-TREE version 2.1.2, but see below for alternative tree 
1929) and C. e. birmanica (Bryk, 1949). In recent literature C. inference analyses that we also performed (Nguyen et al. 2015, Minh 
menciana is confusingly treated as either a subspecies of C. erminea et al. 2020). Using our concatenated dataset of 666 AHE loci, we per-
(Schintlmeister 2008) or as a valid species (Dai et al. 2015), but based formed 100 independent IQ-TREE analyses. In all analyses, 1,000 
on morphology and biogeography, we consider these distinct species. Ultrafast Bootstrap (UFBoot) and 1,000 Shimodaira-Hasegawa ap-
The Asian taxa formosana and birmanica are therefore treated as proximate likelihood ratio test (SH-aLRT) replicates were calculated 
subspecies of C. menciana based on morphological similarities if the for support values (Minh et al. 2013, Hoang et al. 2018a) and the 
name menciana is given specific status, as in the present paper. command ‘-bnni’ was used in all cases to reduce risk of branch sup-
port overestimation. Before the tree inference step, we performed 
IQ-TREE with the command ‘-m TESTNEWMERGEONLY’ which 
Molecular Data employs ModelFinder to identify the best partitioning scheme and 
This study includes phylogenomic data derived from (i) newly gen- model of nucleotide evolution of our 1,998 a priori (666 loci, each 
erated anchored hybrid enrichment (AHE; Lemmon et al. 2012) split into codon position) locus partitions (Kalyaanamoorthy et 
sequences using the Lep1 probe set (up to 855 highly conserved al. 2017). The a priori partitions were consolidated into 128 by 
genomic loci) described in Breinholt et al. (2018) and (ii) publicly ModelFinder. All partitioning schemes and selected best fit models 
available transcriptomes and genomes of Cerurinae, Notodontidae of nucleotide evolution are available on Dryad along with raw 
outgroups, and non-Notodontidae outgroups. For our dating ana- sequencing reads (St Laurent et al. 2023). The best tree of the 100 
lyses we include outgroups from Kawahara et al. (2019) since this independent runs was selected according to likelihood value. A ‘ro-
was also the source of secondary calibrations (see Divergence time bustly supported’ clade, as per IQ-TREE documentation, is defined 
estimation below). as having UFBoot ≥ 95 and SH-aLRT ≥ 80; a clade is considered 
Newly generated AHE data were compiled in the same manner ‘moderately supported’ with UFBoot ≥ 95 or SH-aLRT ≥ 80.
as numerous recent Lepidoptera studies (St Laurent et al. 2018, To test the robustness of our dataset with other tree inference 
2020; Homziak et al. 2019). While various methods of DNA ex- methods, we performed a multispecies coalescent (MSC) analysis 
traction have been used for previous AHE studies, we extracted with ASTRAL v. 5.7.8 (Zhang et al. 2017). Individual gene trees were 
DNA from all samples for the present article by soaking abdo- inferred with IQ-TREE (unpartitioned) as above and ASTRAL was 
mens (with one to three legs for some samples) in a 1:50 solution used to infer a consensus tree considering the multispecies coales-
of proteinase K and lysis buffer for ~18 h. An adapted OmniPrep cent. Branch support is expressed as posterior probability ASTRAL 
DNA extraction protocol (G-Biosciences, St. Louis, MO) was used support values (ASV), with values >0.95 considered to be evidence 
as described in Hamilton et al. (2019). Library preparation and of robust support. Finally, we employed MPBoot, a fast parsimony 
sequencing were conducted by Rapid Genomics (Gainesville, FL, method, on our concatenated supermatrix of 666 loci, with 1,000 
USA). The bioinformatic pipeline for AHE locus-trimming, as- MPBoot bootstrap replicates (Hoang et al. 2018b).
sembly, and ortholog selection follows Breinholt et al. (2018). In All post-assembly analyses, except ASTRAL and MPBoot, 
summary, raw reads were filtered with trimgalore! v.0.4.0 (bio- which were performed locally on a MacBook Pro (macOS 
informatics.babraham.ac.uk), and assembled using a custom it- Catalina), were carried out on the Smithsonian Institution High 
erative baited assembly python script, which employs USEARCH Performance Computing Cluster, Smithsonian Institution (https://
(Edgar 2010) and Bridger (Chang et al. 2015). Assembled AHE doi.org/10.25572/SIHPC) or the University of Florida’s HiPerGator 
loci were aligned with MAFFT version 7.407 (Katoh and Standley High Performance Computing Cluster. FigTree v. 1.4.4 was used for 
2013), and consensus sequences were built in FASconCAT-G ver- phylogenetic tree visualization (Rambaut and Drummond 2009).
sion 1.02 (Kück and Longo 2014), which was also used for con-
catenation. The only modification to this process was that MAFFT Divergence Time Estimation
alignment also included the ‘–adjustdirectionaccurately’ com- Due to difficulties in inferring divergence times with large genomic 
mand to permit alignment of reverse complimented sequences. datasets, we performed ML analyses that could accommodate large 
Transcriptomes were assembled as above to ensure seamless com- datasets of hundreds of loci at once. For more traditional, compu-
bination with the AHE dataset. The genomes, however, proved too tationally intensive Bayesian dating approaches, we subsampled our 
data-rich to assemble to the AHE loci using the IBA process, and dataset, reducing the number of loci used in downstream analyses 
therefore the ‘genome_getprobe_BLAST.py’ script from Breinholt (Jarvis et al. 2014, Misof et al. 2014, Prum et al. 2015, Smith et 
et al. (2018) was used to extract the AHE probe regions from the al. 2018, Kawahara et al. 2019, St Laurent et al. 2021). Regardless 
available genome assemblies, downloaded as complete chromo- of the analysis employed, we used nine secondary calibrations de-
some assemblies from The European Nucleotide Archive (ENA) rived from Kawahara et al. (2019), the most comprehensively fossil-
and made available by the Darwin Tree of Life project (Wellcome calibrated chronogram of Lepidoptera. Dates were assigned to the 
Sanger Institute 2022). AHE assembly was carried out on the following nodes: (i) The most recent common ancestor (MRCA) of 
University of Florida’s HiPerGator High Performance Computing the pyraloid Galleria (Fabricius) + Macroheterocera, (ii) crown of 
Cluster (http://www.hpc.ufl.edu/). Mimallonoidea, (iii) Mimallonoidea + remaining Macroheterocera, 
All alignments were examined manually with AliView version (iv) crown of Drepanoidea, (v) crown of Bombycoidea + 
1.18.1 (Larsson 2014) to ensure all sequences were in reading frame Lasiocampoidea, (vi) crown of Geometroidea, (vii) an internal 
and lacked artifacts introduced by misalignment. We used only AHE Noctuoidea node (MRCA of noctuid Helicoverpa Hardwick + nolid 
loci recovered from 60% or more taxa for phylogenetic inference Manoba Hampson), (viii) the Notodontidae root, and (ix) an in-
and all downstream analyses. In total 666 AHE loci were included in ternal Notodontidae node (MRCA of Pheosia Hübner + Notoplusia 
the final dataset. Table S2 summarizes the number of loci (of the 666 Schaus). The uniform prior dates, based on the 95% confidence 
used in analyses) for each taxon in our dataset. intervals in Kawahara et al. (2019) are shown in Table S3.
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Insect Systematics and Diversity, 2023, Vol. 7, No. 2 5
Our maximum likelihood dating approach used the program (i) one molecular clock, Yule; (ii) one molecular clock, Birth-death; 
TreePL (Smith and O’Meara 2012). TreePL provides a single tree (iii) three molecular clocks (one per codon position), Yule; (iv) three 
with single dates for each node as output, and therefore does not molecular clocks, Birth-death; (v) 15 molecular clocks (one per par-
permit the calculation of confidence intervals at the nodes. To infer a tition), Yule; and (vi) 15 molecular clocks, Birth-death. Path sam-
range of dates for all nodes, we used a custom set of Python scripts pling and steppingstone marginal likelihood estimations (MLE) were 
that enabled us to independently run TreePL many times, permit- calculated for all analyses and Bayes factors (Table S4) were used 
ting a summary of all analyses in the form of a maximum clade to identify the best clock/tree prior pair for use in downstream ana-
credibility (MCC) tree with date intervals at all nodes. This method lyses. For each of these combinations, three separate runs of 200 
was recently used by St Laurent et al. (2021) and in summary, uses million generations each were performed to confirm each run ran to 
RAxML (Stamatakis 2006) to generate 100 bootstrap replicates convergence, with all 600 million runs per analysis combined with 
from a supermatrix (which in this case is our 666-locus dataset, par- LogCombiner and TreeAnnotator (to generate the MCC tree). We 
titioned as above), and then uses these bootstrap replicates to infer confirmed all effective sample size (ESS) values were above 200 for 
branch lengths on a user-supplied tree that lacked branch lengths. all parameters across all analyses in Tracer v. 1.7.0 (Rambaut et al. 
The user-supplied tree was our best-of-100 (highest likelihood) 666- 2018). Only one statistic, ‘(tmrca(Mimallonoidea))’, was found to be 
locus (128 partitions) IQ-TREE output. The 100 resulting RAxML below 200 in some analyses, but this particular lineage presents dif-
trees, which by design are identical in topology, with only branch ficulty in dating analyses because of the extremely long branch sub-
lengths allowed to vary, were then input into 100 independent tending the Mimallonoidea (St Laurent et al. 2021). Although both 
TreePL analyses. The custom Python scripts allowed parameters the BEAST MCC and TreePL trees resulted in similar divergence 
to be optimized using the ‘prime’ command of TreePL and we per- time estimates (see Results), for simplicity, we use the best BEAST 
formed a randomly sampled cross-validation analysis to determine MCC tree for all downstream analyses and related discussion.
the best smoothing parameter for downstream TreePL analyses. The 
cross-validation step was run three times, allowing the smoothing 
parameter to vary to check for consistency across each run. All ana- Biogeography
lyses were run using the ‘thorough’ command of TreePL. The 100 re- We inferred the geographic origin for the Cerurinae using ancestral 
sultant trees dated by TreePL were summarized with TreeAnnotator range reconstruction and calibrated the direction and relative timing 
v. 1.10.4 in the BEAST v. 1.10.4 package (Drummond et al. 2012, of colonization. Considering the presence of two distantly related 
Suchard et al. 2018), in an MCC tree with divergence time intervals cerurine genera in the Americas, we hypothesize at least two inva-
at each node. These intervals are not confidence intervals calculated sions of the Americas from the Old World and sought to test this 
over a posterior distribution, but instead are the range of dates re- with our biogeographic reconstructions.
