Sawflies out of Gondwana:

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Argidae (Hymenoptera) 1st Edition
Leonardo A. Malagón-Aldana
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Systematic Entomology (2021), DOI: 10.1111/syen.12527

Sawflies out of Gondwana: phylogenetics and

biogeography of Argidae (Hymenoptera)
L E O N A R D O A . M A L A G Ó N- A L D A N A 1,2 , A R N R . J E N S E N 2,3 ,
D A V I D R . S M I T H 4 , A K I H I K O S H I N O H A R A 5 and
L A R S V I L H E L M S E N2
1
Museo entomológico UNAB, Grupo Sistemática de Insectos Agronomía SIA, Facultad de Ciencias Agrarias, Universidad Nacional
de Colombia, Bogotá, Colombia, 2 Natural History Museum of Denmark, Science, University of Copenhagen, Copenhagen, Denmark,
3
Department of Biology and Biotechnologies ‘Charles Darwin’, Sapienza University of Rome, Rome, Italy, 4 Systematic Entomology
Laboratory, Agricultural Research Service, U. S. Department of Agriculture, c/o National Museum of Natural History, Smithsonian
Institution, Washington, District of Columbia, U.S.A. and 5 Department of Zoology, National Museum of Nature and Science,
Tsukuba, Japan

Abstract. We present the first phylogenetic hypothesis for the cosmopolitan family
Argidae based on both molecular and morphological data. Furthermore, we present
a biogeographic scenario based on a dated phylogeny and interpret the evolutionary
history of the family. Information from sequences of eight genes is analysed to recon-
struct the phylogeny of the Argidae based on maximum likelihood and Bayesian infer-
ence. Total evidence Bayesian analyses are also conducted, combining the molecular
dataset with data from adult and larval morphology. For the historical biogeographic
reconstruction, divergence times are estimated using node dating with uncorrelated
relaxed-clock analysis, and ancestral biogeographical distributions are estimated apply-
ing a Dispersal Extinction Cladogenesis model and a Bayesian binary model. Argidae
s.s. is retrieved as monophyletic in all analyses and the clade Zenargidae + (Argidae +
Pergidae) is better supported when combining molecular and morphological charac-
ters, and when excluding the saturated third codon position in the molecular analysis.
Within Argidae, two large clades corresponding to the subfamilies Arginae and Ster-
ictiphorinae sensu Benson are retrieved as monophyletic. The ancestral distribution of
Arginae and Sterictiphorinae is estimated to be the Australian and Neotropical regions.
Divergence of Argidae-Pergidae and Arginae-Sterictiphorinae is estimated to occur in
the middle-upper Jurassic before or during the east-west Gondwana breakup. Diversi-
fication of Argidae may be associated with the radiation of angiosperms in the Early
Cretaceous, especially for the Neotropical Sterictiphorinae after the separation of South
America and Africa. Arginae were probably introduced to the northern hemisphere
by dispersal to Africa and/or India and subsequent continental collision with Eura-
sia in the Cenozoic. The occurrence of Sterictiphorinae in the northern hemisphere is
more difficult to explain. Maternal care and brood guarding behaviour evolved inde-
pendently in Argidae and Pergidae, with a single origin in a subclade (Dielocerini) of
Sterictiphorinae.

Correspondence: Leonardo A. Malagon-Aldana, Museo entomológico UNAB, Grupo Sistemática de Insectos Agronomía SIA, Facultad de
Ciencias Agrarias, Universidad Nacional de Colombia, Cra. 30 #45-03, Bogotá, D.C., Colombia. E-mail: landresmalagon@gmail.com

© 2021 The Royal Entomological Society 1

2 L. A. Malagon-Aldana et al.
Introduction
Argidae is the second largest family in the Tenthredinoidea or
true sawflies. The Tenthredinoidea in turn is the largest clade
among the primarily herbivorous ‘Symphyta’, which are para-
phyletic in relation to the predominantly carnivorous Apocrita
(Vilhelmsen & Turrisi, 2011). Argidae are distributed world-
wide, being absent only from Antarctica, Madagascar, Green-
land and New Zealand. Unlike most other families of true
sawflies, they have a strong representation in the southern hemi-
sphere, especially in the Neotropics. The closely related Pergi-
dae is almost exclusively Gondwanan in distribution, occurring
mostly in Australia and South America, with only the genus
Acordulecera extending into North America. Argidae + Pergi-
dae have been placed as sister groups in virtually all comprehen-
sive phylogenetic treatments of ‘Symphyta’ (Vilhelmsen, 1997,
2001; Schulmeister et al., 2002; Schulmeister, 2003a; Schul-
meister, 2003b; Ronquist et al., 2012a; Malm & Nyman, 2015).
The split between Pergidae and Argidae has been estimated to
have occurred at approx. 133 Ma (116–158 Ma), around the time
of the Gondwana break up (Schmidt & Walter, 2014; Nyman
et al., 2019). Until recently, no comprehensive phylogenetic
hypothesis was available for Argidae and no biogeographical
scenario in a cladistic framework has been proposed.
Malagón-Aldana et al. (2021a, 2021b) analysed comprehen-
sive data sets of Argidae based on adult (all 56 extant genera
represented) and larval morphology (21 genera included). Their
main finding was that the monotypic Australian genus Zenarge
(Fig. 1K) that was previously placed in Argidae did not group
with the remainder of the family (Argidae s.s.), but instead was
placed as either sister to Pergidae, or most frequently sister to
(Argidae s.s. + Pergidae); regardless of its position, Zenarge
was always in a monophylum with Argidae s.s. and Pergidae
(the ZAP clade). Consequently, Malagón-Aldana et al. (2021a)
placed Zenarge in a separate family, the Zenargidae. This con-
trasts with the results of the few previous molecular analy-
ses of ‘Symphyta’ that included Zenarge in their taxon sam-
ple (Malm & Nyman, 2015), where the genus was retrieved as
sister to Argidae s.s.; comparatively few exemplars of Argidae
were included in these analyses. Malagón-Aldana et al. (2021a,
2021b) retrieved two major clades within Argidae s.s., corre-
sponding to the subfamilies Arginae (Fig. 1A–J) and Steric-
tiphorinae (Fig. 2) as defined by Benson (1963) (Table 1). Other
classification schemes for Argidae recognizing more subfami-
lies and tribes have been proposed (e.g., Smith, 1992; Taeger
et al., 2010). Malagón-Aldana et al. (2021a) were unable to cor-
roborate these, although some previously suggested tribes (e.g.,
Trichorhachini, Atomacerini, Dielocerini and Sterictiphorini)
were retrieved as monophyletic. Revising the tribal classifica-
tion of Argidae s.s. will require a denser taxon sampling than
was obtained in Malagón-Aldana et al. (2021a).
The fossil record of Argidae includes only seven represen-
tatives (Nel, 2004), all of them relatively young and from
the northern hemisphere: the compression fossil Sterictiphora
konowi (Rohwer) from the Florissant formation of Colorado
(37.2–33.9 Ma, upper Eocene – lower Oligocene); the Domini-
can amber fossils Didymia dominicana Smith, D. davisi Smith,
© 2021 The Royal Entomological Society, Systematic Entomology, doi: 10.1111/syen.12527
D. protera Smith, D. poinari Smith and D. ebena Smith
(20.4–13.6 Ma, lower Miocene), and the compression fos-
sil Mioarge azari Nel, from volcano-sedimentary maar of
Mont-Charay in France (7.2–5.3 Ma, uppermost Miocene)
(Rohwer et al., 1908; Smith & Poinar, 1992; Nel, 2004; Pale-
obiology database, fossilworks.org).
Some Neotropical genera of Argidae s.s. (Themos, Dielocerus,
Pachylota, Digelasinus and Sericoceros) present a characteristic
subsocial behaviour, where the females guard the eggs at least
until they hatch, defending them from predators. The larvae stay
together, feeding gregariously and make a mass cocoon when
they pupate. However, this general behaviour includes variation
within several traits among these genera (defence mechanisms
of the female, host plant species, egg location and distribution on
leaves, total period of guarding, mass cocoon structures, etc.)
(Benson, 1938; de Souza Dias, 1975; de Souza Dias, 1976;
Smith, 1992; Ciesla, 2002; Smith & Janzen, 2003; Boraschi
et al., 2005). A similar maternal guarding behaviour is also
present in some Pergidae (Perga, Pseudoperga, Philomastix,
Cladomacra and Syzygonia) (Benson, 1938; Shinohara, 1986;
Schiff, 2004; Schmidt et al., 2006). Larvae of Argidae s.s. Feed
on a diverse variety of angiosperms. Although the larvae of
most argids remain unknown, they also present various degrees
of gregariousness, chemical defence, feeding behaviour and
morphological adaptations to their host plants (Smith, 1992;
Schmidt et al., 2000; Boevé et al., 2018). Contrary to the
known Pergidae and Argidae s.s., the larvae of Zenarge feed
on gymnosperms, including the native Australian Callitris spp.
(Cupressaceae) (Moore, 1963a, 1963b).
Here we aim to reconstruct a phylogenetic hypothesis for the
Argidae, based on molecular characters including mitochon-
drial, ribosomal and nuclear gene fragments, for the highest
number of argid terminals sequenced so far. We also analyse
the combined morphological (adults and larvae) and molecu-
lar data sets in a total evidence approach to further explore the
phylogeny of Argidae, including node dating calibration of the
phylogeny using available fossil information for the family and
outgroup (other Tenthredinoidea families). Finally, we attempt
to elucidate the biogeographical and ecological patterns (e.g.,
brood guarding behaviour) for Argidae.
Methods
Specimens examined
Adult and larval specimens of 45 species and 18 different
genera of Argidae and five outgroup taxa of Tenthredinoidea
(Pergidae, Tenthredinidae) were successfully extracted for
DNA (Table S1). The majority of the specimens were pre-
served in 96% ethanol, while some of them were dry-pinned
specimens. Exemplars were obtained from the following col-
lections: Instituto de Investigaciones Biológicas Alexander
von Humboldt, Colombia (IAvH); Colección de Zoología,
Instituto de Ciencias Naturales ICN, Universidad Nacional de
Colombia; Bogotá, D.C. (ICN-MHN); Museo Javeriano de
Historia Natural, Colombia (MPUJ); Statens Naturhistoriske
 Evolution of Argidae 3

A B C

D E

F G

H I

J K
Fig. 1. (A–C) Head, frontal view of males, subfamily Arginae. (A) Cibdela janthina. (B) Pampsilota dahomeyanus. (C) Spinarge fulvicornis.
(D–K) Lateral habitus of females. (D–J) Subfamily Arginae. (D) Antargidium dentivalve. (E) Scobina lepida. (F) Pampsilota afer. (G) Triarge nigra.
(H) Athermantus imperialis. (I) Tanyphatnidea microcephala. (J) Spinarge fulvicornis. (K) Family Zenargidae, Zenarge turneri.

Museum, University of Copenhagen, Denmark (NHMD);
National Museum of Natural History, Smithsonian Institution,
Washington D.C., U.S.A. (NMNH); National Museum of
Nature and Science, Tsukuba, Japan (NMNS); Senckenberg
Deutsches Entomologisches Institut, Müncheberg, Germany
(SDEI); Museo entomológico UNAB Facultad de Ciencias
Agrarias Bogota, Universidad Nacional de Colombia (UNAB).
DNA extraction, amplification and sequencing
One hindleg (or piece of larval tissue) was removed from each
specimen (Table S1), breaking it at the coxa and trying to include
parts of the basal extrinsic muscles of the appendage. The dis-
section forceps were cleaned with bleach (10%) and ethanol
(70%) prior to working with a new specimen. The samples were

© 2021 The Royal Entomological Society, Systematic Entomology, doi: 10.1111/syen.12527

4 L. A. Malagon-Aldana et al.

A B C

D E

F G

H I

J K
Fig. 2. Head, frontal view of males, subfamily Sterictiphorinae. (A) Trichorhachus nitidus. (B) Aproceros leucopoda. (C) Atomacera coerulescens.
Habitus of females of Argidae, subfamily Sterictiphorinae. (D) Trichorhachus nitidus. (E) Atomacera coerulescens. (F) Sterictiphora furcata. (G) Ptenos
leucopodus. (H) Duckeana prodiga. (I) Pachylota audouinii. (J) Schizocerella pilicornis. (K) Didymia martini.

transferred to separate vials and then stored in 96% ethanol.
DNA extraction and amplification were carried out in the Geo-
genetics lab of the Globe Institute, University of Copenhagen,
Denmark. Total DNA was extracted using the DNeasy® Blood
and Tissue kits using spin columns manufactured by Qia-
gen (QIAGEN Aarhus, Denmark) with a slight modification
entailing soaking the samples overnight for around 12 h at 56∘ C
with gentle agitation. To remove surface contamination from old
dry-pinned specimens, the legs were removed and initially put in
a tissue lysis buffer (ATL buffer + proteinase K) for 1 h at 56∘ C
with gentle agitation. The lysis buffer was then discarded, and
fresh lysis buffer was added. The samples where then incubated

© 2021 The Royal Entomological Society, Systematic Entomology, doi: 10.1111/syen.12527

 Evolution of Argidae 5

Table 1. List of taxa examined specimens of Argidae and Zenargidae taxa.