covered for each node across 100 independent TreePL analyses. Specimen distribution data were gathered by the first author from 
Our Bayesian approach used the methodology recently em- the Cornell University Insect Collection, Ithaca (CUIC), the MGCL, 
ployed by St Laurent et al. (2021), which uses SortaDate (Smith et the NHMUK, and the USNM. Additional data were provided 
al. 2018) to select AHE loci best suited for divergence time estima- from the ANHRT; Collection Pe. Jesus S. Moure, Departamento 
tion according to the following criteria: (i) clock-like evolution; (ii) de Zoologia, Universidade Federal do Paraná, Curitiba, Paraná, 
maximizing tree length; and (iii) least topological conflict with a pre- Brazil (DZUP); the collection of Tim McCabe, Albany, New York, 
defined species-tree. As in St Laurent et al. (2021), we prioritized USA (TM), and the Vitor Becker Collection, Camacã, Brazil (VOB). 
the parameters 3-1-2 in that order since we used a constrained tree Literature resources, particularly those that included distribution 
topology and the previous study found little impact when changing maps (Schintlmeister 2002, 2008, 2020; Schintlmeister and Witt 
the order of parameter priority if up to 75 loci were selected. The 2015; Miller et al. 2018), were consulted, as well as the current maps 
best species tree according to likelihood among 100 independent on iNaturalist.org with Cerurinae sightings continuously curated by 
IQ-TREE runs based on the full, 666-locus AHE dataset, partitioned the first author (iNaturalist 2022).
as described above, was used as the constraint tree for testing cri- Since Cerurinae are found on all continents except Antarctica, 
terion 3. Upon identifying the 75 loci for divergence time estima- we partitioned the globe into seven biogeographic regions, fol-
tion, we ran ModelFinder in IQ-TREE as above to identify the best lowing Kawahara et al. (2022) who investigated the biogeography 
consolidated partitioning scheme and models of nucleotide evolu- of butterflies, which have a global distribution similar to that of 
tion from the 75 a priori locus partitions, resulting in 15 new parti- Notodontidae. The regions are shown in Fig. 3, and include Nearctic 
tions. However, for this step we restricted models to those available (R), Neotropical (N), East Palearctic (E), West Palearctic (W), Africa 
in BEAST (Drummond et al. 2012, Suchard et al. 2018) since that (A), Indomalaya (I), and Australasia (U). The delimitation of the bio-
program would be used for dating. We then carried out 100 inde- geographic regions in Kawahara et al. (2022) was used here as well, 
pendent IQ-TREE runs, as above, on a concatenated supermatrix and we coded all cerurine species in our tree according to those re-
of the 75 loci partitioned in the 15 partitions to ensure that the top- gions. However, Neocerura liturata (Walker, 1855) was treated as 
ology of the subset of data did not conflict with the full 666-locus exclusively Indomalayan since only one record in Schintlmeister 
dataset. The resulting topology was effectively identical to our 666- (2008) was from the Indomalayan-East Palearctic boundary. This 
locus dataset (Fig. S2). SortaDate then tests each locus independently species is not generally considered Palearctic in distribution [based 
for criteria 1 and 2 using gene trees, which we inferred using an on recent observations from (iNaturalist 2022)]. The complicated 
unpartitioned alignment in IQ-TREE, and criterion 3 against the full taxon F. furcula (Clerck, 1759) is coded as occurring in both the 
data species tree, to check for topological conflict. The ‘pxrr’ com- East and West Palearctic. The single representative of F. furcula is 
mand in Phyx (Brown et al. 2017) was used to root all gene trees to from the United Kingdom where the West Palearctic subspecies F. f. 
the Pyraloidea outgroup. Twenty-four loci that were not represented furcula is present, but the complex taxonomic history and uncertain 
in the Pyraloidea outgroup were excluded. validity of some subspecies, which range across the Palearctic results 
For the Bayesian analysis with BEAST, using our 75-locus dataset, in our treating this taxon as more widespread for biogeographic re-
we performed six sets of molecular clock/ tree prior combinations: construction purposes.
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6 Insect Systematics and Diversity, 2023, Vol. 7, No. 2
For ancestral range reconstruction we used the ML methods into southern South Africa) we alternatively coded them as ‘both’ in-
implemented in BioGeoBEARS v. 1.1.2, specifically the Dispersal stead of ‘tropical’ to test whether this impacted model selection. We 
Extinction Cladogenesis (DEC) and Likelihood equivalent of the tested five models with GeoSSE: (i) dispersal parameters free to vary, 
Dispersal-Vicariance model (DIVALIKE) (Matzke 2013). These with no range-dependent diversification; (ii) a so-called ‘canonical 
analyses were run either unconstrained (no adjacency nor dispersal GeoSSE’ model with a range effect on diversification; (iii) a hidden-
multiplier matrices) or constrained (with both adjacency and dis- state GeoSSE (GeoHiSSE) model that includes one hidden trait and 
persal multiplier matrices) to compare results, though all analyses no range-dependent diversification; (iv) a GeoHiSSE model with one 
were stratified across four time periods (0.1, 5.33, 23.03, and 31.38 hidden trait, and range-dependent diversification; (v) and a Multi-
Ma). The matrices and time slices were adapted from Kawahara state Speciation and Extinction-like Model (MuSSE) with no hidden 
et al. (2022) with the oldest time slice reflecting the crown age of trait and no cladogenetic effects, but anagenetic changes allowed. 
Cerurinae. The maximum range size, which is defined as the number For model 5, anagenetic change refers to changes along the branches 
of areas any single species is allowed to occupy, was set to two be- rather than during cladogenesis. Habitat-temperature specialization, 
cause higher values resulted in an inability to infer ancestral ranges therefore, is not explicitly linked to speciation events (cladogenesis), 
at most nodes. but potentially to extirpation of widespread species (Goldberg et al. 
We also performed time-stratified Bayesian biogeographic ana- 2011). The best fit model was determined with Akaike information 
lyses in RevBayes (Höhna et al. 2016). We adapted the RevBayes criterion (AIC), corrected AIC (AICc), and AIC weight. GeoSSE re-
code employed by Thode et al. (2019) which incorporates adja- quires a sampling fraction of taxa in the tree from each of three 
cency matrices into the dispersal rate matrix (as opposed to a dis- states to correct for incomplete sampling from each region. We deter-
tance scaling factor), in some ways resembling our BioGeoBEARS mined these fractions to be 33% of temperate cerurines are present 
analyses with adjacency matrices. The Markov chain Monte Carlo in our study, 51% of tropical species are present in our study, and 
(MCMC) chain was set to 100,000 and maximum areas set to two 33% of cerurines that inhabit both types of environments are pre-
as for BioGeoBEARS. A burnin of 25% was set before generating sent in our study. Our GeoSSE code is available on Dryad. Table S1 
the MCC tree from RevBayes. Our RevBayes inputs are available on presents temperate/tropical assignments for all cerurine species and 
Dryad with the specific Rev code from Thode et al. (2019) available subspecies, including those not in our trees.
at that source.
Nomenclature
Food Plants and Latitude This paper and the nomenclatural act(s) it contains have been registered in 
Zoobank (www.zoobank.org), the official register of the International 
We conducted an ancestral state reconstruction (ASR) of food plants 
Commission on Zoological Nomenclature. LSID: urn:lsid:zoobank.
(here the term ‘food plants’ refer to plants fed upon by caterpillars 
org:pub:26CDD49C-BB73-415D-8DC4-C1623936B9CB.
in nature) using SIMMAP in the R v. 3 package phytools with the 
symmetrical (‘SYM’) model and 10,000 simulations (Bollback 2006, 
Revell 2011, R Core Team 2020). Food plant data were compiled Results
from various published and online sources and are presented to-
gether for the first time (Table S5). SIMMAP incorporates multiple Monophyly of Cerurinae
states (in case of polyphagous species) per tip by the user assigning All phylogenetic analyses support monophyly of Cerurinae (notwith-
equal prior probabilities to each of the known food plants. The ma- standing the erroneous assignment of some American cerurine taxa 
jority of cerurines feed on Salicaceae (Table S5), but a few are pol- to Tecmessa Burmeister, 1878; see below). Although family-wide sys-
yphagous on Salicaceae and other plants in nature, or feed only on tematics are not addressed pending ongoing sequencing efforts, we 
non-Salicaceae plants. Thus, states were assigned for the following provide a basic subfamilial backbone of Notodontidae to determine 
plant families for which there are unambiguous Cerurinae feeding the monophyly of Cerurinae and to improve the accuracy of diver-
records: Salicaceae, Burseraceae, Rosaceae, Betulaceae, Fagaceae, gence time calibration. The summary of the current backbone phyl-
Proteaceae, Malvaceae, Rutaceae, and Combretaceae; prior tip states ogeny of Notodontidae, including the placement of Cerurinae, based 
are found in Table S6. This ASR was intended simply to illustrate the on our ML results is shown in Fig. 4. Figs. S1–4 show Notodontidae 
conserved nature of this association across the tree and test the hy- relationships from a variety of phylogenetic inference methods.
pothesis that Cerurinae fed on Salicaceae ancestrally. Based on our phylogenetic results, Cerurinae are not nested 
Because Cerurinae reach their highest species richness in tem- within Notodontinae, but are sister to two genera usually asso-
perate climates (~60% of named cerurine taxa occur in temperate ciated with Dicranurinae: Shachia Matsumara and Liparopsis 
regions), we sought to examine whether environmental variables im- Hampson (Schintlmeister 2020). Harpyia Ochsenheimer, another 
pacted cerurine diversification rates. We approached this question genus often assigned to Dicranurinae, is then sister to ([Liparopsis, 
from the perspective of present day temperate vs. tropical distribu- Shachia, Cerurinae]). Because we include the type genera (using type 
tions using the Geographic State Speciation and Extinction Model species of those genera) of Cerurinae (Cerura) and Notodontinae 
(GeoSSE) with the R packages hisse and diversitree (Goldberg et (Notodonta Ochsenheimer), the separation of the two subfamilies 
al. 2011, FitzJohn 2012, Beaulieu and O’Meara 2016, Caetano is unequivocal, rejecting the arrangement of Notodontinae with 
et al. 2018). Taxa may be considered habitat temperature-limited, tribe Dicranurini containing cerurine genera, as in Kobayashi and 
occurring in either temperate or tropical regions, or widespread, Nonaka (2016). Based on results presented here, Dicranurinae as 
occurring in both. Taxa were assigned a ‘temperate’ state (i) if they it is currently understood is polyphyletic. Ongoing phylogenomic 
inhabit Nearctic, East or West Palearctic biogeographic regions, research including all subfamilies of Notodontidae and Dicranura 
and a ‘tropical’ state (ii) if they inhabit the Neotropics, Africa, Reichenbach (type genus of Dicranurinae) indicates that the true 
Indomalaya, or Australasia. Taxa inhabiting both a temperate and concept of Dicranurinae excludes all genera currently assigned to 
tropical biogeographic region were treated as ‘both’ (0). In Southern that subfamily except Dicranura due to the placement of that genus 
Hemisphere taxa that could be considered temperate (e.g., that range sister to nearly all other Notodontidae subfamilies (St Laurent et al. 