Examined species (or morphospecies)

Family Genus Species Molecular Morphology
Argidae (Arginae) †Mioarge Nel, 2005 1 0 1
Antargidium Morice, 1919 6 3 2
Arge Schrank, 1802 395 a 23 10
Asiarge Gussakovskij, 1935 3 0 1
Athermantus Kirby, 1882 2 a1 1
Cibdela Konow, 1899 14 1 1
Kokujewia Konow, 1902 3 1 1
Pseudarge Gussakovskij, 1935 6 0 1
Pseudosinarge Saini, Thind & Kaur, 1998 1 0 1
Scobina Lepeletier & Serville, 1828 56 a5 2
Sinarge Forsius, 1934 1 0 1
Sjoestedtia Konow, 1907 2 0 1
Spinarge Wei, 1998 13 a5 1
Styphelarge Benson, 1938 1 0 1
Tanyphatnidea Rohwer, 1912 2 0 1
Triarge Forsius, 1931 9 a1 2
Zhuhongfuna Wei & Nie, 1999 1 0 1
Argidae (Sterictiphorinae) Acrogymnia Malaise, 1941 11 0 2
Acrogymnidia Malaise, 1955 4 0 1
Adurgoa Malaise, 1937 4 0 1
Aproceros Malaise, 1931 10 a1 1
Aprosthema Konow, 1899 58 a4 1
Atomacera Say, 1836 35 a3 2
Brachyphatnus Konow, 1906 3 0 1
Brevisceniana Wei, 2005 1 0 1
Dacrogymnia Smith & Malagón-Aldana, 2020 5 a1 1
Didymia Lepeletier & Serville, 1828 26 a1 3
Dielocerus Curtis, 1844 6 a1 1
Digelasinus Malaise, 1937 2 0 1
Duckeana Malaise, 1941 4 0 1
Durgoa Malaise, 1937 4 0 2
Eriglenum Konow, 1900 4 0 1
Mallerina Malaise, 1941 1 0 1
Manaos Rohwer, 1912 12 a1 1
Neoptilia Ashmead, 1898 10 0 1
Neurogymnia Malaise, 1937 7 0 2
Ortasiceros Wei, 1997 6 0 1
Pachylota Westwood, 1841 2 0 1
Pampsilota Konow, 1899 12 a1 4
Pseudaprosthema Gussakovskij, 1935 3 0 3
Ptenos Norton, 1872 32 a3 5
Ptilia Lepeletier, 1823 7 a2 2
Schizocerella Forsius, 1927 5 2 1
Sericoceros Konow, 1905 21 1 2
Sphacophilus Provancher, 1888 52 a1 3
Sterectiphora Billberg, 1820 45 a4 2
Subsymmia Malaise, 1955 2 0 1
Tanymeles Konow, 1906 23 a1 2
Themos Norton, 1867 13 1 2
Topotrita Kirby, 1882 3 0 1
Trailia Cameron, 1878 2 0 1
Trichorhachus Kirby, 1882 4 0 2
Triptenus Malaise, 1937 2 0 1
Trochophora Konow, 1905 2 1 1
Yasumatsua Togashi, 1970 3 0 1
Zynzus Smith, 1992 19 0 1
Sterictiphorinae males, unknown genus - 1 0
Genus sp. - 0 1
Zenargidae Zenarge Rohwer, 1918 1 1 1
a Genera with at least one molecular sequence produced in this study.

© 2021 The Royal Entomological Society, Systematic Entomology, doi: 10.1111/syen.12527

6 L. A. Malagon-Aldana et al.

Table 2. Primers and annealing temperature used for gene fragment amplification.

Gene Primer Name Primer sequence (5′ –3′ ) References Annealing T∘ C

16S A Hym (F) TRACTGTRCAAAGGTAGC Schulmeister (2003b) 47–50

B Hym (R) TTAATTCAACATCGAGGTC Schulmeister (2003b) 47–50
28S D2F (F) CGGGTTGCTTGAGAGTGCAGC Schmidt & Walter (2014) 47
D2Ra (R) CTCCTTGGTCCGTGTTTC Schmidt & Walter (2014) 47
CAD 743 nF-ino (F) GGIGTIACIACIGCITGYTTYGARCC Johanson & Malm (2010) 55–60
CAD hym 2F (Fi) GGNAGYTCNATGAARAGYGTNGG Malm & Nyman (2015) 55–60
1028R-ino (R) TTRTTIGGIARYTGICCICCCAT Johanson & Malm (2010) 55–60
COI COI fly-C1-J-1514 (F) CAAATCATAAAGATATTGG Farrell (2001); Malm & Nyman (2015) 56
sym-C1-J-1718 (Fi) GGAGGATTTGGAAAYTGAYTAGTWCC Nyman et al. (2006) 52
A2590 (R) GCTCCTATTGATARWACATARTGRAAATG Normark et al. (1999) 52–56
GLN GLN 1F (F) GTNTAYTTYTGGCARGG Regier et al. (2008) 51
GLN hym 2F (Fi) GCNGGVAAYATGGGHTGGYTNAC Regier et al. (2008) 56
GLN 4R (R) CYCCANCCRTGRAARCAYTT Malm & Nyman (2015) 51–56
PGD PGD hym 3F (F) TGGTRCACAAYGGMATHGARTAYGG Malm & Nyman (2015) 60
PGD hym intRb (R) ATRATRCANCCDCCYCKCCACAT Malm & Nyman (2015) 60

Forward (F = external, Fi = internal) and Reverse (R) sequences.

overnight or over 2 days for around 12–48 h at 56∘ C with
gentle agitation. DNA extracts were stored in a freezer at
−20∘ C. DNA concentration of each sample was measured with
a Qubit™ 4 Fluorometer and a Qubit dsDNA High Sensitivity
Assay Kit.
The following six gene regions were amplified: Mito-
chondrial ribosomal 16S, nuclear ribosomal 28S (Schul-
meister, 2003b; Schmidt & Walter, 2014), mitochondrial
cytochrome oxidase I (COI) and the nuclear protein-coding
genes carbamoyl-phosphate synthetase (CAD), gelsolin (GLN)
and phosphogluconate dehydrogenase (PGD) (Malm &
Nyman, 2015). PCR was performed using 5× HOT FIREPol®
Blend Master Mix (10 mM MgCl2 ) manufactured by Solis Bio-
Dyne (Tartu, Estonia), with a final volume of 25 μL, composed
of: 17 μL of the HOT FIREPol® Blend Master Mix, 4 μL of
molecular biology grade H2 O, 1 μL of Dimethyl sulfoxide
DMSO, 0.5 μL of forward and reverse primer each and 1 μL of
the sample DNA extract. PCR profiles and primer sequences for
COI, CAD, PGD and GLN followed Malm & Nyman (2015)
except that the number of cycles was increased to 50 and the
annealing temperature for each gene was modified (Table 2).
In the case of two forward primers (external and internal) we
used primarily the external (Table 2), while the internal primer
was used only when the external did not work for the specific
sample. For 16S and 28S we used the primers from Schmidt &
Walter (2014) but modifying the PCR profile as follows: denatu-
ration at 94∘ C for 1 min, followed by 40 (16S) or 30 (28S) cycles
of denaturation at 94∘ C for 40 s, annealing for 40 s, and exten-
sion at 72∘ C for 1 min; the last cycle was followed by a final
extension step at 72∘ C for 1 min. The PCR products were visual-
ized under a UV chamber after separating the fragments with gel
electrophoresis using 2% agarose with GelRed® (Merck Milli-
pore, Darmstadt, Germany) and a 100 bp ladder. PCR products
that showed sharp visualized bands were sent for purification
and Sanger sequencing at Macrogen Europe (Amsterdam,
Netherlands).
Additional sequences
GenBank databases https://www.ncbi.nlm.nih.gov/genbank/
(Benson et al., 2017) and Barcode of Life Data System
http://www.boldsystems.org/index.php/ (Ratnasingham &
Hebert, 2007) were employed to obtain 216 additional
sequences, including 83 Argidae (32 species, 17 genera)
and 133 of outgroup taxa (33 species, 23 genera) (Table S1)
for the six different markers we sequenced ourselves, as well
as sequences of the genes TPI (triose-phosphate isomerase) and
IDH (isocitrate dehydrogenase), which were suggested to be
phylogenetically relevant by Malm & Nyman (2015) (Table S1).
The 83 downloaded Argidae sequences were distributed across
genes as follows: COI (29), GLN (8), TPI (8), CAD (9), PGD
(9), IDH (6), 16S (7) and 28S (7). These sequences were
produced by Schulmeister et al. (2002), Schulmeister (2003b),
Linnen & Farrell (2008), Heraty et al. (2011), Leppänen
et al. (2012), Boevé et al. (2013), Klopfstein et al. (2013), Isaka
& Sato (2014), Liston et al. (2014), Schmidt & Walter (2014),
Malm & Nyman (2015), Schmidt et al. (2017), Shinohara
et al. (2016), Endara et al. (2018), Smith et al. (2019) and
Prous et al. (2019). The total number of Argidae species or
morphospecies (i.e., produced in this work and obtained from
databases) included in the molecular analyses were 70 in 25
genera, for a total of 72 terminals (Table S1, Fig. 3). For
those outgroup terminals with several species represented in
the sequence sample (chimeras), the combined terminal taxon
was given as the genus name, e.g., Athalia spp.* consists of
sequences from Athalia circularis (COI, GLN, IDH, and TPI)
and A. scutellariae (28S, CAD and PGD).
Alignment and assembling of sequences
The sequences produced were first submitted to the Basic
Local Alignment Search Tool (BLAST®) on the GenBank

© 2021 The Royal Entomological Society, Systematic Entomology, doi: 10.1111/syen.12527

 Evolution of Argidae 7

Fig. 3. Bayesian tree reconstruction of Argidae s.s. Based on the combined molecular information of first and second codon positions of protein codon
genes using Maximum likelihood (ML) and Bayesian inference (BI) analyses. Capital letters followed by an asterisk show the five calibration points
used in the combined morphological and molecular dataset based on the fossil species to the left. Asterisk after the name of a terminal (e.g., Athalia
spp.) indicates that this taxon is a chimera, i.e., the molecular information from various markers was obtained from different species of the same genus.
PP = posterior probability, BR = Bootstrap resampling support.

© 2021 The Royal Entomological Society, Systematic Entomology, doi: 10.1111/syen.12527

8 L. A. Malagon-Aldana et al.
website to cross-check the identifications and detect poten-
tial contaminations (family or genus level). Chromatograms
of sequences were visualized using finchtv 1.5.0 where
low-quality ends were trimmed. Forward and reverse trimmed
sequence chromatograms of the same sample were aligned,
edited and assembled using codoncode aligner software
9.0.1 (CodonCode Corporation). Low-quality sequences (for-
ward, reverse, or both) were excluded and amplified again.
The assembled sequences of each gene per species were sub-
mitted to GenBank ® (Table S1). Sequences from the sam-
ples and the additional downloaded sequences (Table S1)
were put together and first aligned for each gene in mega
X for macOS (Stecher et al., 2020) using the integrated
multiple sequence alignment algorithm clustalw (Larkin
et al., 2007). For the protein-coding genes (COI, PGD, GLN,
CAD), the amino acid alignments were also checked for stop
codons. Introns were detected by eye as ambiguously aligned
regions with several gaps or inconsistent translations and their
start-end was assessed by a typical nucleotide pattern ‘AG/GT
(start) – AG/CAG (stop)’ that signifies splicing sites (Klopf-
stein & Ronquist, 2013; Malm & Nyman, 2015). The identified
introns were individualized and separated from the exons. Pre-
liminary alignments of the protein-coding genes were realigned
using mafft version 7 online service (Katoh et al., 2019) and
an iterative refinement G-INS-i method with global homol-
ogy. The 16S and 28S rDNA sequences were realigned using
T-Coffee (Notredame et al., 2000), which combines a local
and global pairwise alignment, improving the final alignment
for those sequences with secondary structure information. All
final gene-alignments were concatenated using sequencema-
trix 1.8.0 (Vaidya et al., 2011).
Saturation substitution rate
The level of substitution saturation for the protein-coding
genes was assessed for the first and second codon position
together, and the third codon position separately using dambe7
(Xia & Xie, 2001). First, we visually inspected the plot of the
p-distance in terms of transitions and transversions (observa-
tions/length of sequence) against the corrected genetic distance
pairwise comparison using GTR substitution model (=GTR dis-
tance) (Salemi, 2009). In addition, for each gene we estimated
the random saturation substitution index (Iss) and the corrected
substitution index (Iss.c) (Xia et al., 2003).
Molecular phylogenetic inference
The concatenated alignments were analysed under Maximum
Likelihood (ML) and Bayesian Inference (BI) to reconstruct the
phylogenetic hypotheses and examine topological congruence
between the methods. Gaps were treated as missing data. The
trees were rooted on Blasticotoma filiceti (Blasticotomidae)
in all analyses, a family that has been recovered as sister
group of all other Tenthredinoidea in most previous studies
(Vilhelmsen, 2001; Schulmeister, 2003a; Schulmeister, 2003b;
Malm & Nyman, 2015).
© 2021 The Royal Entomological Society, Systematic Entomology, doi: 10.1111/syen.12527
Data of the concatenated alignments were primarily
partitioned per gene, i.e., 16S, 28S, CAD, GLN, TPI, COI,
PGD and IDH. In addition, the protein-coding genes (CAD,
GLN, TPI, IDH, COI and PGD) were partitioned by codon
position: First and second codon position together and the third
codon position separately. To compare the effect of the third
codon position in the phylogenetic reconstructions, we ran all
analyses including and excluding this codon position for the
protein-coding genes. Similarly, we compared the influence of
the addition or exclusion of the TPI and IDH (Table S1) in the
reconstructed phylogeny.
ML trees were inferred using the Web interface of
W-IQ-TREE (Nguyen et al., 2015; Trifinopoulos et al., 2016).
The best substitution and best partition scheme model for
each partition was estimated in the same programme using the
W-IQ-TREE Modelfinder algorithm (Chernomor et al., 2016;
Kalyaanamoorthy et al., 2017) and selected according to the
Bayesian information criterion scores (Schwarz, 1978), with
an ‘edge unlinked’ partition model allowing a different set
of branch lengths for each partition. Bootstrap resampling
support (BR) was calculated with 100 standard nonparametric
replicates. ML trees were visualized in the Interactive tree of
life (ITOL) v4 portal (Letunic & Bork, 2019) and edited with
corel photo-paint x7 (Corel Corporation).
For the BI, we ran the analyses in the CIPRES Science
Gateway V. 3.3 (Miller et al., 2012) portal. The best substitutions
models and best partition scheme for BI were estimated in
Partition Finder2 (Lanfear et al., 2017) using the ‘greedy’
search algorithm, ‘unlinked branch lengths’ option (Lanfear
et al., 2012) and were selected based on the corrected Aikaike
Information Criterion. A BI tree for the combination of all genes
was reconstructed using MrBayes 3.2.2 (Ronquist et al., 2012b),
while the parameters of substitution model were set according
to the Partition Finder2 estimation. Two runs with four Markov
chains Monte Carlo (MCMC) chains (one cold and three
heated) were run for 20 million generations; diagnostics were
saved every 1000th generation and 25% of the samples were
discarded (burn-in fraction) prior to the summary statistics. The
summarized parameter results of each run were visualized and
examined in tracer 1.7.1 plotting the log likelihood values
against the generation time. Convergence of the Markov chains
Monte Carlo chains within and among runs was considered
sufficient by when there was a correct mixing behaviour in
tracer plots, the potential scale reduction factor was close to 1,
the average standard deviation of split frequencies was below
0.01 and the Effective Sample Sizes for all parameters was
higher than 200. The majority rule consensus tree was visualized
using figtree v1.4.4 (http://tree.bio.ed.ac.uk/software/figtree/)
and edited in corel photo-paint x7 (Corel Corporation).
Total evidence phylogenetic analysis: Molecules
and morphology
An additional total evidence approach using Bayesian tree
inference was performed combining the data matrix of 223 adult
anatomical characters (117 taxa) (Malagón-Aldana et al., 2021a)

and 85 larval characters (48 taxa) (Malagón-Aldana
et al., 2021b) with the molecular dataset. For the adult mor-
phological character matrix, seven characters (41, 45, 87, 89,
161, 162 and 175) were treated as ordered in accordance with
Malagón-Aldana et al. (2021a), and three additional characters
(5, 27, 39) based on the result patterns of the larval morphology
dataset (Malagón-Aldana et al., 2021b). The molecular and
morphological datasets were assembled using mesquite 3.6
(Maddison & Maddison, 2019). For some genera represented by
different species in the molecular and morphological datasets
we composed a terminal taxon (taxon denoted as genus name
+ spp.*) if there was evidence for monophyly of the genus in
previous studies (Malagón-Aldana et al., 2021a). Trees were
rooted on B. filiceti (Blasticotomidae). The parameter settings
for the molecular partition were identical with the above; for
the morphological dataset a standard discrete model was used
with coding set to ‘variable’ and rates to ‘gamma’. The BI
analysis for the combined dataset was run using mrbayes 3.2.2
(Ronquist et al., 2012b) in the CIPRES Science Gateway V.
3.3 (Miller et al., 2012) portal. Two runs with four Markov
chains Monte Carlo (one cold and three heated) were run for
20 million generations, and remaining parameters and test of
convergence were identical to the above-described molecular BI
analysis. The majority rule consensus tree was visualized using
figtree v1.4.4 and edited in corel photo-paint x7 (Corel
Corporation).
Molecular clock calibration and divergence dating
To date the divergence times within Argidae and Pergidae,
we used a node dating (tree calibration by employing priors on
node ages based on the oldest known fossils for correspond-
ing clades) approach (Ronquist et al., 2012a; Zhang, 2017).
Five calibration nodes were assigned (Fig. 3) based on fossil
information on well supported nodes (PP = 90–100) and con-
strained as monophyletic. We used an offset exponential prob-
ability distribution as suggested by Ronquist et al. (2012a),
with a mean age of 235 Ma (maximum oldest fossil age of
Hymenoptera, Triassoxyela Rasnitsyn) and a minimum age for
each selected node according to the fossil (Fig. 3): (A) For the
root (=tree age) Pseudoxyelocerus bascharagensis Nel et al.
(183 Ma, oldest fossil of Tenthredinoidea); (B) for the node
ZAP: Zenarge + (Argidae + Pergidae), Xyelotoma macroclada
Gao et al. (164.7 Ma, oldest fossil closest to Argidae + Pergi-
dae) (Ronquist et al., 2012a); (C) for the node Sterictiphora, the
fossil S. konowi Rohwer (37.2 Ma) (Zhelochovtzev & Rasnit-
syn, 1972); (D) for the node of ‘Neotropical Sterictiphorinae’ the
fossil Didymia davisi Smith & Poinar (20.4 Ma) and (E) for the
Arginae (except Antargidium and Scobina) the fossil Mioarge
azari Nel (7.2 Ma) (Malagón-Aldana et al., 2021a).
For the node dating of the combined dataset, we used mrbayes
3.2.2 (Ronquist et al., 2012b). The analyses were run in
CIPRES Science Gateway V. 3.3., with an IGR (independent
gamma rate at each branch with mean 1.0) model (Ronquist
et al., 2012a). The remaining parameters followed Ronquist
et al. (2012a) for Hymenoptera: Lognormal (−7.08069, 1) and
© 2021 The Royal Entomological Society, Systematic Entomology, doi: 10.1111/syen.12527
Evolution of Argidae 9
prset igrvarpr = exp (37.12). We used two different priors to
compare their effect on the age estimation, an uniform tree
prior (clock:uniform) and a birth-death prior (clock:birth-death).
For the latter, the following settings were used: Speciation
prior = exponential (10), extinction prior beta (1, 1) and
extant sample proportion fixed to 0.1 (our sample of Argidae
taxa was about 10% of all described species). Substitution
models were assigned to each gene partition according to
the models estimated previously with Partition Finder2. The
analysis had two runs, four chains for 20 million generations
and a majority rule consensus tree was estimated, with the same
parameters and test of convergence and effective sample size
used for the above-described Bayesian phylogenetic analysis
(see Section 2.6).
Biogeographic reconstruction
The geographic records of genera and species were obtained
from the Electronic World Catalogue of Symphyta (Taeger
et al., 2010, 2018), labels of the examined specimens, literature
(Smith, 1992; Saini, 2009; Koch et al., 2015) and the WaspWeb
(van Noort, 2020). World biogeographic regions were assigned
according to the zoogeographic realms of Holt et al. (2013)
with some modifications from Morrone (2015) (i.e., the Mediter-
ranean and Africa north of Sahara were considered as parts of
the Palearctic region; New Guinea considered part of the Aus-
tralian region). The ancestral biogeographical states were recon-
structed using rasp 10.15 (Yu et al., 2015) applying S-BGB (Sta-
tistical BioGeoBEARS) and BBM (Bayesian Binary MCMC)
methods, estimated over the ultrametric calibrated consensus
tree of the node dating analysis to compare directly with the
divergence time estimations. Regarding S-BGB (Matzke, 2013a,
2013b, 2014), we used the Dispersal Extinction Cladogenesis
(DEC) approach model that implements a Lagrange (Likeli-
hood Analysis of Geographic Range Evolution) source code
(Ree & Smith, 2008) and estimates likelihoods of ancestral
states at cladogenesis events constraining dispersal multipliers
between regions according to certain geological events (Brown
& Lomolino, 1998; SanMartin & Ronquist, 2004; Hall & Sev-
astjanova, 2012; Müller et al., 2019) from 0 to 1 in a matrix
(File S1). We used two dispersal rates, 0.1 and 1 (e.g., dis-
persal between Afrotropical and the Neotropical regions was
set as 1 before their breakup, around 110 Ma and 0.1 after
it). An additional sample of 1000 trees derived from the node
dating Bayesian analysis was used for the S-BGB analysis.
The Bayesian Binary Analysis (BBM) (Ronquist & Huelsen-
beck, 2003; Yu et al., 2015) was run with 10 chains and 500 000
generations, state sampling every 100 generations; and apply-
ing a F81 model with a site rate variation of gamma, which
is the most complex model implemented in rasp 10.15. The
maximum number of ancestral states was set to three in both
analyses. Both S-BGB and BBM analyses accepted polytomies
in the input consensus tree. For the composite Argidae terminal
taxa = *genus + spp. (two species in the same genus from dif-
ferent datasets) the distribution state was assigned as a combina-
tion of both, e.g., Aprosthema tardum (Palearctic) + Aprosthema
10 L. A. Malagon-Aldana et al.
brunniventre (Nearctic) = Aprosthema spp.* (Palearctic and
Nearctic).
Results
Sequence assemblage and saturation of substitution rate
The final molecular assemblage of aligned sequences included
99 terminal taxa (72 sequences and 70 species of Argidae and
27 sequences of outgroup taxa) (Table S1, Fig. 3) and 5350
characters or total sites, including 2296 parsimony informative
sites, 490 singletons and 2564 constant sites. The most com-
pletely sampled genes were COI (82%), 28S (69%) and 16S
(61%), while TPI (27%) and IDH (19%) were the least sam-
pled (Table 3), the latter two consisting exclusively of sequences
from the GenBank database. There were 46% missing data.
Sixty sequences contained more than 50% gaps when includ-
ing IDH and TPI. The successful amplification of mitochondrial
and ribosomal DNA (COI, 16S and 28S) was higher than for
the nuclear genes. rDNA 16S alignment was highly incongru-
ent among the different alignment methods due to abundance of
ambiguous sections in its structure. The lengths of the aligned
gene regions were 1015 bp (COI), 711 bp (IDH), 513 bp (PGD),
834 bp (GLN), 783 bp (CAD), 453 bp (TPI), 397 bp (16S) and
644 bp (28S), respectively. For the protein-coding genes COI,
IDH and PGD, no intron regions were detected, while for
CAD, GLN and TPI, one or more introns were present and
excluded. We were able to correct one erroneous identification
from the GenBank database, the species Sericoceros nr. tannuus
(KC975751.1, ZSM HYM AE086, Schmidt & Walter, 2014;
Schmidt et al., 2017; Boevé et al., 2018) was recovered inside
the genus Atomacera in all trees; after checking the photographs
of the voucher specimen in the http://www.boldsystems.org/, we
conclude that this specimen is an Atomacera sp. We provide the
first known sequences for the Sterictiphorinae genera Manaos
(28S, COI, and PGD), Dielocerus (16S, CAD, COI, GLN, PGD
and TPI), Sphacophilus (16S, CAD, COI, GLN, PGD and TPI),
Tanymeles (16S) and for the Arginae genera Athermantus (16S,
COI), Pampsilota (16S, 28S, CAD and COI), Triarge (28S, 16S,
and COI); as well as for several species (Table S1).
Among the protein-coding genes there was a clear difference
in the variation of sites among the first and second codon
positions relative to the third (Table 3), the third codon having
a high proportion of unique and parsimony informative sites
(>98%, >90%) and few invariant sites (= high variation) (<8%).
The rDNA genes were intermediately conserved in terms of
unique and invariant sites, while first and second codon positions
of coding genes were the most conserved partitions (Table 3).
The substitution saturation index (Iss) (Table 3) shows little
saturation (Iss < Iss.c) in the first and second codon position
of all genes, except for COI, which is saturated according
to this index in all three codon positions. A similar pattern
was observed in the graphics of the substitution’s saturation
(Fig. S1). Here the asymptotic behaviour of the observed third
codon of p-distance against the corrected genetic distance
indicates that the substitution rate is saturated, while for the first
© 2021 The Royal Entomological Society, Systematic Entomology, doi: 10.1111/syen.12527
and second position, it shows a linear growth and low saturation
in all genes, including COI.
Gene trees
When analysing individual ML gene trees and all codon posi-
tions (Fig. S2), the monophyly of Argidae s.s. and Pergidae was
recovered only for CAD, GLN, COI, PGD and IDH. The rela-
tionship (Zenarge + Argidae s.s.) was retrieved when analysing
GLN; (Zenarge + Pergidae) for the individual analyses of COI,
PGD and CAD. Zenarge + (Pergidae s.s. + Argidae s.s.) was
recovered for IDH. The 16S gene tree did not recover the mono-
phyly of any of these families, while for the gene 28S only the
family Pergidae was retrieved as monophyletic and for TPI the
family Argidae s.s. was retrieved as monophyletic (Fig. S2).
These families were generally not recovered when using only
the first and second codon for each individual gene tree. An addi-
tional example of contrasting results between the phylogenetic
reconstruction of individual genes was observed for the genus
Atomacera, which was recovered either as being the sister group
of Arginae for CAD, GLN and IDH, or as sister group of Ster-
ictiphorinae for PGD and 28S (see below).
Combined molecular dataset
The branch support was lower in all cases in the ML trees
(BR) relative to the BI trees (posterior probability), even with
similar topology (Fig. S3). The BI trees reconstructed with
only the first and second codon position were less resolved
and supported to species level (e.g., Arge) (Fig. 3), while the
inclusion of the third codon position in the analyses improved
the resolution and support (>50%) at this level. On the other
hand, excluding the saturated third codon position improved the
support of deep nodes (e.g., at the base of the ZAP clade, see
below) (Fig. S3).
The ML and BI showed different topologies for Argidae s.s.,
Pergidae and Zenarge relationships, although in all cases they
formed part of the same clade with high support (BR or posterior
probability PP = 89–100) (Fig. S3). (Zenarge + Argidae s.s.)
was retrieved when using ML and BI with all codon positions
(Fig. S3: A, C). On the other hand, Zenarge + (Pergidae +
Argidae s.s.) was obtained in both analyses when using only the
first and second codon positions (Fig. S3: B, D).
Regarding the internal relationships of Argidae s.s., in the
combined molecular data set using either all codon positions
or only the first and second codon position we recovered two
main clades that correspond to the subfamilies Arginae and Ster-
ictiphorinae s. Benson (1963) (Fig. 3); the clades within each
subfamily are also highly congruent with the morphological
hypotheses (Malagón-Aldana et al., 2021a, 2021b). For Ster-
ictiphorinae, Atomacera was retrieved as sister group of the
remaining genera (Fig. 3). Other clades recovered within the
Sterictiphorinae were the sister relationship between the Hol-
arctic clade Sterictiphora + (Aproceros + Aprosthema) and a
Neotropical clade comprising a number of genera but whose
 Evolution of Argidae 11

Table 3. Descriptive parameters of aligned sequences with corresponding substitution model assigned (IQ-tree and partition finder).