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Insect Systematics and Diversity, 2023, Vol. 7, No. 2 7
in prep.). Future work will establish new subfamily or tribal level named taxon expected to fall within this clade, but, based on its 
taxa for the clades including Harpyia, Shachia, and Liparopsis. genitalia, this South African endemic appears to be a close relative 
The same relationship of Cerurinae to the aforementioned of C. natalensis (Schintlmeister and Witt 2015). Oreocerura is the 
Dicranurinae s.l. genera is also recovered with the parsimony ana- only cerurine genus for which food plant information is unknown. 
lysis in MPBoot (Fig. S4). And while our ASTRAL analysis recovers Cerurella natalensis, on the other hand, has been recorded from 
a nearly identical topology for the ingroup (the placement of the Homalium rufescens (Salicaceae) (Kroon 1999).
clade containing Oreocerura Kiriakoff, 1963, stat. rev. + Cerurella 
Kiriakoff, 1962 shifts, however) with robust bootstrap support for Furcula
its monophyly (Fig. S3), the relationship of Cerurinae to Shachia and 
The data support the need to reassess use of the name Furcula fur-
Liparopsis is much less well-supported (ASV = 0.59 for [Cerurinae, 
cula in North America, as recently proposed by Miller et al. (2018). 
Shachia]).
These authors determined that the North American F. occidentalis 
An important issue highlighted by our results is that the genus 
(Lintner, 1878) represents two species. Based on Cytochrome c oxi-
Tecmessa, which was assigned to Cerurinae by Schintlmeister (2013) 
dase subunit I (COI) ‘barcoding’ (Hebert et al. 2003), the western 
and retained there by Becker (2014) and Miller et al. (2018), is 
North American populations are more closely related to Palearctic 
polyphyletic, with representatives appearing in both Cerurinae 
F. furcula than to North American F. occidentalis. In our analyses 
and Heterocampinae in all phylogenetic analyses (Figs. S1–4). 
we included the following three relevant taxa: nominotypical F. fur-
Importantly, the type species of Tecmessa, T. annulipes (Berg, 1878), 
cula furcula from the United Kingdom, what was called ‘F. furcula’ 
does not fall within Cerurinae, but within Heterocampinae, and dis-
in Miller et al. (2018) from Colorado, USA, and F. occidentalis 
plays at least one apomorphy (albeit in a reduced form) of that sub-
from New York, USA. The phylogenetic results recover the two 
family: a tuft of elongated scales that extend below the head, termed 
North American taxa as sister species with Old World F. furcula 
a ‘beard tuft’ by Miller et al. (2018). Therefore, a new genus is pro-
furcula sister to them, rendering ‘Furcula furcula’ sensu Miller 
posed below to remedy the issue of polyphyly of Tecmessa and, by 
et al. (2018) paraphyletic. Fortunately, two available names for 
extension, Cerurinae.
American populations exist: deorum Dyar (TL: USA: Colorado, 
In summary, the recognized genera of Cerurinae are (in order dis-
Manitou) and gigans McDunnough (TL: Canada: Alberta, Head 
played in Fig. 2): Pararethona Janse, 1920 (Figs. 1A, 2A, 5A, 6A, 7A), 
of Pine Creek, Calgary), both of which were named in 1922, 
Pseudorethona Janse, 1920 (Figs. 1A, 2B, 5B, 6B, 7B), Oreocerura 
but with gigans being named earlier (Dyar 1922a, McDunnough 
stat. rev. (Figs. 2D, 5C, 6D, 7D), Cerurella (Figs. 2C, 5D, 6C, 7C), 
1922). We therefore revalidate F. gigans stat. rev. to be applied 
Notocerura Kiriakoff, 1963 (Figs. 1B, 2E, 5E, 6E, 7E), Hampsonita 
to North American populations and synonymize F. deorum syn. 
Kiriakoff, 1963 (Figs. 1C, 2F, 5G, 6F, 7F), Cerurina (Figs. 1D, 2G, 
nov. with F. gigans. Although we sampled F. gigans from Colorado, 
5H, 6I, 7G), Afrocerura Kiriakoff, 1963 (Figs. 2H, 5J, 6J, 7H), 
nearer to the type locality of deorum than to that of F. gigans from 
Neoharpyia Daniel, 1965 (Figs. 2I, 5I, 6G, 7I), Furcula (Figs. 1E, 2J, 
Alberta, ongoing phylogenomic efforts include a near topotype of 
5F, 6H, 7J), Neocerura Matsumura, 1929 (Figs. 1F, 2K, 5K, 6K, 7K), 
F. gigans from Alberta, which is recovered as sister to, with weak 
Americerura gen. nov. (Figs. 1G, 2L, 5M, 6N, 7L, 11–14), Kamalia 
genetic divergence from, the F. gigans from Colorado included in 
(Figs. 1I, 2M, 5L, 6L, 7M), and Cerura (Figs. 1H, 2N, 5N, 6M, 7N). 
the present study, further supporting this synonymy (St Laurent in 
The monophyly of each of these 14 genera is robustly supported 
prep.). Miller et al. (2018) provided extensive morphological evi-
with bootstrap values across all analyses (Fig. 2, Figs. S1–4).
dence supporting the separation of North American populations 
For discussion purposes we refer to the clade containing African 
of F. gigans (as ‘F. furcula’) from F. occidentalis, but erroneously 
genera Notocerura, Hampsonita, Afrocerura, and Cerurina as ‘Clade 
stated that the caterpillar of ‘F. furcula’ from North America was 
A,’ which is sister to ‘Clade B’: Furcula, Neoharpyia and ‘Clade 
unknown. McDunnough (1922) provided a detailed description 
C’: Neocerura, Americerura gen. nov., Cerura, and Kamalia. These 
of the caterpillars of F. gigans, and it closely matches the known 
clades are denoted in Fig. 2.
caterpillars of F. furcula and F. occidentalis. Future efforts should 
In general, species-level taxonomy of Cerurinae has been well-
continue to examine fine-scale population genetics of F. furcula, its 
studied except for the faunas of the Tropical Americas and parts 
numerous Palearctic subspecies, and the American F. gigans and 
of Africa. Ongoing taxonomic revisionary efforts, including among 
F. occidentalis, since this is a group with complex taxonomy and 
African genera, are the focus of other studies (Mulvaney et al. 
biogeography.
unpubl. data; Schintlmeister pers. comm.). Males of each genus of 
Cerurinae are shown in Fig. 5 and their terminalia in Figs. 6 and 7.
Tecmessa and Americerura
Tecmessa contains the type species Thosea annulipes Berg, 1878 
Oreocerura and Cerurella from Argentina (later moved to Tecmessa by Burmeister (1878)) and 
Oreocerura stat. rev. (type species Cerura dissodectes Kiriakoff, Tecmessa elegans from southeastern Brazil (Berg 1878, Burmeister 
1958) is reinstated as a valid genus with the sole species being 1878, Schaus 1901). Tecmessa annulipes and T. elegans are both 
Oreocerura dissodectes, comb. rev. There is a rather significant (~10 similar in patterning to cerurines and have simple genitalia, superfi-
Ma) divergence between Oreocerura and the sister genus Cerurella cially similar to those of some cerurine genera (e.g., Cerura), see Figs. 
(type species Cerurella natalensis Kiriakoff, 1962) with which it had 8 and 9. Schintlmeister (2013), in his comprehensive checklist of 
been synonymized by Schintlmeister and Witt (2015). This diver- global Notodontidae and Oenosandridae, transferred all American 
gence is greater than that between most other pairs of sister genera representatives placed in Cerura since the time of Draudt (1932) to 
in the Cerurinae. We note that the terminalia of Oreocerura are re- Tecmessa based on these superficial similarities. This classification 
markably morphologically distinct from those of Cerurella (compare was followed by Becker (2014) and Miller et al. (2018). However, 
Figs. 6C, D and 7C, D) and from other cerurine genitalia, being one the transfer of species from Old World endemic Cerura to Tecmessa 
of the most complex genitalia structures in the subfamily. We did by Schintlmeister (2013) was erroneous, although these species do 
not sample Cerurella whittakeri (Kiriakoff, 1981) the only other not belong in Cerura either. Morphologically, true Tecmessa display 
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8 Insect Systematics and Diversity, 2023, Vol. 7, No. 2
Fig. 2. Best IQ-TREE result of 100 independent analyses. Absence of bootstrap values means UFBoot ≥ 95 and SH-aLRT ≥ 80. See Fig. S1 for outgroup relationships 
and full support values. Caterpillar in bottom left is Americerura scitiscripta showing a defense display (photo J. Miller). Adult cerurine moths shown to the 
right: A, Pararethona argentescens (photo D. Fischer); B, Pseudorethona albicans (photo M. FitzPatrick); C, Cerurella natalensis (photo Q. Grobler); D, Oreocerura 
dissodectes (photo R. St Laurent, NHMUK); E, Notocerura spiritalis (photo W. Roland); F, Hampsonita cf. esmeralda (photo N. Voaden); G, Cerurina marshalli 
(photo L. Mulvaney); H, Afrocerura sp. (photo M. FitzPatrick); I, Neoharpyia verbasci (photo F. Romão); J, Furcula furcula (photo G. Kunz); K, Neocerura liturata 
(photo S. Lamberts); L, Americerura scitiscripta (photo A. Sourakov); M, Kamalia sp. (photo G. Kunz); N, Cerura vinula (photo A. Hardacre).
the following characters not shared with Cerurinae: narrowly pec- et al. 2021). The caterpillar morphology (Fig. 10) and behavior, 
tinate (not plumose) antennae and somewhat developed ‘beard tufts’ which are known for both T. annulipes and T. elegans, are also 
below the haustellum, the latter of which is a derived condition of highly distinct from those of Cerurinae. The caterpillars of these 
Heterocampinae (Miller et al. 2018). We also note the strong simi- two species lack the main larval apomorphies of Cerurinae, namely 
larity of Tecmessa genitalia (see revised genus diagnosis below) to stemapods and enlarged thoracic segments (Berg 1878, Oleiro et al. 
some Heterocampinae genera, such as Coelodasys Packard (Miller 2011, St Laurent pers. obs., Wheeler pers. comm.); and both feed on 
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Insect Systematics and Diversity, 2023, Vol. 7, No. 2 9
Fig. 3. Biogeographic reconstruction of Cerurinae in RevBayes using a constrained, time-stratified DEC model. Nodal symbols represent the highest posterior 
probabilities (PP) for ancestral ranges, the node symbol size reflecting the PP value given in the inset legend. State codes are as follows: A = Africa, R = Nearctic, 
N = Neotropical, W = West Palearctic, E = East Palearctic, I = Indomalaya, and U = Australasia; with combined states given as a combination of any two codes, 
which are further denoted on key nodes for clarity.