Parsimony
Number Length Unique informative Invariant Substitution Substitution
Gene sequences (bp) sites (%) sites (%) sites (%) model IQ-tree model PF Iss Iss.c P-value

16s 61 397 70.5 44.1 41.8 GTR + F + I + G4 GTR + I + G - - -

28s 69 644 62.3 39.3 46.4 SYM + I + G4 SYM + I + G - - -
COI12 83 677 48.9 26.0 63.8 TIM + F + I + G4 GTR + I + G 1.11 0.72 0.000
COI3 83 338 97.6 89.1 4.4 TIM + F + G4 TIM + G 1.65 0.69 0.000
IDH12 20 474 29.3 15.6 77.0 TIM3 + F + I + G4 GTR + I + G 0.25 0.71 0.000
IDH3 20 237 98.7 93.7 3.0 TNe + G4 GTR + I + G 0.72 0.63 0.005
PGD12 45 342 40.1 19.0 73.4 SYM + I + G4 GTR + I + G 0.39 0.69 0.000
PGD3 45 171 98.8 92.4 7.6 K2P + I + G4 GTR + I + G 0.78 0.70 0.260
TPI12 29 302 43.0 19.5 64.2 K2P + I + G4 SYM + I + G 0.26 0.67 0.000
TPI3 29 151 99.3 95.4 3.3 TPM3 + F + I + G4 TRN + I + G 0.66 0.61 0.139
CAD12 54 522 50.6 18.4 71.5 TIM3 + F + I + G4 GTR + I + G 0.44 0.71 0.000
CAD3 54 261 99.2 93.9 5.7 TPM2 + F + I + G4 GTR + I + G 0.73 0.68 0.035
GLN12 48 556 45.7 14.2 71.9 GTR + F + I + G4 GTR + I + G 0.57 0.71 0.000
GLN3 48 278 98.2 89.9 4.7 HKY + F + I + G4 TRN + I + G 0.79 0.68 0.000

internal relationships were not resolved (Fig. 3). Concerning
Arginae, the Australian genus Antargidium was recovered as the
sister group of the remaining genera. The Neotropical genus
Scobina is the sister group of the remaining Arginae, minus
Antargidium.
Total evidence analysis and phylogenetic relationships
The combined morphological and molecular dataset com-
prised 164 terminal taxa (52 genera of Argidae out of 55) and
5658 characters (File S2). The genera Pseudosinarge, Zhuhong-
funa and Mioarge (fossil) were excluded from the morpholog-
ical matrix and the analyses due to the low number of scored
characters and their negative effect in the resolution of Arginae.
When combining molecular and morphological data using BI
and all codon positions, we retrieved Zenarge + (Pergidae +
Argidae s.s.) (Figs 4, 5). Pergidae and Argidae were recovered
as monophyletic sister groups, both with high branch support
(PP = 100). With regard to Pergidae (Fig. 5), the new world
subfamily Acordulecerinae is retrieved as the sister group
of the remaining genera, which in turn are divided in two
main clades (Fig. 5). One clade includes Phylacteophaginae +
(Perginae + Parasyzygoninae) (PP = 96), while the second clade
has Loboceratinae (Aulacomerus) (PP = 89) as sister group
of the remaining subfamilies: (Styracotechynae + Heteroper-
reyia) + ((Pterygophorinae + (Philomastiginae + (Euryinae +
(Syzygoninae + Perreyinae)). Remarkably, the South American
Perreyinae was recovered as paraphyletic, Heteroperreyia being
outside the subfamily.
Inside Argidae, two main clades were retrieved for all the dif-
ferent reconstructed phylogenies. They correspond to the sub-
families Arginae (PP = 100) and Sterictiphorinae (PP = 66)
(Figs 4, 5), as in the above molecular analyses and previ-
ous morphological phylogenetic hypotheses (Malagón-Aldana
et al., 2021a, 2021b). For the subfamily Arginae (Fig. 4), the
Australian genus Antargidium (Fig. 4, clade I) was retrieved
as the sister group of the rest, followed by the Neotropi-
cal Scobina (Fig. 4, clade II). Clade III (Fig. 4) comprises
exclusively African taxa (PP = 100), i.e., the genus Tri-
arge, Pampsilota and African Arge spp. In clade III, Pamp-
silota afra was recovered as sister group of the remaining
taxa. Pampsilota is paraphyletic, P. afra being apart from the
other species, P. dahomeyanus, which is in a clade with Arge
congrua and A. fenestralis. Triarge is recovered as mono-
phyletic and sister group of Arge montana and A. stuhlmanni.
Clade III includes two subclades, one consisting of Atherman-
thus + Sinarge (Oriental-Sino-japanese) and a second compris-
ing Cibdela + (Tanyphatnidea + Sjoestedtia) (Oriental-African)
(Fig. 4). Clade III was recovered as the sister group of the last
major clade of the subfamily that comprises the majority of the
ubiquitous genus Arge (Fig. 4, clade V), with a number of other
genera embedded in the clade (Asiarge, Kokujewia, Brevisceni-
ana, Pseudarge and Spinarge).
In Sterictiphorinae (Fig. 5), the Australian Tri-
chorhachus + Styphelarge (clade VI) were retrieved as sister
group of the remaining genera, followed by Atomacera (clade
VII) from the Nearctic and Neotropical regions. A third
clade includes the Holarctic genera (clade VIII), among them
Sterictiphora was the sister group of the remaining genera
Aprosthema + (Pseudaprosthema, Ortasiceros and Yasumat-
sua + Aproceros). Finally, the remainder of the Sterictiphorinae
were placed in a large, predominantly Neotropical clade
(clade IX) with males having the flagellum furcate, Brachy-
phatnus (southern South America) being the sister group of
the remaining genera. The deeper relationships of the latter
are poorly resolved, but some subclades have high support
(PP ≥ 70) (Fig. 5). These subclades are: clade X, Ptenos
(paraphyletic), Manaos and Zynzus; clade XI, Acrogymni-
dia + (Acrogymnia + Trailia); clade XII, the ‘Dielocerini s.l.’
clade (Smith, 1992) including Sericoceros, Subsymmia, Durgoa,
Neoptilia, Themos, Mallerina, Topotrita, Pachylota, Dielocerus
and Digelasinus and clade XIII, Schizocerella, Didymia, Ptilia,
Trochophora, Triptenus and Neurogymnia + Eriglenum, where

© 2021 The Royal Entomological Society, Systematic Entomology, doi: 10.1111/syen.12527

12 L. A. Malagon-Aldana et al.

Fig. 4. Total evidence Bayesian tree of subfamily Arginae (Argidae) highlighting the current biogeographical distribution. Main clades are labelled
with numbers in circles. Asterisk after the name of a terminal (e.g., Athalia spp.) indicates that this taxon is a chimera, i.e., the morphological and
molecular information was obtained from different species of the same genus.

© 2021 The Royal Entomological Society, Systematic Entomology, doi: 10.1111/syen.12527

 Evolution of Argidae 13

Fig. 5. Total evidence Bayesian tree of subfamily Sterictiphorinae (Argidae) highlighting the current geographical distribution. Main clades are labelled
with numbers in circles. Asterisk after the name of a terminal (e.g., Athalia spp.) indicates that this taxon is a chimera, i.e., the morphological and
molecular information was obtained from different species of the same genus. Colours of occurrence in Pergidae and Zenargidae (Neotropical, Australian
and Nearctic region) are lighter than in legend to emphasize the distribution in Argidae s.s.

© 2021 The Royal Entomological Society, Systematic Entomology, doi: 10.1111/syen.12527

14 L. A. Malagon-Aldana et al.
the genus Didymia was not recovered monophyletic. We were
unable to properly resolve the exact position of Sphacophilus.
Node dating and ancestral distribution (Figs 6, 7)
The estimated mean age of the clades was slightly older when
using the uniform tree prior than with the birth-death prior. In
the same way the 95% highest posterior density (HPD interval)
was wider for the uniform tree prior (Fig. 6; Table 4); here we
describe the results using the shorter HPD interval. The age of
the ZAP clade was estimated to be 177.1 Ma (164.7–215.8),
e.g., early Jurassic; this agrees with the estimate of Ronquist
et al. (2012a). The biogeographical reconstructions (both
S-BGB/DEC and BBM) indicate an ancestral distribution of
the ZAP ancestor in the Neotropics or Australia + Neotropics
(Fig. 7). The divergence between Pergidae and Argidae s.s.
occurred approx. 168.5 Ma (144.9–205.4). This interval of
time overlaps with other estimates (Schmidt & Walter, 2014;
Nyman et al., 2019) and is contemporary with the beginning of
the breakup between west and east Gondwana. The first split
within Argidae s.s. is dated from the middle to late Jurassic
around 155.1 Ma (127.4–191.1 Ma). The age of the Steric-
tiphorinae crown group is estimated to be around 148.2 Ma
(118.6–183.2 Ma) in the beginning of the Cretaceous, signif-
icantly older than crown group Arginae at approx. 123.3 Ma
(92.1–159.5). The ancestral area of Argidae s.s. was inferred
to be Australian/Neotropical (BBM) or Australian/African
(S-BGB/DEC); for each subfamily the ancestral distribu-
tion was estimated also as Australian/Neotropical (BBM and
S-BGB/DEC) (Fig. 7).
Discussion
Although for many taxa the overall gene sampling was low
(Table S1), especially for the nuclear protein-coding genes, the
phylogenetic relationships retrieved with the available molec-
ular data (Fig. 3) are highly congruent with previous hypothe-
ses, e.g., monophyly of Argidae, Pergidae, Sterictiphorinae and
Arginae. However, for more than half of the genera of Argidae,
molecular information is still absent, and therefore the relation-
ships of these taxa rely exclusively on morphological evidence.
Saturation of substitution rate, additional data and their
phylogenetic effect
Our results show how the type of data differentially affect
the placement of taxa, with special effect on isolated taxa,
e.g., Zenarge turneri (Zenargidae) (Fig. S3). The placement
of Zenarge was highly influenced by the third codon position,
and when excluding it, the phylogenetic position of Zenarge
was more similar in the molecular and morphological trees
(Malagón-Aldana et al., 2021a, 2021b), Zenarge being the
sister group of Pergidae + Argidae s.s. However, for more
distal nodes in the topology, e.g., within Arge, the resolution
© 2021 The Royal Entomological Society, Systematic Entomology, doi: 10.1111/syen.12527
decreased substantially when excluding the third codon position
(Fig. 3). This indicates that while the third codon position
might have achieved saturation for the deeper nodes, this may
not be the case for the more distal nodes; a similar effect
occurs in implied weights analyses, where different k-values
might resolve some parts of the phylogeny better than others
do. Using the highly saturated nucleotides in phylogenetic
analyses is still controversial (Källersjö et al., 1999; Strimmer
& von Haeseler, 2009; Breinholt & Kawahara, 2013). Malm
& Nyman (2015) retrieved Zenarge + Argidae s.s. even if
excluding the third codon position for most of the protein-coding
genes they used, except for CAD and GLN.
Additionally, when we combined the molecular and morpho-
logical datasets, including all codon positions, Zenarge was
recovered as sister group of Argidae + Pergidae. This shows that
morphological characters, in spite of their significantly lower
number relative to molecular sites, can still influence the results
of the combined analyses when using few genes. The sampling
of additional fresh material of Zenarge (we relied on GenBank
sequences), together with a phylogenomic approach would help
to test its current placement. For now, based on the morpho-
logical (adult and larval stages) and molecular evidence anal-
ysed here, we uphold placing Zenarge in a separate family, the
Zenargidae (Malagón-Aldana et al., 2021a).
Zenargidae
The combined analysis supports the topology Zenargidae
+ (Argidae + Pergidae) or ZAP clade (Figs 4, 5). Zenarge
turneri, the sole representative of Zenargidae, only occurs in
New South Wales in Australia. It is the only taxon of the ZAP
clade that feeds on gymnosperms, the endemic Cypress pine
Callitris as well as the introduced Cupressus (Cupressaceae)
(Moore, 1963a, 1963b; Smith, 1992). The position of Zenarge as
sister to the rest of the ZAP clade and our estimation of the age
of the split between Zenargidae and Argidae + Pergidae around
180 Ma in the early Jurassic, prior to the Gondwana breakup
(Fig. 6), is apparently in conflict with the inference of Nyman
et al. (2019) that the common ancestor of the ZAP clade (indeed,
of all Tenthredinoidea) was feeding on angiosperms and the
feeding habits of Zenarge a result of a secondary shift to gym-
nosperms. The origin of the Zenargidae is according to our esti-
mates of similar age as the earliest putative angiosperm flower
fossil (Nanjinganthus dendrostyla; Fu et al., 2018 of 174 Ma),
older than the age estimate (120 Ma, i.e., Early Cretaceous) of
the split between Zenarge and the remaining Argidae accord-
ing to Nyman et al. (2019). Different lines of evidence sup-
port the pre-Cretaceous origin of angiosperm plants (Hochuli &
Feist-Burkhardt, 2013; Fu et al., 2018); however, fossil evidence
indicates that the major radiation of angiosperms did not take
off until the Early Cretaceous (Silvestro et al., 2015; Herendeen
et al., 2017; Nyman et al., 2019). The evolution of the Tenthredi-
noidea seems to display a similar pattern (Nyman et al., 2019)
and they may still have co-evolved with angiosperms even if
their common ancestor in the early Jurassic was feeding on gym-
nosperms. However, the estimated age of Callitris, the current
 Evolution of Argidae 15

Fig. 6. Node dating chronogram of Argidae using the combined dataset (morphology and molecules). Horizontal bars show the 95% highest posterior
density interval (HPD) of the estimated node age, in blue for the combined dataset and red for the molecular data. Vertical bars show relevant geological
events in that transition period of time. Main clades are labelled with numbers in circles.

© 2021 The Royal Entomological Society, Systematic Entomology, doi: 10.1111/syen.12527

16 L. A. Malagon-Aldana et al.

Fig. 7. Biogeographic ancestral reconstruction of Argidae, subfamily Arginae (left) and Sterictiphorinae (right) showing Bayesian Binary Analysis
(BBM) estimations in colour charts, and S-BGB/DEC Dispersal Extinction Cladogenesis (DEC) in white pentagons. Main clades are labelled with
numbers in circles.

© 2021 The Royal Entomological Society, Systematic Entomology, doi: 10.1111/syen.12527

 Evolution of Argidae 17

Table 4. Divergence times of main clades (Fig. 8) based on node dating Bayesian analysis, using a uniform tree prior (UN) and a birth-death prior
(BD).

95% Highest
posterior density Age 95% HPD
Node Age (Ma) UN (HPD) (Ma) UN (Ma) BD (Ma) BD

Zenarge + (Argidae + Pergidae) 187.3 164.7–246.9 177.1 164.7–215.8

Argidae + Pergidae 182.1 156.4–241.8 168.5 144.9–205.4
Sterictiphorinae + Arginae 170.2 141.3–226.3 155.1 127.4–191.1
I + remaining 139.4 108.3–189.8 123.3 92.1–159.5
II + remaining 120.7 87.1–167.1 109.8 78.7–140.4
III + remaining 75.5 54.5–103.1 88.2 63.1–122.6
IV + remaining 72.6 52.7–95.2 77.8 54.1–109.8
V + remaining 67.6 49.1–89.5 66.9 46.2–90.1
VI + remaining 164.8 135.9–220.6 148.2 118.6–183.2
VII + remaining 154.7 127.4–207.5 135.7 110.5–170.7
VIII + remaining 138.2 110.9–186.8 124.5 102.3–156.9
IX 117.4 88.8–157.3 112.5 86.7–145.2

host plant of Zenarge, indicates that gymnosperm feeding in
the latter might be secondary rather than ancestral. Callitris is
a genus of drought-resistant plants that evolved around 30 Ma,
probably in response to climate change in the middle Cenozoic
as Australia drifted northwards (Pittermann et al., 2012; Larter
et al., 2017). The Zenarge lineage cannot have switched to
Callitris earlier than this, whether it was from an angiosperm
or a gymnosperm host.
Pergidae
The reconstruction of ancestral biogeographic area of Pergi-
dae (Fig. 7, BBM, S-BGB/DEC) is congruent with the hypoth-
esis of Gondwanan origin (Neotropical or a combination of
Neotropical and Australia) with a mean age of 150 Ma for the
crown group, similar to the estimation (140 Ma) of Schmidt &
Walter (2014). Our results suggest, in accordance with Schmidt
& Walter (2014), a pattern of repeated dispersals between
South America and Australia through Antarctica before the final
separation of the three continents around 35 Ma (Figs 6, 7).
Schmidt & Walter (2014) suggested that several pergid lineages
(Perginae, Phylacteophaginae, and Pterygophorinae) radiated on
Myrtaceae (Eucalyptus and their relatives) as the Australian cli-
mate became more arid during the Cenozoic era, while other
lineages experienced reduced diversity or went extinct.
The combined analysis (Fig. 5) placed the new world sub-
family Acordulecerinae (Acordulecera + Anathulea), as sis-
ter group of the remaining genera; this in contrast to Schmidt
& Walter (2014) that placed Loboceratinae (Aulacomerus)
as sister to all other Pergidae. Although both subfamilies
have a similar distribution in South America, Acordulecerinae
have a higher diversity, especially within the genus Acordule-
cera that also extends into North America. Previous studies
(Smith, 1990; Schmidt & Walter, 2014; Malm & Nyman, 2015;
Malagón-Aldana et al., 2021a) suggested that the subfamily
Acordulecerinae belongs in a clade with Phylacteophaginae and
Perginae from Australia. Regarding Loboceratinae, all studies to
date have only included one genus, Aulacomerus, and the mono-
phyly of the subfamily itself remains uncertain.
The two main clades of Pergidae retrieved (Fig. 5) are mostly
congruent with previous studies (Schmidt & Walter, 2014;
Boevé et al., 2018) in terms of subfamily composition, although
their relationships partly differ; this might be caused by dif-
ferences in taxon and sequence sampling. Remarkably, we
retrieved the Australian Heteroperreyia as sister group of the
Australian Styracotechys (Styracotechyinae); this makes the
Neotropical subfamily Perreyinae paraphyletic since Heteroper-
reyia is placed apart from Perreyia + Decameria, similar to what
was found in larval morphological analysis (Malagón-Aldana
et al., 2021b); they differ substantially in the ovipositor structure
(well developed in Heteroperreyia, reduced in, e.g., Perreyia
and Decameria) (Smith, 1990; Malagón-Aldana et al., 2021a)
and larval life history (saprophagous/mycophagous in Per-
reyia/Decameria, phytophagous in Heteroperreyia) (Schmidt &
Walter, 2014; Smith et al., 2019).
The Neotropical Parasyzygoniinae was retrieved as sister
group of Perginae (Australia), and Syzygonia (Syzygoninae)
as sister group of Perreyinae (Neotropics). Parasyzygoniinae is
a monotypic subfamily with only two species (Parasyzygonia
cyanoptera and P. pallidor) from Venezuela and Brazil, which
has been previously suggested to be close to Perginae by
Smith (1990) and Malagón-Aldana et al. (2021a). With regard
to the Neotropical Syzygoninae, Schmidt & Walter (2014)
included only the genus Lagideus, and they retrieved it as sister
group of Philomastiginae (Philomastix). The monophyly of the
subfamily Syzygoninae (e.g., Syzygonia and Lagideus) has not
been properly tested.
Argidae relationships
Our phylogenetic results (Figs 4, 5), based on the combined
analysis, are highly similar to the morphological phylogenetic
hypotheses (Malagón-Aldana et al., 2021a, 2021b) with two
large clades, i.e., the subfamilies Arginae and Sterictiphorinae.

© 2021 The Royal Entomological Society, Systematic Entomology, doi: 10.1111/syen.12527

18 L. A. Malagon-Aldana et al.
In general terms, Arginae correspond to the taxa where males
have an undivided antennal flagellum (Fig. 1A–C) and Ster-
ictiphorinae to those where males have a furcate antenna
(Fig. 2A,B). The genus Atomacera is the only genus where the
antenna of males is simple (Fig. 2C) but recovered inside the
Sterictiphorinae, see below.
Within Arginae, the position of Antargidium (clade I) and
Scobina (clade II) as successive sister groups to the rest of
the subfamily is stable and well supported by both molecular
and morphological data (Figs 3, 4); these two genera retain
some plesiomorphies, including the presence of a complete
cupula in the male genitalia and two separated suprapedal
lobes in their larval stage (Malagón-Aldana et al., 2021a,
2021b). Clades I and II (Fig. 4) are congruent with clades
1 and 2 of the adult morphology analysis (Malagón-Aldana
et al., 2021a) and partly congruent with clade A of the larval
analysis (Malagón-Aldana et al., 2021b). We retrieve a clade of
African taxa (Pampsilota, Triarge and Arge) (clade III) that is in
conflict with the current genus classification and questions the
diagnostic value of some of the morphological characters that
this classification is based on, e.g., the occurrence of preapical
tibial spurs (absent in Pampsilota, present in Arge) and the
presence of the 2r-m vein in the fore wing (absent in Triarge,
present in Arge). According to the adult morphological analysis
of Malagón-Aldana et al. (2021a, fig. 20H), the presence of long
volsellar teeth on the digitus of the males could be considered a
putative synapomorphy for clade III; more Afrotropical Arginae
need to be examined to confirm this.
The clades III and IV (Fig. 4) are partly congruent with
clades 6a and 6b of Malagón-Aldana et al. (2021a) and the
tribe Athermantini of Benson (1963). The genera of clade
IV lack the preapical tibial spur in mid and hind tibiae and
are restricted to the Oriental/Sino-Japanese region, except for
Sjoestedtia (Afrotropics). Clade IV includes a subclade with
two small genera, Athermantus (two species) and Sinarge (one
species), for which we were only able to sequence mitochon-
drial markers (COI and 16S) for Athermantus. In the sec-
ond subclade within clade IV, Cibdela is retrieved as the sis-
ter group of the remaining genera. Pampsilota scutellis is
retrieved inside Sjoestedtia + Tanyphatnidea; the three taxa
have parapenis lobes in the male genitalia, the third valvu-
lae of females strongly angulated dorsolaterally and an epis-
tomal sulcus present but weak (Malagón-Aldana et al., 2021a).
Additionally, P. scutellis also has the anal cell present in
the hind wing, as in Tanyphatnidea. The genera Tanyphat-
nidea and Pampsilota (recovered paraphyletic here) have been
defined differently by taxonomists (Togashi, 1975; Wei, 1997a;
Saini, 2009); however, in Pampsilota the parapenis lobes of
males and epistomal sulcus are absent, and dorsolateral cor-
ners of the third valvulae of females are not angulated. In
addition, Tanyphatnidea is restricted to southern China, Taiwan
and the Oriental region, while Pampsilota species are endemic
to the Afrotropics (Liston et al., 2017). We did not examine
the Chinese species Pampsilota cenchra, although the descrip-
tion by Wei (1997a) suggested high similarity with P. scutellis.
Regarding hostplants of clade IV, Cibdela janthina larvae feeds
on Rubus alcifolius (Rosacae) (Florens et al., 2017), whereas
© 2021 The Royal Entomological Society, Systematic Entomology, doi: 10.1111/syen.12527
Tanyphatnidea sinensis feeds on Polygonum sp. (Polygonaceae)
(Togashi, 1980).
The remaining, non-Afrotropical Arge spp. (clade V, Fig. 4)
are placed in a monophylum that includes several other small
genera from the Palearctic region. Of these, Kokujewia and
Spinarge have been sampled for both molecular (only COI for
Kokujewia) and morphological data; Asiarge, Brevisceniana
and Pseudarge only for morphology. Kokujewia, Asiarge and
Brevisceniana form a clade primarily defined by the absence of
preapical tibial spurs and the short apical maxillary palpomere.
Due to the large number (approx. 400) of described species of
Arge, it is standard practice to define species groups to facilitate
its classification (e.g., Smith, 1989); we retrieved some of these
as monophyletic (Fig. 4): The pectoralis group, ochropus group
and clavicornis group. Malagón-Aldana et al. (2021a) retrieved
Arge ochropus as sister to a large clade comprising the majority
of the Arginae included, which could be due to the retention
of plesiomorphic conditions in the ovipositor with a simple,
flat third valvula, from which more complex shapes of the third
valvula might have evolved. However, here the Arge ochropus
group (Fig. 4) (Smith, 1989; Yan et al., 2009) is in a polytomy
with other species of Arge that have a highly modified ovipositor
sheath. Host plants associations within clade V seem to follow
the oligophagous patterns of other temperate herbivores (Haack
& Mattson, 1993) with preference for woody, shrubby and
herbaceous food plants belonging predominantly to the families
Rosaceae, Betulaceae, Fagaceae, Ericaceae and Salicaceae
(Boevé et al., 2018); Some examples of argids with high host
specificity are Kokujewia ectrapetala on Rumex spp. (Polyg-
onaceae), Arge berberidis on Berberidis spp. (Berberidaceae)
and the Arge ochropus group on Rosa (Rosaceae).
A large amount of evidence supports the inclusion of Spinarge
in Arge (Figs 3, 4, clade V) (Boevé et al., 2018; Malagón-Aldana
et al., 2021a, 2021b). However, a clear delimitation of Arge s.s.
is still necessary. Since Afrotropical species of Arge are more
closely related to other genera (Triarge and ‘Pampsilota’) than
to the rest of Arge species, it further complicates the taxonomic
delimitation of the genus. One alternative could be to define Arge
s.s. as the non-Afrotropical species (clade V, Fig. 4). A larger
sample of Afrotropical species of Arge than ours (four out of
125 species) (Taeger et al., 2010) is needed to corroborate this.
The branch support (PP = 66) for the Sterictiphorinae is lower
than for Arginae (PP = 100). This might be caused by the
lack of molecular data for Trichorhachus and Styphelarge, and
the retention of several plesiomorphic character states in adult
and larval morphology in Trichorhachus (e.g., preapical tib-
ial spurs present in adults and simple setae in epipharynx and
lacinia of larva) (Malagón-Aldana et al., 2021a, 2021b). Nev-
ertheless, these two genera form clade VI (Fig. 5) which is
congruent with clade 7 of Malagón-Aldana et al. (2021a) and
the tribe Trichorhachini of Benson (1963). Clade VI is the sis-
ter group of the remaining Sterictiphorinae. The Neotropical
genus Atomacera (clade VII, Fig. 5) is recovered within Ster-
ictiphorinae as suggested by Benson (1963) (tribe Atomacerini)
and Malagón-Aldana et al. (2021a, 2021b), although in some
individual gene trees it was placed closer to Arginae. It was
retrieved as sister group of Arginae + Sterictiphorinae in Malm