Anacardiaceae, not Salicaceae as is typical of Cerurinae (Biezanko the genitalia of the two species are nearly identical (compare Fig. 
et al. 1974, Oleiro et al. 2011). We also note that these two species 9C and D). Although DNA extraction of these two species failed 
have gregarious caterpillars, whereas those of Cerurinae are nearly to provide enough DNA for AHE sequencing, the genitalia provide 
always solitary in all stages, except for the aposematic caterpillars of sufficient evidence for the placement of T. pedrana in Tecmessa and 
Cerurina (Mulvaney 2021, St Laurent pers. obs.). maintenance of Becker’s (2014) assignment of T. pica to Tecmessa. 
We transfer Tecmessa from Cerurinae to Heterocampinae based Furthermore, among the series of T. pica, we found specimens inter-
on our phylogenetic results (Figs. S1–4) and the characters con- mediate in coloration and patterning between T. pica and T. pedrana, 
sidered apomorphic for Heterocampinae and Cerurinae by Miller suggesting a close relationship between these two species. So far as 
et al. (2018). Two additional taxa, following our examination, are is known, true Tecmessa under this concept of the genus are en-
also included in Tecmessa on the basis of genitalia and external demic to South America, inhabiting Brazilian Atlantic Forest, grass-
morphology: the monotypic genus Corania Schaus, 1939, syn. nov., lands in Brazil and Uruguay, arid regions of Argentina, and Chilean 
assigned to Cerurinae in Becker (2014), is hereby synonymized with mountain ranges. Habitus and male genitalia of all known true 
Tecmessa: Tecmessa pedrana comb. nov. (TL: Argentina). Tecmessa Tecmessa are shown in Figs. 8 and 9, and caterpillars of two spe-
pica (TL: Chile), which was already transferred to Tecmessa by cies in Fig. 10. The two generic synonyms listed under Tecmessa in 
Becker (2014), is highly similar morphologically to T. pedrana and Becker (2014), Eucerura Schaus, 1901 and Eunaduna Dognin, 1901, 
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10 Insect Systematics and Diversity, 2023, Vol. 7, No. 2
Americerura St Laurent and Goldstein, gen. nov.
(Figs 11–14)
(urn:lsid:zoobank.org:act:CFE2F544-A9FD-4872-B026-
D7D7501E68FD)
Type species.
Cerura scitiscripta Walker, 1865: 408, by present designation.
Etymology.
The name is a combination of ‘America’ and ‘Cerura’ referencing that 
this is the only American contingent of a principally Old World clade 
containing Cerura and its closest relatives. The name is feminine.
Diagnosis.
As is typical of most cerurines, adult Americerura are primarily 
black and white, with the white of the wings deriving from lustrous, 
reflective scales in most species. Antennae are feathery, bipectinate to 
the tips in both sexes, with those of males bearing longer pectinations. 
The male genitalia (Fig. 12) are rather simple, and within Cerurinae, 
most similar to the related Old World genera Cerura and Kamalia, 
on the basis of smoothly upwardly curving rounded valvae, an ellips-
oidal vinculum, a short but well-sclerotized blunt uncus, and heavily 
sclerotized, short, curled fingerlike socii. Typically, Americerura 
valvae are narrower and more widely splayed than in Cerura, and 
the phallus and eighth sternite less robust than in Kamalia. The 
phallus is variable, but usually pointed. The eighth sternite (Fig. 13) 
nearly always bears a trifid plate forming a ‘W’, a similar trait seen 
in Kamalia. Female genitalia are generally simple (Fig. 14), without 
Fig. 4. Relationships among Notodontidae subfamilies inferred from the a heavily sclerotized ostium, prominent antevaginal plate, or a not-
present study based on the best of 100 IQ-TREE analyses. The full tree can 
able differentiation between the vaginal plates. Most species of 
be found in Fig. S1. Bootstrap values for all branches shown are 100/100 
(UFBoot/SH-aLRT) except for few noted nodes. Americerura are also smaller moths, in general, and thus genitalia 
are smaller overall than those of the large Cerura and Kamalia.
remain synonyms of Tecmessa because the type species of both were Americerura may be confused with some of the paler Furcula 
examined and determined to be Tecmessa species. Namely, the type species in North America where they are sympatric, but Furcula can 
of Eucerura, Drymonia pica Butler, 1882, is Tecmessa pica and the be easily recognized by the presence of dark, metallic scales on the 
type of Eunaduna, Eunaduna cerurata Dognin, 1901, is a synonym thorax that give it a characteristic sheen. Americerura could also 
of Tecmessa annulipes. continue to be confused with members of Tecmessa s.l. as they have 
Three other species transferred to Tecmessa by Schintlmeister been since Schintlmeister (2013), but the genitalia of Tecmessa s.l. 
(2013) belong neither in Cerurinae nor in Tecmessa, but to another are distinct, with narrower valvae, a narrower phallus, and a less 
as yet unnamed heterocampine genus. These species, T. olindata modified eighth sternite lacking a W-shaped structure. The antennae 
(Schaus, 1939), T. gonema (Schaus, 1905), and T. laqueata (Schaus, provide an immediate clue to their differentiation given that in 
1911) are each very similar to one another but together are divergent Tecmessa s.l. they are longer and less feathery. Americerura also lack 
in morphology (Figs. 8E, F, 9E) and phylogenetically (Figs. S1–4) the beard-tufts of the head and the cteniophores of some species of 
from true Tecmessa. Only T. gonema is included in our analyses, but Tecmessa s.l. as well.
external and genitalia morphology of these three species are nearly All known last instar Americerura caterpillars are green or 
identical. They differ greatly from Cerurinae and from Tecmessa s.s. yellow, with a variably shaped and colored dorsal saddle on the 
in that they bear cteniophores, a widespread trait in Heterocampinae thoracic and abdominal segments, and broad thoracic segments into 
which is absent in Cerurinae (Miller et al. 2018) and Tecmessa s.s. which the head can partially retract. Usually, the head is edged by 
Maintaining our focus on Cerurinae, we do not formally transfer red or pink on the prothoracic segment, with false eyespots flanking 
these taxa from Tecmessa at this time, but they will require a new the head on each anterior corner of the prothorax. All species of 
genus to maintain monophyly of Tecmessa and this is the topic of a Americerura have stemapods with eversible internal whips that are 
future work (St Laurent et al. in review). yellow, orange, or red (or some combination). Americerura caterpil-
Below we describe a new genus to accommodate the cerurine con- lars are only likely to be confused with Furcula, and only in Canada, 
tingent of species formally assigned to Tecmessa s.l. and transfer them the U.S.A., and Mexico where the genera co-occur. Furcula cater-
accordingly. Adult habitus and male genitalia are shown for all of the pillars generally lack colored prothoracic margins around the head 
transferred species in Figs. 11–13. Description of Americerura gen. nov. (and are less contrasting if present), have darker stemapods, and are 
ensures that all genera in Cerurinae, and the subfamily itself, are mono- generally much more mottled in overall appearance.
phyletic. Miller et al. (2018) provide a detailed description of ‘Tecmessa’ 
consisting of the North American species of Americerura, thus we pro- Description.
vide a new description here taking into consideration all Neotropical Adult. Male. Head: Width more than half that of thorax, frons col-
species and excluding the species now established as heterocampines. oration white, gray, black or bicolored black and white with black 
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Insect Systematics and Diversity, 2023, Vol. 7, No. 2 11
Fig. 5. Males of each genus of Cerurinae, type species shown unless otherwise indicated. A, Pararethona hierax (ANHRT); B, Pseudorethona albicans (ANHRT); 
C, Oreocerura dissodectes primary type (NHMUK); D, Cerurella natalensis (NHMUK); E, Notocerura spiritalis (ANHRT); F, Furcula furcula (USNM); G, Hampsonita 
esmeralda (NHMUK); H, Cerurina marshalli marshalli (NHMUK); I, Neoharpyia pulcherrima, not type species (MGCL); J, Afrocerura leonensis (ANHRT); K, 
Neocerura liturata (NHMUK); L, Kamalia cf. priapus, not type species (NHMUK); M, Americerura scitiscripta (MGCL); Cerura vinula (MGCL). Scale bar = 1 cm.
ventrally, eyes large, naked, occupying more than 2/3 area of head, dorsal surface, bipectinate to tip with rami increasing in length from 
eyes usually bordered posteriorly by dark brown or black scales; antennal base to roughly half antennal length where rami length 
labial palpus reduced, not extending beyond frons, two-segmented, abruptly become shorter, remaining short until terminus. Thorax: 
coloration usually as for frons; haustellum very short, not typically overall patterning gray or black and white, prothorax often with 
visible, antennal scape scaled in erect tuft of white or gray scales, black margin or fully clothed in black scales, mesothorax and meta-
antennae brown to black with white and/or black scales coating thorax with white ground color and black spots; thickly scaled. 
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12 Insect Systematics and Diversity, 2023, Vol. 7, No. 2
Fig. 6. Male genitalia of each genus of Cerurinae, type species shown unless otherwise indicated. The excised phallus is shown below the ventral aspect of 
the genitalia. A, Pararethona hierax, gen. prep. ANHRTUK00105411 (ANHRT); B, Pseudorethona albicans, gen. prep. ANHRTUK00006221 (ANHRT); C, Cerurella 
natalensis, gen. prep. STAUDE001 (HS); D, Oreocerura dissodectes, gen. prep. LEP33752 (MGCL); E, Notocerura spiritalis, gen. prep. LEP33758 (MGCL); F, 
Hampsonita esmeralda, gen. prep. LEP33751 (MGCL); G, Neoharpyia pulcherrima, not the type species, gen. prep. LEP33766 (MGCL); H, Furcula gigans, not the 
type species, gen. prep. LEP33768 (MGCL); I, Cerurina marshalli argentata, gen. prep. NHMUK010316654 (NHMUK); J, Afrocerura leonensis, gen. prep. 6-21-22:2 
(MGCL); K, Neocerura liturata, gen. prep. 5-14-21:1 (MGCL); L, Kamalia tattakana, gen. prep. LEP33760 (MGCL); M, Cerura vinula, gen. prep. LEP33762 (MGCL); 
N, Americerura scitiscripta, gen. prep. 5-12-22:6 (USNM). Scale bar = 1 mm.
In species with darker (not white) forewings, thorax tends to ap- spurs thin, short, clothed in fine white scales, in formula 0-2-2. 
proximate darker wing ground color. Legs mostly concolorous with Tarsal claws simple. Forewing length 11–24  mm (measured from 
thorax, basal segments black and white, tarsi typically black. Tibial edge of thorax to wing tip), wingspan 23–46 mm (measured wing 
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Insect Systematics and Diversity, 2023, Vol. 7, No. 2 13
Fig. 7. Male eighth sternite of each genus of Cerurinae, all sternites correspond to specimens figured in Fig. 6. A, Pararethona hierax, B, Pseudorethona albicans; 
C, Cerurella natalensis; D, Oreocerura dissodectes; E, Notocerura spiritalis; F, Hampsonita esmeralda; G, Cerurina marshalli argentata; H, Afrocerura leonensis; I, 
Neoharpyia pulcherrima; J, Furcula gigans; K, Neocerura liturata; L, Americerura scitiscripta; M, Kamalia tattakana; N, Cerura vinula. Scale bar = 1 mm.
tip to wing tip); subtriangular, outer margin weakly convex, apex Tegumen weakly defined. Uncus simple, bifid, or distally widened, 
not pronounced. Forewing dorsum ground color ranging from gray thickly sclerotized, blunt, covered in setae. Socii simple, fingerlike, 
(A. duonumennia [Dyar, 1912]), gray-brown (A. annulifera [Berg, upcurved and heavily sclerotized, thickness variable, externally 
1878]), to off-white or lustrous immaculate white (all other species). ranging from smooth to rugose. Gnathos absent. Valvae simple, 
Antemedial, medial, and submarginal ground color concolorous. rounded to somewhat acutely angled apically. Juxta typically a thin 
Overall forewing pattern variable but always with some combination sclerotized strip, partially fused to phallus. Phallus somewhat vari-
of wavy or zigzagging lines ranging from extremely reduced or nearly able, but typically broadest mesally, downcurved distally, with sharp 
absent (e.g., A. splendens [Jones, 1908], A. candida [Lintner, 1878]) apex, vesica thin, simple, bag-like without cornutus.
to nearly covering the forewings (some forms of A. scitiscripta). Female. Sexual dimorphism in general not greatly pronounced. 
Antemedially proximal to thorax usually a weakly defined trans- Head: As for male but antennae with shorter rami overall, rami 
verse black line present, antemedial band usually present, may range longest basally and becoming gradually shorter along antennal 
from complete and filled with light blue-gray scales to incomplete or length to apex. Thorax: As for male. Forewing length 18–26 mm, 
replaced by yellow splotches. Wing with variable number of zigzag- wingspan 23–50 mm; subtriangular, outer margin weakly convex, 
ging black lines medially. Postmedial line usually most well-defined apex not pronounced. Forewing dorsum as for male but wing 
line of wing, typically thickest subapically with a secondary outer broader. Hindwing dorsum gray, gray-brown to black, rarely white 
submarginal line paralleling postmedial line to varying degrees, both (e.g., A. rarata). Hindwing venter as for hindwing dorsum, usually 
postmedial and submarginal lines sharply crenulate, forming points without markings. Frenulum with numerous tightly packed bris-
at vein intersections. Discal spot usually absent but may be weakly tles. Abdomen: As for male but bulkier, terminal segments with 
defined as black mark, thick markings usually present along costa corethrogyne in some species. Eighth sternite simple, lacking the 
and anal angle. Most species bearing black markings between veins W- or five-pronged structure present in males. Genitalia (Fig. 14) 
along wing margin, with wing fringe further invaded by black scales (n = 4) simple; tergite eight a narrow, sclerotized bar. Apophyses 
aligned with intervenular marks along wing margin. Forewing venter anteriores reduced to thick prominences; apophyses posteriores thin, 
usually lacking markings but may be variously smudged by darker relatively short, not extending well beyond margin of eight, if at all. 
scales, particularly subapically. Hindwing dorsum generally de- Antevaginal plate a simple sclerotized bar; ostium a narrow longi-
void of markings, although dark discal marks, marks at anal angle, tudinal opening or more circular; postvaginal plate either a simple 
and along margin may be present; hindwing coloration ranging sclerotized region or forming more of a sclerotized pocket. Ductus 
from gray, blackish, to pure white. Hindwing venter as for hind- bursae indistinct; corpus bursae thin, bag-like, lacking signa, smaller 
wing dorsum, usually without marks except along costa. Frenulum or only slightly larger than remaining genitalia complex. Papillae 
a single bristle. Wing venation as for other Cerurinae, but forewing anales broad, covered in long, fine setae.
bearing a small accessory cell and radial veins arising from its apex. 
Abdomen: Clothed in white, gray, or black scales, separation be- Egg.
tween abdominal segments usually poorly defined but may be delim- Circular, compressed, coloration variable, including white, brown, 
ited by margins of hoary scales; terminal segment often with lighter pink, or green, usually with reticulate pattern. Laid singly on adaxial 
scales. Eighth sternite (Fig. 13) either with W-shaped sclerotization leaf surface.
(lacking the medial prong in A. bratteata (Draudt, 1932)) or with 
more complex five-pronged structure (A. rarata [Walker, 1865] and Caterpillar.
A. tehuacana [Draudt, 1932]). Central prong variable in width and (Figs. 1G and 2 inset) Head large relative to prothorax, reddish 
length depending on species. Genitalia (Fig. 12) (n = 25) simple in brown to black, head withdrawn deep into prothorax when not 
overall structure, with ovoid, weakly connected vinculum ventrally. feeding and especially during threat displays; body ground color 
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14 Insect Systematics and Diversity, 2023, Vol. 7, No. 2
Fig. 8. Tecmessa adult males. A, T. annulipes, type species (NHMUK); B, T. elegans (NHMUK); C, T. pica (USNM); D, T. pedrana comb. nov., primary type (photo 
V. Verdecia, CMNH); E, ‘Tecmessa’ gonema (USNM); F, ‘Tecmessa’ laqueata (USNM). Scale bar = 1 cm.
Fig. 9. Male genitalia of Tecmessa. The excised phallus is shown below the ventral aspect of the genitalia and the eighth sternite is shown to the right of each 
set of genitalia. A, T. annulipes type species, gen. prep. USNMENT01771001 (USNM); B, T. elegans, gen. prep. USNMENT01771030 (USNM); C, T. pica, gen. prep. 
USNMENT01771031 (USNM); D, T. cf. pedrana comb. nov., gen. prep. USNMEN01771034 (USNM); E, ‘Tecmessa’ gonema, gen. prep. 5-12-22:3 (USNM). Scale 
bar = 1 mm.
yellow to bright apple-green, becoming reddish brown when saddle varying in color from brown, white, purple, red, maroon, 
prepupal; prothorax usually with pink or red coloration along or green, saddle widest on T1 and A4 where it may become dis-
margin with head, forming a hood when head retracted, in all but continuous or lengthened towards the second pair of prolegs; some 
the final instar, each anterior corner of prothorax with spiked knob, species with prominent white subspiracular line (A. scitiscripta and 
these knobs absent and may be replaced by black eye spots in final A. candida), spiracles white, black, or orange; body usually speckled 
instar, one on either corner of triangular prothorax above head; laterally; stemapods black, brown, or with coloration of saddle 
metathorax usually with dorsal single or pair of knob-like protru- bleeding into them basally, with a yellow, orange, or red eversible 
sions; thoracic and abdominal segments with contrasting dorsal component that can be shunted outwards with fluid during threat 
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Insect Systematics and Diversity, 2023, Vol. 7, No. 2 15
Fig. 10. Tecmessa caterpillars. A, T. annulipes, Uruguay (photo J. S. López); B, T. elegans, Argentina (photo G. Wheeler).
displays. A10 bears thick green or black paranal spines (paraprocts) host genera (Biezanko 1962a, 1962b; Biezanko et al. 1974; Oleiro et 
which assist in catapulting fecal pellets far from the body. True legs al. 2011). Recent sightings of both species are on Anacardiaceae as 
vary from green to black or are striped green and black. Five to six well (iNaturalist, 2022). Americerura annulifera, however, has been 
larval instars. verified with images only from Salix (Bourquin 1939). Biezanko et 
al. (1974) also report several Americerura species from introduced 
Pupa. Eurasian Salix species. We recommend interested readers to search 
Obtect, dark red-brown, cremaster absent. Formed inside rigid co- for Americerura caterpillars on Salicaceae to have the best chances of 
coon spun lengthwise along partially excavated notch in wood, with finding them. Reports from Asteraceae, Euphorbiaceae, and Poaceae 
chewed wood and other material that may have coated the wood should be confirmed. The single record from Rosaceae is thought to 
(e.g., moss) integrated in or on matrix of silk. represent confusion with Furcula borealis (Guérin-Méneville, 1844) 
which is obligate on that host (Miller et al. 2018).
Natural history.
Available host records for Americerura species are in the Salicaceae, Systematic remarks.
as is typical of Cerurinae. Feeding records include the food plant The long branch length between the species pair A. scitiscripta and 
genera Azara Ruiz and Pav., Banara Aubl., Hasseltia Kunth, Laetia A. candida and the remaining Americerura species, is similar to the 
Loefl. ex L. (one record, this genus was recently synonymized with divergence time (over 15 Ma) between the two Old World genera 
Casearia Jacq.), Populus (L.), Salix (L.), and Xylosma G.Forst. Kamalia and Cerura. However, given the American taxa are mono-
(Bourquin 1939, Janzen and Hallwachs 2017, Miller et al. 2018, phyletic, and genitalia morphology is consistent across species, we 
Samarakoon and Alford 2019, iNaturalist 2022). A handful of re- do not feel the need to introduce yet another genus at this time. Upon 
ported cases of non-Salicaceae hosts, such as Asteraceae, Alchornea examination of all named species assigned to Americerura, we de-
(Sw.) (Euphorbiaceae), Prunus serotina Ehrh. (Rosaceae), Saccharum termine that A. argynnis and A. argentina (Dognin, 1911), syn. nov. 
officinarum L. (Poaceae), Lithraea Miers ex Hook. & Arn., and are synonymous. The male genitalia of the type of A. argentina are 
Schinus spp. (Anacardiaceae) are questionable, possibly due to within the range of specific variation among other A. argynnis geni-
nomenclatural confusion or wandering caterpillars (Biezanko talia preparations and are comparable to those of a paralectotype 
1962a, 1962b; Biezanko et al. 1974; Parasitoid-Caterpillar-Plant of A. argynnis. Americerura argynnis is a common, widespread spe-
Interactions in the Americas’ 2014; Janzen and Hallwachs 2017; cies in Brazil and Argentina (St Laurent unpubl. data). Other cases 
Miller et al. 2018; Prada Lara 2022). References to caterpillars of seemingly closely related Americerura species (e.g., A. dandon 
using Anacardiaceae (Biezanko 1962a, 1962b; Biezanko et al. 1974) [Druce, 1894] and A. grandis [Schaus, 1901]) were named from 
are most likely due to confusion of the names T. annulipes and A. widely separate type localities with major geographic barriers be-
annulifera, which were both described in the same paper a few pages tween populations; we do not consider such species pairs synonyms 
apart but in different original genera (Berg 1878). Schreiter (1943) as we do A. argentina and A. argynnis.
is apparently the source of the error, who incorrectly used the name We do not propose additional taxonomic changes within 
Cerura annulifera when describing and figuring Tecmessa annulipes Americerura but hasten to point out that the illustration of A. xicona 
larvae and an adult that he reared on Schinus. He confusingly also (Dyar, 1924) in Draudt (1932) is incorrect, and instead, a weakly 
referred to a true Americerura species near rarata as “Cerura argen- marked A. cf. rarata is figured (similar to our specimen LEP34078 
tina” in the same work. Tecmessa annulipes is a true Tecmessa and from MGCL). The type of A. xicona in the USNM is in actuality 
is reported to feed on Schinus and/or Lithraea (Anacardiaceae) (Berg very similar to A. scitiscripta as pointed out in Miller et al. (2018). 