& Nyman (2015). In contrast to the putative reversal to a simple
flagellum antenna in the males, a number of characters justify
the placement of Atomacera in Sterictiphorinae, e.g., the pres-
ence of setae multiple branched in the lacinia and epipharynx of
larva (Malagón-Aldana et al., 2021b). The conflicting evidence
from individual gene trees (Fig. S2) as well as adult and larval
anatomy indicates that the exact position of Atomacera needs
further clarification.
Clade VIII (Fig. 5) is well supported (PP = 100) and retrieved
with molecular and total evidence (Figs 3, 5), as well as
previous morphological analyses (clade 8 of Malagón-Aldana
et al., 2021a, clade D of Malagón-Aldana et al., 2021b).
Clade VIII is partly congruent with the tribe Sterictiphotini of
Wei (1997b) but includes the genus Ortasiceros as well. For
the small genera Pseudaprosthema (three species), Ortasiceros
(six species) and Yasumatsua (three species) only type material
exists, so the amount of information that can be extracted from
them is limited. A characteristic larval feeding behaviour has
been described for larvae of some species of clade VIII; they
make sinusoidal channels or zig-zag feeding tracks on their
host plants, which are Prunus spp. (Rosaceae) [Sterictiphora
serotina, S. prunivora] and Ulmus spp. (Ulmaceae) [Aproceros
leucopoda] (Blank et al., 2014; Eiseman, 2015). In contrast,
Aprosthema tardum does not display this behaviour when
feeding on herbaceous Leguminosae (Liston et al., 2018).
For clade IX (Fig. 5) (PP = 100) that includes all the Neotrop-
ical Sterictiphorinae except Atomacera, internal resolution and
support is weak, even if morphological synapomorphies can
be inferred for some subclades (Malagón-Aldana et al., 2021a,
2021b). This Neotropical clade comprises more than half of
all genera of Argidae (29 out of 55) while molecular infor-
mation (mostly COI) is only available for 12 genera. This is
not enough to resolve the polytomy of well supported inter-
nal subclades (Fig. 5, clades X, XI, XII and XIII). Within
subclades of clade IX, some hostplant associations patterns
are evident. For clade X, larvae are associated mainly with
the Fabaceae Inga + Zygia clade (Smith, 1970; Smith, 1992;
Smith & Janzen, 2003; Smith et al., 2013; Endara et al., 2018;
Malagon-Aldana et al., 2019). Endara et al. (2018) suggested a
coevolutionary arms race between Ptenos and Inga, where Inga
defence chemistry played a role in cospeciation with Ptenos.
In clade XIII, the larvae feed on members of Connaraceae
(Kimsey & Smith, 1985; Smith, 1992; Smith & Janzen, 2003;
Smith et al., 2013).
Biogeography of Argidae
The ancestral biogeographic reconstruction of extant Argidae
s.s. is congruent with a Gondwanan distribution with the same
probability for the Australian and the Neotropical region (Fig. 7,
S-BGB/DEC and BBM) during the early breakup between
east and west Gondwana in the middle-upper Jurassic (around
155 Ma, crown group age, Fig. 6). While this corresponds to the
pattern displayed by the Pergidae (Schmidt & Walter, 2014), it
ignores the distributional information from Xyelotoma macro-
clada, a fossil taxon possibly closely related to the ZAP clade
© 2021 The Royal Entomological Society, Systematic Entomology, doi: 10.1111/syen.12527
Evolution of Argidae 19
and used to calibrate this node in the divergence dating analyses
(see Methods). This fossil is from the late Jurassic (164.7 Ma)
and occurs in the Daohugou Formation from China, as well
as Kazakhstan (see http://fossilworks.org/?a=taxonInfo&taxon_
no=139523), i.e., the northern hemisphere (Gao et al. 2009). We
have not been able to study this fossil and integrate it in our
morphological data set; the estimate of its phylogenetic posi-
tion is from Ronquist et al. (2012a). However, even if Xyelo-
toma macroclada was included in our biogeographic analyses,
the otherwise strong Gondwanan signal in the deeper nodes of
the ZAP clade would probably not be overturned.
The radiation of Argidae s.s. in the Early Cretaceous was
probably related to radiation of the angiosperms at the same
time (Sun et al., 1998; Soltis et al., 2008), as it has been
suggested for other sawflies (Nyman et al., 2019). Inside
each subfamily, the same ancestral distribution in Gond-
wana is inferred (Fig. 7), either Australian/Neotropics (BBM)
or Australian/Africa/India-Oriental (S-BGB/DEC). In both
Arginae and Sterictiphorinae, the sister groups of the extant
genera, clades I and VII, Antargidium (Arginae) and Tri-
chorhachus + Styphelarge (Sterictiphorinae), respectively,
are endemic to Australia, while the next clades to branch
off, i.e., II (Arginae: Scobina) and VII (Sterictiphorinae:
Atomacera) (Figs 4, 5), respectively, are Neotropical. Tri-
chorhachus + Styphelarge are restricted to Western Australia,
while Antargidium is mainly present in Eastern Australia; it is
possible that the habitat of their ancestors contracted substan-
tially during the increased aridity in the interior of Australia,
when the Australian plate drifted towards the Equator in the
Miocene (Wilford & Brown, 1994). Some of the first extant
genera to branch off have as current host plants taxa that evolved
much more recently than the ancestors of the sawflies. Antargid-
ium feed on the Australian Sapindaceae Arytera and Alectryon
(20–30 Ma; Muellner-Riehl et al., 2016). Trichorhachus feed
on flowers of the endemic Australian Proteaceae Conospermum
(70 Ma; Lamont & He, 2012) from fire-adapted vegetation
in western Australia; the adults have a long proboscis and
probably use it to reach the nectar of Conospermum small
flowers at the base of the calyx tube, similar to some colletid
bees (Houston, 1989).Unlike some Australian Pergidae that
have apparently experienced modest diversity increase as a con-
sequence of feeding on Myrtaceae (Schmidt & Walter, 2014),
the endemic Australian argid genera have not switched to this
family, another possible cause of their current low diversity.
The early diversification in the Neotropics of both Arginae
and Sterictiphorinae present a similar pattern, occurring in the
lower Cretaceous, i.e., ancestors of Scobina (Arginae clade
II, 78.6–140.35 Ma, Fig. 6) and Atomacera (Sterictiphorinae
clade VII, 86.7–145.2 Ma). This is congruent with the isola-
tion of South America from Africa around 110 Ma (SanMartin
& Ronquist, 2004) in a vicariance event. Both genera are
associated with Malvaceae (Smith, 1992), which presumably
also arose in Gondwana in the late lower Cretaceous or
early upper Cretaceous (Areces-Berazain & Ackerman, 2017;
Hernandez-Gutierrez & Magallon, 2019). Atomacera apparently
dispersed from the Neotropical to the Nearctic region, although
the estimated age of A. debilis, the Nearctic species in our
20 L. A. Malagon-Aldana et al.
sample, is much older than the formation of the Panamian
Isthmus (13–15 Ma) (Montes et al., 2015). However, the cur-
rent host genera of these argid taxa are much younger than
the ancestors of clade II and VII: Scobina larvae feed on the
pantropical Sida rhombifolia (30–40 Ma) and Atomacera on
Hibiscus (30–50 Ma) (Areces-Berazain & Ackerman, 2017).
The major radiation of Sterictiphorinae occurred in the
Neotropical clade IX (Figs 5–7) after diverging from the ances-
tors of the Palearctic clade VIII in the lower Cretaceous. This
probably happened after the isolation of South America from
Africa around 110 Ma. South America can be considered an area
of endemism for Sterictiphorinae, whose genera have a diverse
range of host plant families (Smith, 1992), and the angiosperm
radiation might have influenced the diversification of Steric-
tiphorinae.
The early evolutionary history of Argidae and Pergidae are
similar (Fig. 7), being restricted to Gondwana (Schmidt &
Walter, 2014). The main difference between the two families is
the diversification of Argidae in the rest of the world, the north-
ern hemisphere in particular. How Argidae colonized Eurasia
is somewhat difficult to explain, and the correlation of split-
ting events in the family with tectonic events is hampered by
the wide HPD intervals for the estimated divergence times for
the former (Fig. 6). The dispersal to the northern hemisphere
might have happened through dispersal and diversification in
Africa and/or India in the Cretaceous, which separately drifted
northwards to collide with Eurasia during the early Eocene
(India) and Oligocene (Africa-Arabian Peninsula) (Brown &
Lomolino, 1998). The breakup between West (South America,
Arabia, and Africa) and East Gondwana (Australia, India, and
Antarctica) began around 160 Ma, several Ma after the esti-
mated divergence of Argidae + Pergidae. Subsequently, ances-
tors of extant lineages of Argidae s.s. might have dispersed to
India and/or Africa. However, according to the ancestral biogeo-
graphic reconstructions of Argidae (Figs 7, 8), the Afrotropical
and/or Indian/Oriental distribution can only be inferred for the
ancestors of Arginae but not for Sterictiphorinae, e.g., clades I,
II, III and IV (Fig. 7). Although the African continent collided
with the Palearctic region (through the Arabian Peninsula) later
than India (Brown & Lomolino, 1998), both events might have
influenced the dispersal of clade V to Eurasia, since either of the
two continents might have housed Arginae ancestors (Fig. 8).
According to the HPD interval of the divergence times
(Fig. 6), the ancestors of clade I (92.1–159.5 Ma) and clade
II (78.7–140.3 Ma) could precede the separation of Africa
from South America and the separation of India from Africa
(130-110 Ma), as well as their dispersion to these continents.
However, the ancestors of clades III and IV (distributed cur-
rently in Africa and India/Oriental, respectively) might have
evolved after India and Africa separated from the rest of Gond-
wana, since their estimated age is younger than those events
(75–63 Ma). Assuming that all extant Afrotropical Arge (125
species) belong to this clade, the radiation of the exclusively
Afrotropical clade III could be related to the diversification
of the African austro-temperate and arid floras in the Paleo-
gene and Eocene, although their ancestors might have evolved
together with the radiation of lowland forest angiosperm flora
© 2021 The Royal Entomological Society, Systematic Entomology, doi: 10.1111/syen.12527
in the Cretaceous (Linder, 2014). In clade III, Pampsilota and
Afrotropical Arge spp. have a wide distribution in sub-Saharan
Africa (Koch et al., 2015; Liston et al., 2017), while the genus
Triarge is restricted to South Africa and its austro-temperate
flora. Host plants for Triarge are unknown, while larvae of
Afrotropical species groups of Arge feed on plants of Gera-
niaceae found in different natural biomes of shrub, bush and
grasslands (Koch et al., 2015). Contrary to the Austral king-
dom hypothesis suggested mainly by phytogeographers (Lin-
der, 2014; Morrone, 2015), we did not find taxa with affinities
across southern Africa (e.g., Triarge), southern South America
and Australia.
The extant Arginae occurring in the India-Oriental region
(clade IV) are restricted to northern India, adjacent and par-
allel to the Himalayas (Fig. S4) to the north. Possible expla-
nations for the current absence of clade IV in central and
southern India are: (i) Climatic variation: their ancestors in
southern India moved northwards or went extinct due to a
substantial climatic change to much drier conditions in cen-
tral and southern India, shaped by its northward drift from
Gondwana and the formation of the Himalayas (Chatterjee
et al., 2017) in the early Eocene; (ii) Cretaceous interchange:
their ancestors could have dispersed from northern Africa
(and the Arabian Peninsula) through a land bridge that con-
nect only to northern India from the late Cretaceous before
India collided with Eurasia (Oman-Kohistan-Ladakh [OKL] arc,
Fig. 8C) (Reinert, 1970; Chatterjee & Bajpai, 2016; Chatterjee
et al., 2017); (iii) Mio-Pliocene interchange: their ancestors
could have dispersed much more recently from Africa during
a Miocene interchange after to the collision of the Arabian plate
with Eurasia around 30–15 Ma and before/during the aridifi-
cation of Sahara, as it has been suggested for other organisms
with similar distribution (Bibi, 2011; Portik & Papenfuss, 2015).
Given the absence of any evidence for the occurrence of Arginae
in India south of the Indo-Gangetic Plain, it is perhaps most
likely that they arrived in the subcontinent from the west some-
time in the Tertiary. Nonetheless, according to our divergence
time estimation, ancestors of clade IV (mostly Oriental taxa)
would be older (54.1–109.8 Ma) than the Miocene interchange
scenario. Records of Argidae in the Arabian Peninsula region
are unknown. Hypotheses 2 and 3 above could also partly
explain the sister relationship between the Afrotropical Sjoest-
edtia and the Oriental (Pampsilota scutellis + Tanyphatnidea) in
clade IV.
Regarding the distribution pattern of clade IV south and par-
allel to the Himalayan chain (Fig. S4), it is possible that the
uplift of this mountain range functioned as a natural barrier
for dispersal of the ancestors of this clade. Oriental represen-
tatives of clade IV are usually collected over 3000 m of eleva-
tion (Saini, 2009), indicating that they have adapted to cooler
conditions. This is in contrast to the occurrence of Argidae in
the Neotropics (e.g., Colombia, Ecuador), where they are only
recorded below 2500 m (Malagon-Aldana et al., 2019). The
genus Cibdela (clade IV, Figs 6–8G) is the only Indian-Oriental
genus that reaches the Australian region in New Guinea, which
could be a more recent dispersal event (e.g., during the collision
of Australia-Wallacea-SE Asia in the early Miocene or during
 Evolution of Argidae 21

A D

B E

C F

Fig. 8. Hypothetic sequence of ancestral node distribution for subfamilies Arginae (A–C) and Sterictiphorinae (D–F) of Argidae in different geologic
times. (G) Current distribution of subfamily Arginae. (H) Current distribution of subfamily Sterictiphorinae. Main clades are labelled with numbers in
circles.