1878; Biezanko 1962a, 1962b; Biezanko et al. 1974) and the re- The synonym of A. scitiscripta, platea (Schaus, 1890) named from 
lated species T. elegans has also been reported from the same two Veracruz, Mexico, may actually be more properly placed as a 
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16 Insect Systematics and Diversity, 2023, Vol. 7, No. 2
Fig. 11. Males of each species of Americerura gen. nov. A, A. annulifera comb. nov. (NHMUK); B, A. argynnis comb. nov., holotype of A. argentina syn. nov. 
(USNM); C, A. argynnis comb. nov. (photo E. Orlandin, DZUP); D, A. rivera comb. nov. (photo E. Orlandin, DZUP); E, A. trigonostigma comb. nov., holotype 
(USNM); F, A. purusa comb. nov., holotype (photo V. Verdecia, CMNH); G, A. splendens comb. nov. (photo E. Orlandin, DZUP); H, A. bratteata comb. nov. 
(NHMUK); I, A. dandon comb. nov. (USNM); J, A. grandis comb. nov. (NHMUK); K, A. presidio comb. nov., holotype (USNM); L, A. lancea comb. nov. (USNM); M, 
A. scitiscripta comb. nov. (USNM); N, A. xicona comb. nov., holotype (USNM); O, A. duonumennia comb. nov. (NHMUK); P, A. candida comb. nov. (MGCL); Q, A. 
rarata comb. nov. (MGCL); R, A. cf. tehuacana comb. nov. (NHMUK). Scale bar = 1 cm.
synonym of A. xicona, named from Distrito Federal, Mexico. But scitiscripta and may not warrant specific status. Further research on 
pending sampling that includes topotypical A. scitiscripta, A. xicona, these matters, as well as the species-level taxonomy of Neotropical 
and platea, we do not make those changes here. Our results show Americerura, is needed. The figure of Tecmessa gonema (as Cerura 
that the taxon A. candida, the validity of which was considered gonema) in Draudt (1932) is also incorrect, and instead shows a fe-
uncertain by Miller et al. (2018), is only weakly divergent from A. male of A. rivera (Schaus, 1901).
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Insect Systematics and Diversity, 2023, Vol. 7, No. 2 17
Fig. 12. Male genitalia of each species of Americerura gen. nov.. The excised phallus is shown below the ventral aspect of the genitalia. A, A. annulifera comb. 
nov., gen. prep. NHMUK010316656 (NHMUK); B, A. argynnis comb. nov., holotype of A. argentina syn. nov., gen. prep. USNMENT00991441 (USNM); C, A. 
argynnis comb. nov., paralectotype, gen. prep. 6-23-22:1 (USNM); D, A. rivera comb. nov., gen. prep. USNMENT01771032 (USNM); E, A. splendens comb. nov., 
gen. prep. USNMENT01771033 (USNM); F, A. trigonostigma comb. nov., holotype, gen. prep. USNMENT00991268 (USNM); G, A. purusa comb. nov., gen. prep. 
USNMENT01771008 (CMNH); H, A. bratteata comb. nov., gen. prep. USNMENT01771002 (AMNH); I, A. presidio comb. nov., gen. prep. LEP88199 (MGCL); J, A. 
dandon comb. nov., gen. prep. 5-12-22:2 (USNM); K, A. grandis comb. nov., gen. prep. NHMUK010316655 (NHMUK); L, A. lancea comb. nov., gen. prep. 5-12-22:4 
(USNM); M, A. duonumennia comb. nov., gen. prep. LEP88198 (MGCL); N, A. scitiscripta comb. nov., gen. prep. LEP33761 (MGCL); O, A. candida comb. nov., 
gen. prep. 5-12-22:1 (USNM); P, A. xicona comb. nov., holotype, USNM gen. slide 220 (USNM); Q, A. rarata comb. nov., gen. prep. 5-12-22:8 (USNM); R, A. cf. 
tehuacana comb. nov., gen. prep. NHMUK010316657 (NHMUK). Scale bar = 1 mm.
For a list of species currently included in Americerura, see the A. candida (Lintner, 1878), comb. nov., (Figs. 11P, 12O, 13N)
summary of taxonomic changes in the following section. A. dandon (Druce, 1894), comb. nov., (Figs. 11I, 12J, 13I)
A. duonumenia (Dyar, 1912), comb. nov., (Figs. 11O, 12M, 13P)
Summary of Taxonomic Acts A. grandis (Schaus, 1901), comb. nov., (Figs. 11J, 12K, 13J)
Taxonomic acts in Cerurinae: A. lancea (Schaus, 1905), comb. nov., (Figs. 11L, 12L, 13K)
Americerura St Laurent & Goldstein, gen. nov. (Type species: A. presidio (Dyar 1922b), comb. nov., (Figs. 11K, 12I, 13M)
Cerura scitiscripta Walker) A. purusa (Schaus, 1928), comb. nov., (Figs. 11F, 12G, 13G)
A. annulifera (Berg, 1878), comb. nov., (Figs. 11A, 12A, 13A) A. rarata rarata (Walker, 1865), comb. nov., (Figs. 11Q, 12Q, 13Q)
A. argynnis (Schaus, 1901), comb. nov., (Figs. 11B, C, 12B, C, A. rarata optima (Bryk, 1953), comb. nov.
13B, C) A. rivera (Schaus, 1901), comb. nov., (Figs. 11D, 12D, 13D)
A. argentina (Dognin, 1911), comb. nov., syn. nov. A. scitiscripta (Walker, 1865), comb. nov., (Figs. 1G, 2 inset, 2L, 
A. bratteata (Draudt, 1932), comb. nov., (Figs. 11H, 12H, 13H) 5M, 6N, 7L, 11M, 12N, 13L)
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18 Insect Systematics and Diversity, 2023, Vol. 7, No. 2
Fig. 13. Male eighth sternite of each species of Americerura, all sternites correspond to specimens figured in Fig. 12, gray areas around some sternites are 
composed of membrane from the abdomen. A, A. annulifera comb. nov.; B, A. argynnis comb. nov., holotype of A. argentina syn. nov.; C, A. argynnis comb. nov.; 
D, A. rivera comb. nov.; E, A. splendens comb. nov.; F, A. trigonostigma comb. nov.; G, A. purusa comb. nov; H, A. bratteata comb. nov.; I, A. dandon comb. nov.; 
J, A. grandis comb. nov.; K, A. lancea comb. nov.; L, A. scitiscripta comb. nov.; M, A. presidio comb. nov.; N, A. candida comb. nov.; O, A. xicona comb. nov.; P, A. 
duonumennia comb. nov.; Q, A. rarata comb. nov.; R, A. cf. tehuacana comb. nov.. Scale bar = 1 mm.
A. splendens (Jones, 1908), comb. nov., (Figs. 11G, 12E, 13E) Node-dating analyses recovered a range of 64.8–67.8 Ma 
A. tehuacana (Draudt, 1932), comb. nov., (Figs. 11R, 12R, 13R) (TreePL) and 61.2–70.9 Ma (BEAST) for the crown age of 
A. trigonostigma (Dyar, 1925), comb. nov., (Figs. 11E, 12F, 13F) Notodontidae. The split of cerurines from the sister lineage com-
A. xicona (Dyar, 1924) comb. nov., (Figs. 11N, 12P, 13O) prising Shachia + Liparopsis was estimated at 39.4–42.3 Ma 
(TreePL) and 34.2–41.5 Ma (BEAST). The crown age of Cerurinae 
Furcula gigans (McDunnough, 1922), stat. rev.
was inferred to be 30.5–33.7 (TreePL) and 28.3–34.7 Ma (BEAST). 
F. deorum (Dyar, 1922), syn. nov.
Here and in the discussion below, we refer to the major clades A, B, 
Oreocerura Kiriakoff, 1963, stat. rev. and C within Cerurinae, with the topology (A,[B,C]). The recovered 
Oreocerura dissodectes (Kiriakoff, 1958), comb. rev. crown ages for these are in the Miocene: Clade A: 14.8–16.2 Ma 
(TreePL), 13.5–18.3 Ma (BEAST); Clade B: 17.2–19.9 Ma (TreePL), 
Complete checklist of Tecmessa including taxonomic acts: 14.9–20.4 Ma (BEAST); Clade C: 18.4–19.8 Ma (TreePL), 16.1–
Tecmessa Burmeister, 1878 20.0 Ma (BEAST). All Cerurinae genera, except the recently diverged 
Corania Schaus, 1939, syn. nov. sister genera Afrocerura and Cerurina and the ancient monotypic 
Eucerura Schaus, 1901 Pseudorethona, also had their origins in the Miocene.
Eunaduna Dognin, 1901
T. annulipes (Berg, 1878), (Figs. 8A, 9A, 10A)
Biogeography
T. elegans (Schaus, 1901), (Figs. 8B, 9B, 10B)
T. pedrana (Schaus, 1939), comb. nov., (Figs. 8D, 9D) The Bayesian and ML biogeographic analyses resulted in similar 
T. pica (Butler, 1882), (Figs. 8C, 9C) scenarios for Cerurinae over the ~30 My since their origin, espe-
The following species are provisionally maintained in Tecmessa: cially when constraints on dispersal and adjacency were imposed. 
T. gonema (Schaus, 1905), (Figs. 8E, 9E) For purposes of discussion, we consider the constrained analyses 
T. laqueata (Schaus, 1911), (Fig. 8F) that include both adjacency and dispersal matrices the most real-
T. olindata (Schaus, 1939) istic approaches since they reflect paleogeography, though they do 
introduce user-imposed assumptions on historical biogeography. 