© 2021 The Royal Entomological Society, Systematic Entomology, doi: 10.1111/syen.12527

22 L. A. Malagon-Aldana et al.

A B C

D E

Fig. 9. Mass cocoons constructed by individuals of Sterictiphorinae species. (A) Sericoceros dimidiatus Konow, 1908 (NMNH). (B) Themos olfersii
(Klug, 1834) (NMNH). (C) Dielocerus diasi Smith, 1975 (NMNH). (D) Digelasinus diversipes (Kirby, 1882) (NMNH). (E) Pachylota audouinii
Westwood, 1841 (NHMUK). NMNH = National Museum of Natural History, Smithsonian Institution, Washington D.C., USA, NHMUK=Natural
History Museum, London, UK.

the glaciations of Pleistocene caused by falling sea levels and
exposition of glacial land bridges) (Brown & Lomolino, 1998;
Hall & Sevastjanova, 2012), since their ancestral distribu-
tion is estimated to be Indian-Oriental (S-BGB/DEC, BBM,
Figs 4, 7).
The radiation of the subfamily Arginae is most prevalent in the
non-Afrotropical Arge (clade V) (+250 species) in the Palearc-
tic and Sino-Japanese regions and occurred between 38 and
61 Ma, around the time estimation of the collision between
India and Asia, 45–50 Ma, that formed the Himalaya-Tibetan
plateau (Fig. 6) (Chatterjee et al., 2017) and produced severe
climatic shifts in the Oriental region. Clade V has a current dis-
tribution in northern India similar to clade IV, although with
a substantial lower number of species (20 taxa, 13 endemic)
(Saini, 2009) than in the Sino-Japanese region where there
are more than 200 species (Wei et al., 2006; Hara & Shino-
hara, 2018; Taeger et al., 2018). This includes several Spinarge
spp. that diversified mainly in Japan after the opening of the
Japanese sea (Fig. 6) around 15.3–33.7 Ma. More recently,
Arge spp. of clade V reached the Nearctic, at least in two dif-
ferent dispersal events, represented by Arge cyra (15.7 Ma)
and A. quidia + A. pectoralis (16.2 Ma) (Figs 4, 7), proba-
bly through one of the Bering land bridges that occurred at
different times in the Cenozoic (Hopkins, 1959; SanMartin
et al., 2001).
In Sterictiphorinae, no African or Indian taxa have been
found that can link the primarily Neotropical clades VII and
IX and the mostly Palearctic clade VIII. Only one species of
Sterictiphorinae has been recorded from the Oriental region,
Aproceros sikkimensis Saini & Thind, 1992 (Saini, 2009); this
species occurs in northernmost India and was not included here.
No reliable records of Sterictiphorinae exist from the Afrotrop-
ical region, except for one of the otherwise Neotropical Spha-
cophilus (recorded as Sterictiphora afra by Pasteels, 1963);
this record is probably based on a mislabelled specimen (Lis-
ton et al., 2017). The virtual absence of Sterictiphorinae from
the Afrotropical and Oriental regions may have been caused
by extinction or failure to colonize these regions. Either way,
it is not possible to establish a connection between Neotropi-
cal and Eurasian members of the subfamily. The ancestral dis-
tribution of clade VIII is estimated to be Palearctic (Figs 7,
8). The clade is distributed throughout the Holarctic and com-
prises more than 100 species, primarily in the genera Ster-
ictiphora (41 spp.) and Aprosthema (55 spp.). Ortasiceros (6
spp.) and Yasumatsua (3 spp.) are endemic to the Sino-Japanese
region, as are 8 out of 10 species of Aproceros; the species

© 2021 The Royal Entomological Society, Systematic Entomology, doi: 10.1111/syen.12527


Aproceros leucopoda has been introduced to Europe (Blank
et al., 2014). Sterictiphora and Aprosthema reached the Nearctic
in the Miocene (Figs 5, 8), presumably through a Beringian
connection.
Subsocial behaviour and maternal care
According to our results, the subsocial behaviour of females
within Argidae and Pergidae evolved independently; this
behaviour cannot be ascribed to the ground plan of either
family. In Pergidae, subsociality occurs in genera in different
subfamilies (Perginae, Philomastiginae, Perreyinae and Syzy-
goninae) representing at least two different lineages (Schmidt
et al., 2006). In Argidae, it occurs only in clade XII (Fig. 5),
corresponding to the tribe Dielocerini of Benson (1938) or
subfamily Dielocerinae of Smith (1992); the behaviour is not
developed to the same degree in all genera. According to our
phylogeny, clade XII includes several subclades with unre-
solved relationships. Maternal care has not been recorded for
subclade XIIa, Neoptilia + Durgoa, although the larvae of some
species display signs of subsocial behaviour, e.g., Durgoa mat-
togrossensis and Neoptilia tora are gregarious feeders (Smith
et al., 2013) and also build small irregular mass cocoons (D. R.
Smith, pers. observation). In subclade XIIb, Sericoceros shows
maternal care (Ciesla, 2002), while for its sister group Sub-
symmia the life history is unknown. In subclade XIIc, Themos
clearly shows maternal care and subsocial behaviour, together
with the genera Pachylota, Dielocerus and Digelasinus, while
for Topotrita and Mallerina, the biology is unknown (de Souza
Dias, 1975; de Souza Dias, 1976; Smith, 1992; Ciesla, 2002;
Smith & Janzen, 2003; Boraschi et al., 2005).
In Sericoceros the maternal care behaviour is more weakly
developed than in other genera of subclade XIIc that show
this trait (Ciesla, 2002; Smith & Janzen, 2003). The less
developed subsocial behaviour of Sericoceros (construction of
small and irregular mass cocoons, low levels of aggressiveness
when guarding the eggs by the female and shorter duration of
maternal care) might suggest that it displays the initial steps of
evolving this behaviour complex. However, within clade XIIc
the same characters do not seem to follow a transformation
series (number of days of guarding and degree of defence
aggressiveness by the female), except for the construction of
much more complex and organized mass cocoons in all genera
(Fig. 9). One probable ancestral trait in this behaviour complex
present in Sericoceros and Themos (sister group of the remaining
taxa in clade XIIc) is that females lay the eggs directly on
the leaf epidermis and in circular/oval egg clusters instead of
inside the mesophyll of the leaves and scattered over the entire
leaf as in Dielocerus and Digelasinus (Benson, 1938; de Souza
Dias, 1975; de Souza Dias, 1976; Smith, 1992; Ciesla, 2002;
Smith & Janzen, 2003; Boraschi et al., 2005). Larvae of clade
XII share the presence of two or more non-eversible coxal
glands (Malagón-Aldana et al., 2021b); however, it is unknown
if this feature is related to gregarious or subsocial behaviour.
A morphological convergence between adult females that show
maternal care in Pergidae (e.g., Pseudoperga) and clade XII
© 2021 The Royal Entomological Society, Systematic Entomology, doi: 10.1111/syen.12527
Evolution of Argidae 23
are the long plantulae on the tarsi, mainly on the basitarsomere
(Malagón-Aldana et al., 2021a, char. 59).
Conclusions
Our study is the most comprehensive phylogenetic analysis for
the Argidae to date, although the limited amount of fresh mate-
rial and the prevalence of monotypic or small genera represented
only by type material made it impossible to reconstruct a better
resolved phylogeny. We provide the first molecular sequences
of some genes for the genera Athermantus, Triarge, Pamp-
silota, Dielocerus, Sphacophilus, Tanymeles and Manaos, as
well as some outgroup species, e.g., Perreyia tropica (Pergidae),
Andeana farcta and Waldheimia amazonica (Tenthredinidae).
We hope that new technologies and collecting efforts will help
to increase the amount of data and evidence for understanding
the evolution of the family.
The placement of Zenarge in a separate family as suggested
by Malagón-Aldana et al. (2021a, 2021b), as indicated by the
relationship Zenargidae + (Argidae + Pergidae), is better sup-
ported when combining molecular and morphological charac-
ters or when excluding the saturated third codon position in the
molecular analysis. Furthermore, the morphological (adult and
larva) and ecological features (e.g., gymnosperm host plant) of
Zenarge clearly makes it unique. According to the calibrated
phylogeny presented here, the more basal position of Zenarge
turneri, which currently is the only species of the ZAP clade
known to feed on gymnosperms, might challenge the hypothetic
scenario of an ancestral angiosperm host for Tenthredinoidea
proposed by Nyman et al. (2019). Alternatively, the host plant
choice of Zenarge is a secondary reversal to feeding on gym-
nosperms, as indicated by the estimated Cenozoic age of its
current native host plant, see above.
Our results agree mostly with previous phylogenetic results
for the family Argidae s.s. showing two main clades, i.e.,
the subfamilies Arginae and Sterictiphorinae (Malagón-Aldana
et al., 2021a, 2021b), both of them with ancestral distribution
in Gondwana, and with an origin coinciding with the breakup
between east and west Gondwana. The biogeographic recon-
struction suggests Arginae dispersed to Eurasia most likely
through northern Africa (and the Arabian Peninsula); however,
for Sterictiphorinae similar evidence is absent. We consider that
the rise of the Himalayas played a substantial role in the diversi-
fication of Arginae, and perhaps Sterictiphorinae, in the Palearc-
tic region, as has been suggested for other organisms in conjunc-
tion with uplift of mountain ranges (Wen et al., 2014; Luebert
& Muller, 2015; Klaus et al., 2016) (Fig. 9A). Regarding sub-
social behaviour, we find that maternal care evolved only once
within Sterictiphorinae; however, further research is necessary
to understand evolution of specific traits of this behaviour com-
plex.
Much more work still needs to be done to resolve the rela-
tionships of Neotropical Sterictiphorinae and the megadiverse
genus Arge, for which we only obtained low resolution. Besides,
Arge in particular needs to be sampled much more densely, and
the genus will probably have to be subdivided, as indicated by
24 L. A. Malagon-Aldana et al.
our results for Afrotropical Arginae. At the same time, a num-
ber of small northern hemisphere genera nested inside Arge
(clade V, Fig. 4), e.g., the ‘prefix’ genera Asiarge, Pseudarge
and Spinarge, might eventually have to be synonymized with the
nominal genus. The genus Sphacophilus is retrieved as mono-
phyletic, contrary to previous hypotheses. Molecular informa-
tion is also needed to confirm the placement of a number of
genera, e.g., Trichorhachus, Tanyphatnidea, Sjoestedtia, Pseu-
daprosthema, Yasumatsua, Ortasiceros, most of the Afrotropical
species of Arge and Neotropical Sterictiphorinae, including the
status of the New World genera Ptenos and Didymia currently
retrieved as paraphyletic. Similarly, a revision of the tribal clas-
sification of Argidae must await a denser sampling than the one
achieved here.
Supporting Information
Additional supporting information may be found online in
the Supporting Information section at the end of the article.
Figure S1. Substitution saturation per codon position (first
and second codon together and third codon separately) per
protein-coding gene. Observed transitions and transversion
vs. corrected genetic divergence.
Figure S2. Maximum likelihood Individual Gene Trees.
Figure S3. Phylogenetic position of Zenarge turneri Rohwer
when using different gene datasets. First and second codon
position together (first-second) and third codon position
(third) separately. Maximum likelihood ML and Bayesian
inference (BI) analyses.
Figure S4. Current occurrence of Oriental genera of clade
IV.
File S1. Dispersal matrix, Dispersal Extinction Cladogenesis
(DEC) analysis.
File S2. Total Evidence Matrices for Node dating.
Table S1. List of taxa used for molecular information and
GenBank accession number per gene (Sequences starting
with OK were produced in this work).
Acknowledgements
We thank COLCIENCIAS Institution of Colombia and the
Natural History Museum of Denmark (NHMD), which funded
the first author’s PhD programme at the University of Copen-
hagen. We are especially grateful to the curators of the differ-
ent entomological collections that provided specimens to this
work: Stephan Blank (SDEI), Fernando Fernández (ICN-MHN),
Dimitri Forero (MPUJ), Andrew Liston (SDEI), Claudia Peña
(ICN-MHN), Andreas Taeger (SDEI), and Valentina Vergara
(UNAB). Thanks to Tobias Malm, Tommi Nyman, and Saskia
Wutke who shared with us all their own original data of Argi-
dae, including the 16S and 28S rDNA sequences of Cibdela
© 2021 The Royal Entomological Society, Systematic Entomology, doi: 10.1111/syen.12527
janthina. Also, thanks to Seraina Klopfstein, Tamara Spasojevic
and Liping Yan, who provided several suggestions and construc-
tive criticism of the maximum likelihood and Bayesian inference
methods applied, as well as to Katja Kramp for her relevant
suggestions on the DNA extraction techniques. Finally, we thank
Seraina Klopfstein and Gavin Broad who reviewed an early ver-
sion of the manuscript. Additional input was provided by Bonnie
Blaimer and two anonymous reviewers. Mention of trade names
or commercial products in this publication is solely for the pur-
pose of providing specific information and does not imply rec-
ommendation or endorsement by the USDA. USDA is an equal
opportunity provider and employer. The authors declare no con-
flict of interest.
Author contributions
Leonardo A. Malagón-Aldana (LAMA) and Lars Vilhelmsen
(LV) conceived and designed the study. David R. Smith and Aki-
hiko Shinohara provided crucial material for DNA extraction.
LAMA and Arn R. Jensen generated the molecular data. LAMA
conducted all analyses. LAMA wrote the first draft of the paper,
which was then reviewed by LV. All authors contributed to the
final version of the paper.
Data availability statement
NCBI Sequence accession numbers are provided in Table S1.
The final total evidence matrix, including the used concatenated
nucleotide alignement and the morphological data set are pro-
vided in File S2. The rest of data that support the findings of this
study are available from the corresponding author upon reason-
able request.
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Tree scale: 1