We focus our discussion on the Bayesian, constrained DEC analysis 
Divergence Time Estimation in RevBayes (Fig. 3) and the BioGeoBEARS analyses are found in 
Overall, maximum likelihood (Fig. S5) and Bayesian divergence time Figs. S7–10.
estimates were largely congruent for deep nodes. The Birth-death We infer that Cerurinae originated in the Afrotropics and re-
tree prior with three unlinked molecular clocks, one per codon pos- mained there exclusively until ~25 Ma when they most likely dis-
ition, was the preferred BEAST analysis (Fig. S6). Tree files showing persed to the East Palearctic. The most recent common ancestor of 
age ranges, means, and medians for all nodes from each dating ana- all Cerurinae except the Afrotropical genera (the ancestor of clades 
lysis are available on Dryad. B and C) therefore likely inhabited the East Palearctic. Clade B is 
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Insect Systematics and Diversity, 2023, Vol. 7, No. 2 19
no hidden trait and anagenetic effects. Importantly, turnover rates 
(temperate, τ00 = 2.40 vs. tropical, τ11 = 0.55) are higher in tem-
perate lineages, but because we found anagenetic rather than clado-
genetic effects there is not an explicit link between shifting from 
temperate to tropical or tropical to temperate ranges and speciation. 
An alternative coding strategy that treats Southern Hemisphere taxa 
that range into cooler regions as inhabiting both temperate and trop-
ical climates also recovered the MuSSE-like model with no hidden 
trait and anagenetic effects as the best model.
Discussion
Composition of Cerurinae
The genera assigned to Cerurinae by previous authors, except 
Tecmessa and Corania (here a synonym of Tecmessa) (Schintlmeister 
2008, Becker 2014, Schintlmeister and Witt 2015, Miller et al. 
2018), are monophyletic under a variety of phylogenetic inference 
methods. To alleviate the polyphyly of Tecmessa (Heterocampinae), 
we propose Americerura and transfer 17 species to it from Tecmessa. 
Based on our phylogenomic results, which included all genera of 
Cerurinae, the generic classification of this subfamily is likely to be 
stable.
At the notodontid subfamily level, the nearly comprehensive 
(lacking only Scranciinae and Dicranurinae s.s.) outgroup sam-
pling showed that Cerurinae is monophyletic and not nested within 
Notodontinae. Its relationship with Dicranurinae is yet unresolved. 
In the ML analyses, Cerurinae shares a common ancestor with the 
Dicranurinae s.l. genera Shachia, Liparopsis, and Harpyia. The 
ASTRAL analysis (Fig. S3) only weakly supports (ASV = 0.59) the 
sister relationship of Cerurinae with Shachia, and does not recover 
Liparopsis and Shachia as monophyletic, illustrating the instability 
Fig. 14. Ventral view of female genitalia of Americerura scitiscripta comb. of the ((Shachia, Liparopsis), Cerurinae) grouping recovered in 
nov., type species of Americerura gen. nov., gen. prep. 5-12-22:7. Scale bar ML analyses. While caterpillars of both Shachia and Liparopsis 
= 1 mm.
are similar to those of Cerurinae (e.g., by virtue of the presence of 
stemapods), they are physiologically distinct, feeding on Fagaceae 
found entirely in the Northern Hemisphere and is inferred to have and Juglandaceae rather than Salicaceae (Schintlmeister 1989, 2008, 
been restricted ancestrally to the East Palearctic, dispersing thence to 2020; Miller 1991; Funamoto and Sugiura 2017).
the Americas in the late Miocene with subsequent back-colonization The composition of the Dicranurinae will ultimately depend 
to the Old World and later a second colonization of New World on the placement of its type genus Dicranura in a more exhaust-
much more recently in the Pleistocene. Alternatively, considering that ively sampled phylogenetic analysis of the family. This will require 
we permitted up to two areas per species, we can also report these the reclassification and re-assignment of various ‘Dicranurinae’ to 
results in terms of a widespread East Palearctic and Nearctic dis- subfamilies better circumscribed by a combination of genomic and 
tributed ancestor of all Furcula, that diverged into a New World morphological data (St Laurent et al. unpubl. data). Pending add-
clade and a mostly Old World Clade, with the ancestor of F. fur- itional sampling, it is conceivable that the concept of Cerurinae 
cula, F. gigans, and F. occidentalis becoming widespread across the could eventually be broadened to include other taxa. Regardless, 
East Palearctic and Nearctic, and then subsequent isolation of the the monophyly of the genera here considered to compose Cerurinae 
North American lineage leading to F. gigans and F. occidentalis. The is well-established on the basis of our analyses and corroborated 
remaining Cerurinae (Clade C) had an equally complex biogeo- morphologically; establishing higher taxa is outside the scope of the 
graphical history, with an ancestral range in the East Palearctic, and present work.
subsequent dispersal to Indomalaya (Neocerura and Kamalia spp.), 
the Americas (Americerura gen. nov.), and Palearctic (Cerura). Divergence Time Estimation
The inferred origin of Notodontidae (61.2–70.9 Ma) straddles the 
Food Plant Use and Diversification Cretaceous-Paleogene (K-Pg) boundary, an important extinction 
Our SIMMAP ASR expectedly recovers Salicaceae as the ancestral event that immediately preceded the radiation of other globally dis-
host at all nodes in Cerurinae (Fig. S11), including as the ancestral tributed clades of Lepidoptera (Espeland et al. 2018, Kawahara et al. 
condition of the subfamily. Shifts to non-Salicaceae hosts occurred 2019). This result is an important advance in our understanding of 
at three tips: Hampsonita esmeralda (to Proteaceae), Furcula bor- the age of Notodontidae because the divergence time analysis is the 
ealis (to Rosaceae), and Furcula bicuspis (Borkhausen, 1790) (to first to include the majority of Notodontidae subfamilies. Previously, 
Betulaceae). the only available sources of divergence times for Notodontidae were 
Of the five GeoSSE models tested (Table S7) the best supported those reported in Wahlberg et al. (2013), which recovered a crown 
based on AIC, AICc, and AIC weight was a MuSSE-like model with age of Notodontidae at ~69 Ma, and Kawahara et al. (2019), which 
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20 Insect Systematics and Diversity, 2023, Vol. 7, No. 2
recovered the most recent common ancestor of Pheosia Hübner cycles and sporadic Beringian land bridges rather than long distance 
(Notodontinae) and Notoplusia Schaus (Nystaleinae), an internal dispersal (the latter being a more likely scenario in Americerura, for 
node of Notodontidae, to be ~50 Ma. The study by Wahlberg et example). Previous research suggested possible affinities between 
al. (2013) had more comprehensive taxon sampling but still lacked American ‘F. furcula’ and Old World F. furcula subspecies found in 
Platychasmatinae, an enigmatic subfamily thought to be sister to all Siberia and the Russian Far East (Miller et al. 2018), but further 
other Notodontidae (Kobayashi and Nonaka 2016). We included research is needed to contextualize the biogeography of each lin-
one representative of Platychasmatinae, corroborating its placement eage in this clade because we lack other F. furcula subspecies and 
as sister to all other Notodontidae. Therefore, the inferred crown topotypical F. gigans, which may be more closely related to Old 
age of 61.2–70.9 Ma, as found here, can be seen as a reasonable World F. furcula than Colorado populations (e.g., deorum) based 
hypothesis for Notodontidae considering these previous studies and on COI barcodes (Miller et al. 2018). Ongoing work with nuclear 
our inclusion of the earliest diverging lineage. However, comprehen- AHE phylogenomics, including both Alberta and Colorado F. gigans 
sive family-wide sampling was not the focus of the present study; populations, however, seems to contradict the mitochondrial data (St 
we caution that these dates are provisional and will be reassessed Laurent unpubl. data).
with more complete sampling. Since Cerurinae were absent from Concurrent with the invasion of the Americas in Clade C, the pre-
both Wahlberg et al. (2013) and Kawahara et al. (2019), we pre- dominantly Indomalayan and Australasian genus Kamalia and the 
sent the first estimated crown age for the subfamily in the Oligocene principally Palearctic genus Cerura evolved from an East Palearctic 
(30.5–33.7 Ma [TreePL] and 28.3–34.7 Ma [BEAST]). ancestor. While we only sampled one species of the Australasian 
contingent of Kamalia (K. amoa [Holloway, 1979] from New 
Biogeography of Cerurinae Caledonia), it is worth noting that it was found to be sister to the re-
An African origin for Cerurinae makes sense because all extant mainder of the genus. This makes determining the ancestral range of 
species of the sequentially sister lineages of Cerurinae until Clades Kamalia, and thereby the ancestral range of the most recent common 
(B+C) are Afrotropical. There is no evidence of secondary dispersal ancestor of Kamalia and Cerura, more complicated because the ma-
back into Africa south of the Saharan desert from any of the bio- jority of Kamalia species are Indomalayan and not Australasian. 
geographic analyses. The Mediterranean African Cerura delavoiei To further complicate matters, K. malaysiana jakli (Schintlmeister, 
(Gaschet, 1876) is the only taxon believed to have recolonized 2002) occurs on both sides of Wallace’s Line in Bali and Sumbawa, 
Africa, but only north of the Sahara in Morocco, Algeria, Tunisia, suggesting a second invasion of Australasia by the relatively wide-
and the Canary Islands (Schintlmeister 2008), a region we treat bio- spread K. malaysiana (Holloway, 1982). Kamalia and Cerura were 
geographically as part of the West Palearctic. considered a single genus until Schintlmeister (2002) split them on 
Outside of Africa, the biogeography of Cerurinae is more com- morphological grounds. Due to the estimated 15 My divergence and 
plicated, with several major dispersal events. However, the most the pronounced morphological differences between them, treating 
likely scenario is that Cerurinae invaded the Palearctic from Africa these as separate genera is warranted and conveys historical bio-
via the Balkans or Asia Minor, from whence they spread throughout geography and morphological information.
the East and West Palearctic and spawned all subsequent radiations 
of Cerurinae. Two major lineages evolved in the Palearctic, the first 
(Clade B) leading to the sister genera Neoharpyia and Furcula, which Cerurinae Food Plants and Global Temperature
spread from the Palearctic to the Americas; and the more speciose Cerurinae are nearly entirely oligophagous on Salicaceae, a plant 
Clade C which independently colonized the Americas (Americerura), family which originated much earlier than Notodontidae in the 
Indomalaya and Australasia (Neocerura and Kamalia), and radiated Mesozoic (Li et al. 2019, de Mestier et al. 2022). Most molecular 
in situ within the Palearctic (Cerura). phylogenies that include Salicaceae do not sample densely across the 
The dispersal events that led to the colonization of the Americas family, with available research examining specific clades or genera 
by Americerura and Furcula occurred during the Miocene. (de Mestier et al. 2022, Wang et al. 2022) or the parent order, 
Americerura are found from Canada to Uruguay and have radiated Malpighiales, with limited Salicaceae sampling (Xi et al. 2012, Cai 
in the Neotropical realm, whereas Furcula are found exclusively in et al. 2020). Li et al. (2019) however, did examine the timing of di-
the Northern Hemisphere. The ancestor of Americerura arrived in vergence based on plastid genomes and sampled 18 genera across 
the Americas via North America some 15 Ma, requiring either long Salicaceae. There is also some disagreement as to what constitutes 
distance dispersal from the Old World or shorter dispersal routes in Salicaceae sensu stricto, with the relevant studies differing on the in-
the Bering region. The ancestor of Furcula, on the other hand, spread clusion of Casearia, which is consistently found to be sister to all or 
from an exclusively East Palearctic range to a more widespread most other Salicaceae (Xi et al. 2012, Li et al. 2019, Cai et al. 2020, 
East Palearctic and Nearctic range during the Miocene, potentially de Mestier et al. 2022). However, based on what is known at this 
overlapping the period when Americerura is inferred to have arrived point, Salicaceae had a Cretaceous origin, with stem ages reported 
in the Nearctic. The ancestor of Furcula eventually led to two major as 102.08–86.05 Ma in de Mestier et al. (2022), ~93 Ma in Li et al. 
lineages in the late Miocene to early Pliocene, one each in the Old (2019), and 78.7–59.8 Ma in Xi et al. (2012). Regardless of whether 
World and New World. Within the Old World Furcula clade, at the or not Salicaceae includes Casearia and some other debated lineages, 
beginning of the Pleistocene, the ancestor of F. furcula, F. gigans, and the crown age of Salicaceae excluding these taxa still antedates the 
F. occidentalis likely spread again to a semi-Holarctic distribution crown of Cerurinae by some 20–30 My (65.1–51.3 Ma in Xi et al. 
(Nearctic and East Palearctic) and in less than 2 My, split into F. (2012)). Based on these ages, the Salicaceae biogeography recon-
furcula in the East and West Palearctic and the ancestor of F. gigans struction in de Mestier et al. (2022), and our own biogeographic 
and F. occidentalis in the Nearctic. These two extant species occur in reconstruction of Cerurinae, Salicaceae had already colonized much 
boreal/subarctic habitats ranging into temperate deciduous forests, of the globe by the time Cerurinae arose.
and the sister to the clade of these two species, Eurasian F. furcula, Within Salicaceae, there is not a clear affinity for a single clade 
is found throughout the northern Palearctic. Thus, the biogeography or genus of plants. For example, the ‘derived’ Asian cerurine genus 
of these high-latitude Furcula species was likely impacted by glacial Neocerura is known to feed on Casearia in the Old World Tropics 
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Insect Systematics and Diversity, 2023, Vol. 7, No. 2 21
(de Mestier et al. 2022), but in the Americas where Casearia are new plants has been a central tenet in attempts to understand 
most species-rich, only two feeding records exist. Janzen and Lepidoptera diversification dynamics (Janz et al. 2006, Ebel et al. 
Hallwachs (2017) report two instances of Americerura rarata on 2015, Fagua et al. 2017, Sahoo et al. 2017, Strutzenberger et al. 
Laetia thamnia L., but the genus Laetia was recently synonymized 2017, Braga et al. 2018, Kergoat et al. 2018, Bruzzese et al. 2019, 
with Casearia and phylogenetic evidence supports the synonymy Toussaint et al. 2019, Allio et al. 2021, St Laurent et al. 2021). We 
(Samarakoon and Alford 2019, de Mestier et al. 2022). Extensive also do not consider food plant shifts within Salicaceae to be a major 
searching for caterpillars on Casearia spp. by the first author has factor driving cerurine diversity since the larvae of this group tend 
not revealed the presence of any cerurines despite their abundance not to specialize, and essentially feed on any available Salicaceae 
on nearby Salicaceae genera Xylosma and Banara (St Laurent pers. (see our earlier discussion about tropical Cerurinae feeding pref-
obs.). Casearia is one of the most ancient lineages of Salicaceae, and erences). Cerurinae are physiologically adaptable, given the range 
originated in the Neotropics ~10 million years before the origin of latitudes and habitats they occupy and their species richness in 
of Cerurinae in Africa (de Mestier et al. 2022). In the Northern the temperate regions. To further understand the mechanisms by 
Hemisphere, relatively unrelated but sympatric Cerurinae clades which Cerurinae so successfully spread across the planet, future re-
(e.g., Cerura and Furcula in the Palearctic; Americerura and Furcula search might investigate the chemical ecology of Salicaceae feeding 
in the Nearctic) feed on Salix and Populus, the two primary, and and how cerurine detoxification of foods may have allowed them to 
often only, Salicaceae genera found in temperate regions (Wang et utilize an abundant, but chemically defended group of food plants. 
al. 2022). There are also cases of non-Salicaceae feeding by Furcula Such a pathway may have been paved by an ancient adaptation to 
borealis (on Rosaceae) and F. bicuspis (Betulaceae) in these areas. feeding on Salicaceae in Africa, deep in the Oligocene, that led a 
In the tropics, where Salix and Populus are not major components curious group of insects with familiar ‘faces’ and whip-like ‘tails,’ all 
of forests, cerurines feed on a wide range of other Salicaceae genera over the world.
from the various accepted Salicaceae subfamilies: Azara, Banara, 
Casearia (=Laetia), Hasseltia, and Xylosma in the New World 
tropics and Casearia, Dovyalis E.Mey. ex Arn., Homalium Jacq., Supplementary Material
Flacourtia Comm. ex L’Hér., Scolopia Schreb., Trimeria Harv., and Supplementary data are available at Insect Systematics and Diversity online.
Idesia Maxim. in the Old World tropics (Kroon 1999, Schintlmeister 
2008, 2020, Chandra et al. 2018). The fidelity to Salicaceae is nearly 
universal among Cerurinae, but utilization of specific food plants Acknowledgments
appears to be opportunistic with respect to local availability of 
Salicaceae species rather than a distinct pattern of coevolution. This We offer our sincere thanks to Vitor Becker, Eduardo Carneiro, Jason 
fact that Salicaceae appear to have antedated Cerurinae supports Dombroskie, David Grimaldi, Guilherme Fisher, Loren Jones, Kevin 
Keegan, Joe Martinez, and Elton Orlandin for assistance gathering distribu-
the scenario that Cerurinae dispersed around the globe opportunis-
tion data used in this project. Ana P. S. Carvalho assisted with BEAST and 
tically due to the widespread availability of Salicaceae food sources. BioGeoBEARS. We also appreciate helpful discussions with Chris Schmidt 
Yet another indication of opportunistic feeding on Salicaceae by and Alberto Zilli. We are particularly grateful to Alberto for hosting Ryan St 
Cerurinae can be found in South America, where there are reports Laurent in London during a research visit to the Natural History Museum, 
of several Americerura feeding on introduced, ornamental Salix spe- London, UK. Alexander Schintlmeister provided many type photos that aided 
cies (Biezanko et al. 1974). The single native Salix in the region (S. in identification of Cerurinae. Numerous individuals supplied photos of 
humboldtiana Willd.), is a host of A. annulifera (St Laurent pers. living Cerurinae adults and larvae used in this study, see the figure legends 
obs.), similar to how A. scitiscripta feed on various Salix and Populus for specific names. Ryan St Laurent thanks the Peter Buck Postdoctoral 
in North America, where other genera of Salicaceae are absent. Fellowship program for funding this research. This research received support 
As inferred from our GeoSSE analyses, Cerurinae turnover rates from the SYNTHESYS+ Project http://www.synthesys.info/ which is financed 
by European Community Research Infrastructure Action under the H2020 
are highest in temperate regions with an anagenetic impact of tem-
Integrating Activities Programme, Project number 823827. We also thank 
perature on turnover (Table S7). This suggests that while Cerurinae three reviewers who improved an earlier version of this manuscript. Mention 
had a tropical African origin, and clearly are present in most tropical of trade names or commercial products in this publication is solely for the 
regions, their ability to diversify in temperate regions may explain purpose of providing specific information and does not imply recommenda-
their relatively high extant diversity in the Northern Hemisphere. tion or endorsement by the USDA; USDA is an equal opportunity provider 
However, the observed effect of range (defined as either widespread and employer.
or restricted to temperate/ tropical climates) on net diversification is 
anagenetic, and so could be explained by local extirpation of wide-
spread species over time rather than cladogenesis explicitly being Author Contributions
the precursor of temperature specialization. The clade containing RAS: Conceptualization-Lead, Data curation-Lead, Formal analysis-Lead, Funding 
Americerura (mostly tropical), Cerura (mostly temperate), and acquisition-Lead, Investigation-Lead, Methodology-Lead, Validation-Lead, 
Kamalia (mostly tropical) exemplify this scenario well, in that they Visualization-Lead, Writing – original draft-Lead, Writing – review & editing-
had a widespread ancestor but regional specialization, possibly due Lead. PZG: Conceptualization-Supporting, Funding acquisition-Supporting, 
to extirpation, leading to three climate-limited genera. Investigation-Supporting, Methodology-Supporting, Project administration-
As shown by our food plant ASR (Fig. S11), the ancestral cater- Supporting, Resources-Supporting, Supervision-Equal, Validation-Supporting, 
pillar feeding condition was unequivocally on Salicaceae, with rare Writing – original draft-Supporting, Writing – review & editing-Supporting. JSM: 
Conceptualization-Supporting, Investigation-Supporting, Supervision-Supporting. 
instances of recently divergent feeding behavior. And while there is 
AM: Data curation-Supporting, Methodology-Supporting, Resources-Supporting, 
evidence that temperature has impacted Cerurinae diversification, Writing – review & editing-Supporting. HSS: Data curation-Supporting, Writing – 
the nearly uniform extant and ancestral Salicaceae-feeding in this review & editing-Supporting. AYK: Data curation-Supporting, Funding acquisition-
group supports the hypothesis that, conversely, food plant shifts on Supporting, Resources-Supporting, Software-Supporting, Supervision-Supporting, 
the scale of plant family did not impact diversification. This latter Writing – review & editing-Supporting. SEM: Data curation-Supporting, 
point is important because food plant shifts and ability to colonize Project administration-Supporting, Supervision-Supporting, Writing – review & 
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22 Insect Systematics and Diversity, 2023, Vol. 7, No. 2
editing-Supporting. RKR: Conceptualization-Supporting, Funding acquisition- Cai L, Xi Z, Lemmon EM, Lemmon AR, Mast A, Buddenhagen CE, Liu L, 
Supporting, Investigation-Supporting, Methodology-Supporting, Project Davis CC. The Perfect Storm: gene tree estimation error, incomplete lin-
administration-Supporting, Supervision-Equal, Visualization-Supporting, Writing eage sorting, and ancient gene flow explain the most recalcitrant ancient 
– original draft-Supporting, Writing – review & editing-Supporting. Angiosperm clade, Malpighiales. Syst Biol. 2020:70(3):491–507. https://
doi.org/10.1093/sysbio/syaa083
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