16S
Blasticotoma filicetiGB
82
Runaria reductaGB
Waldheimia amazonica
Abia sppGB
10
Pterygophorus insignisGB
23
Dracogymnia fernandezi
100 Sericoceros nrtannuusGB
7
Styracotechys dicelysmaGB
Sphacophilus sp1
24
Dielocerus sp
99
Dielocerus larva
Ptenos texanus
8 Aprosthema tardum
89
Aprosthema intermedium
38
14
Aprosthema fusicorne
Sterictiphorinae males
12
Sterectiphorinae males
3
Sphacophilus sp3
14
Sterictiphora geminata
Atomacera pubicornis
16
Aproceros leucopoda
1
Cladius pectinicornisGB
34
Corynis crassicornisGB
Siobla sppGB
23
Sterictiphora angelicae
2 3
Phylacteophaga froggattiGB
19
Pergagrapta sppGB
99
Pseudoperga sppGB
Aulacomerus atriventris
Andeana farcta
3 32
Monoctenus juniperiGB
92
Neodiprion sertiferGB
6
Polyclonus atratusGB
24 Decameria sppGB
99
Perreyia picea
64
Perreyia tropica
Triarge sp
19
Arge berberidisGB
99
Arge berberidis
9 Arge suzuki
3 Cibdela janthina
15
Arge melanochra
51
Arge pagana
2
Athermantus imperialis
24
Arge thoracica
17
Arge gracilicornis
33

4
Arge rejecta
Arge rustica
1
30 Arge obesa
41 Scobina bolivariGB
91
Scobina strophosaGB
92
Scobina thoracicaGB
Arge mali
10 Pampsilota dahomeyanus
34
Antargidium allucenteGB
98
Antargidium atricepsGB
Spinarge metallica
2
36 Spinarge fulvicornisP
85 Spinarge fulvicornisA
87
Spinarge prunivora
35
10 Spinarge affinis
Arge ustulata
80
Arge nigripes
17
Arge aruncus
49 Arge similis
100
Arge similislarva
40
Arge similis2
Tree scale: 0.1

28S
Runaria reductaGB
Euura respondens
Cladius pectinicornisGB
Andeana farcta
Waldheimia amazonica
Siobla sppGB
Corynis crassicornisGB
Neodiprion sertiferGB
Monoctenus juniperiGB
Abia sppGB
Athalia sppGB
Scobina incultaGB
Scobina terminalisMix
Antargidium atricepsGB
Antargidium allucenteGB
Sericoceros nrtannuusGB
Atomacera debilisGB
Atomacera pubicornis
Aproceros leucopodaMix
Aprosthema fusicorne
Aprosthema tardum
Aprosthema intermedium
Sphacophilus sp1
Manaos mulsus
Ptenos texanus
Sterictiphora geminata
Sterictiphora furcataGB
Sterictiphora angelicae
Dracogymnia sp
Tanymeles galumnatus
Sphacophilus sp3
Sterictiphorinae males
Ptilia peleterii
Ptilia versicolor
Phylacteophaga froggattiGB
Acordulecera spMix
Anathulea sp
Aulacomerus atriventris
Pergagrapta sppGB
Pseudoperga sppGB
Styracotechys dicelysmaGB
Pterygophorus insignisGB
Polyclonus atratusGB
Decameria sppGB
Perreyia piceaGB
Perreyia tropica
Arge suzuki
Arge berberidisMix
Arge similis
Cibdela janthina
Arge thoracica
Arge rustica
Arge obesa
Arge gracilicornis
Arge cyanocrocea
Arge ochropus
Arge pagana
Arge captiva
Arge rejecta
Pampsilota dahomeyanus
Triarge sp
Arge scita
Arge melanochra
Arge mali
Arge nigripes
Arge ustulata
Arge metallica
Spinarge pumilla
Spinarge fulvicornisP
Spinarge affinis
Spinarge fulvicornisA
Spinarge prunivora
Tree scale: 0.1

CAD Blasticotoma filicetiGB

Euura respondens
92 Waldheimia amazonica
38 Andeana farcta
48
Siobla sppGB
Neodiprion sertiferGB
49
98
Monoctenus juniperiGB
48
Abia sppGB
33
Athalia sppGB
48
Corynis crassicornisGB
Zenarge turneri
46 Acordulecera spGB
28
Phylacteophaga froggattiGB
Decameria spGB
21
31
Polyclonus atratusGB
28
Pterygophorus insignisGB
47
Pseudoperga spGB
98
Pergagrapta sppGB
Sterictiphora geminata
98 Sterictiphora spGB
80
94
Sterictiphora furcataGB
100
Sterictiphora angelicae
46
Aproceros leucopodaGB
100
Aproceros leucopoda
67
Dielocerus larva
29
Sphacophilus sp1
88 Ptenos texanus
27
Schizocerella pilicornisGB
50
31
Sterectiphorinae males
40
Ptilia peleterii
51
Dracogymnia fernandezi
Atomacera debilisGB
100
Atomacera pubicornis
Scobina terminalisGB
100
31 Scobina terminalis
Arge suzuki
86
Arge rustica
3
Arge obesa
22
Cibdela janthinaGB
12
94 Arge cyanocrocea
10 Pampsilota dahomeyanus
28 Arge rejecta
50
Arge pagana
95
17 Arge ochropus
Arge similislarva
15 Arge scita
28
Arge berberidisGB
100
Arge berberidis
14
Arge melanochra
34
Arge aruncus
Arge nigripesGB
28
97
Arge ustulata
64
Arge cyraGB
42
Arge mali
52 Spinarge metallica

55 Spinarge fulvicornisP

87
Spinarge fulvicornisA
62
Spinarge affinis
Tree scale: 1

COI 100
Runaria reductaGB
Blasticotoma filicetiGB
Athalia sppGB
Waldheimia amazonica
56
Andeana farcta
17
21 Siobla sppGB
44
Euura respondens
89

13
Cladius pectinicornisGB
Neodiprion sertiferGB
31
Monoctenus juniperiGB
7
Abia sppGB
25
Corynis crassicornisGB
41
Heptamelus dahlbomiGB
Zenarge turneri
Heteroperreyia hubrichiGB
46
Styracotechys dicelysmaGB
14 12
Aulacomerus atriventrisGB
24
Acordulecera spGB
34
64
Acordulecera sp
Pergagrapta sppGB
97
Pseudoperga sppGB
25
Pterygophorus insignisGB
36
Perreyia tropica
54
Polyclonus atratusGB
Dracogymnia fernandezi
31 Sericoceros nrtannuusGB
64
Atomacera debilisGB
61
Atomacera pubicornis
Aproceros leucopodaGB
33 Aprosthema tardum
95
Aprosthema austriacumGB
14 100
69
3 Aprosthema intermedium
Sterictiphora longicornisGB
14 Sterictiphora geminata
73
Sterictiphora furcataGB
99
Sterictiphora angelicae
Themos mayiGB
11
40
Dielocerus sp
99
Dielocerus larva
Neurogymnia nigricosta
4
Sericoceros vumirusGB
32

1
Sphacophilus sp1
2 Durgoa mattogrossensisGB
23
Trochophora lobataGB
63
Didymia jonesiGB
Manaos mulsus
Ptenos texanus
13
4
31
Ptenos spGB
100
Ptenos leucopodaGB
7 Ptilia versicolorGB
100
Ptilia versicolor
100

4
Ptilia peleterii
Sterictiphorinae males
13
Schizocerella pilicornisGB
100
Schizocerella lineataGB
Antargidium atricepsGB
93
Antargidium dentivalveGB
99
Antargidium allucenteGB
33
Scobina incultaGB
88 Scobina bolivariGB
96 Scobina terminalisGB
88
Scobina thoracicaGB
48
Scobina strophosaGB
44
Pampsilota dahomeyanus
83
Triarge sp
88
Arge stuhlmanniGB
Cibdela janthinaGB
Arge aruncus
96 13
Arge similislarva
100
Arge similis
Spinarge metallica
17 19 Spinarge affinis
67
75 Spinarge affinisGB
67
Arge naokoaeGB
67
4 Spinarge fulvicornisGB
Arge enodis
39
Arge berberidis
16
12
Arge melanochra
55 Arge nigripes
100
Arge ustulata
95
Arge pullataGB
Arge scita
Athermantus imperialis
12
Arge meliosmae
8 9
Arge thoracica
37
Arge quidia
95
6 Arge pectoralis
Arge captivalarva
Arge cyanocrocea
1
Arge rustica
6 17
Arge rejecta
100

9
Arge captiva
Kokujewia ectrapelaGB
16
Arge bipunctataGB
98
Arge ochropus
Tree scale: 0.1 100
Runaria reductaGB
Blasticotoma filicetiGB

GLN
Athalia sppGB
51
Siobla sppGB
59
Andeana farcta
66
Corynis crassicornisGB
76 Abia sppGB
36
Neodiprion sertiferGB
57
Monoctenus juniperiGB
Phylacteophaga froggattiGB
66
Pseudoperga spGB
100
Pergagrapta sppGB
37
Pterygophorus insignisGB
32 Polyclonus atratusGB
86
Perreyia tropica
80
39 Decameria spGB
Zenarge turneri
Aproceros leucopodaGB
100
Aproceros leucopoda
65
Sterictiphora geminata
96
Sterictiphora spGB
98
59
21
Sterictiphora angelicae
Dielocerus sp
83
Sphacophilus sp1
54
Dracogymnia fernandezi
20
Schizocerella pilicornisGB
46
Atomacera pubicornis
99
Atomacera debilisGB
16 Scobina terminalisGB
99
Scobina terminalis
71
Arge scita
Arge ochropus
100
Arge cyanocrocea
12 Arge suzuki
44
37 Arge obesa
25
Arge rejecta
89
8 Arge captiva
Arge rustica
Arge pagana
2
40
Arge berberidisGB
100
Arge berberidis
2 Cibdela janthinaGB
Arge cyraGB
48
4
Arge nigripes
46
Arge spGB
52
Arge ustulata
38
Arge mali
24
Arge metallica
Spinarge prunivora
42
54
Spinarge fulvicornisA
95
Spinarge fulvicornisP
48
Spinarge pumilla
43
Spinarge affinis
Tree scale: 0.1
Runaria reducta
100
Blasticotoma filiceti
Athalia circularis
80 Siobla ruficornis
62
Corynis crassicornis
56
Abia lonicerae
Acordulecera sp.
Zenarge turneri
55 Sterictiphora sp.
33
Aproceros leucopoda
25
24
Atomacera debilis
13 Scobina terminalis
80
Arge cyra
IDH 28
100
Arge berberidis
Lophyrotoma analis
99
Pterygophorus insignis
57 Phylacteophaga froggatti
42
Decameria sp.
73
Polyclonus atratus
30
Pergagrapta sp.
100
Pseudoperga sp.
Tree scale: 0.1

Runaria reductaGB
PGD 100
Blasticotoma filicetiGB
Corynis crassicornisGB
82
Athalia sppGB
Siobla sppGB
63
49
Euura respondens
54

31
Andeana farcta
Abia sppGB
46
Neodiprion sertiferGB
28
Monoctenus juniperiGB
Scobina terminalisGB
Cibdela janthinaGB
80
Arge scita
100 21
Arge captiva
46
Arge suzuki
51
Arge berberidisGB
60 Spinarge fulvicornisP
95
Arge nigripes
96
Arge spGB
Zenarge turneri
90
Phylacteophaga froggattiGB
32
46 Acordulecera sp
91
Acordulecera spGB
86
Anathulea sp
69
Pergagrapta sppGB
100
Pseudoperga spGB
50
Pterygophorus insignisGB
47 Polyclonus atratusGB
26
76 Decameria spGB
56
Perreyia piceaGB
100
Perreyia tropica
Atomacera pubicornis
100
Atomacera debilisGB
Sterictiphora spGB
99
Sterictiphora angelicae
29 68 Aprosthema fusicorne
100
Aprosthema tardum
68
Aproceros leucopodaGB
100
Aproceros leucopoda
72
Sphacophilus sp
88 Schizocerella pilicornisGB
74
Ptilia versicolor
100
59 Ptilia peleterii
Sphacophilus sp1
67
Dielocerus larva
46
Manaos mulsus
93
Ptenos leucopodaGB
94
Ptenos texanus
92
Ptenos spGB
Tree scale: 1

TPI

Runaria reducta
98
Blasticotoma filiceti
Pterygophorus insignis
Pseudoperga sp
44
Acordulecera sp
Pergagrapta sp
82
82
Phylacteophaga froggatti
Decameria sp
97
27 Polyclonus atratus
Zenarge turneri
16
Athalia circularis
31
Euura humeralis
100
Cladius pectinicornis
35 71
Corynis crassicornis
92
Abia lonicerae
33
Siobla ruficornis
32
Neodiprion sp
24 81
Monoctenus juniperi
Dielocerus sp
99
Schizocerella pilicornis
57
Aproceros leucopoda
Atomacera debilis
52 46 Sphacophilus sp1
66
Sterictiphora sp
100
28 Sterictiphora angelicae
Scobina terminalis
100 Cibdela janthina
100
Arge cyra
83
Arge berberidis
File S1. Dispersal matrix, Dispersal Extinction Cladogenesis (DEC) analysis.

1 0.1 0.1 1 0.1 0.1 0.1

0.1 1 0.1 0.1 0.1 0.1 0.1
0.1 0.1 1 0.1 0.1 0.1 0.1
1 0.1 0.1 1 1 1 1
0.1 0.1 0.1 1 1 1 1
0.1 0.1 0.1 1 1 1 1
0.1 0.1 0.1 1 1 1 1
[0-35Ma Oligocene-Miocene]

1 1 0.1 0.1 0.1 0.1 0.1

1 1 0.1 0.1 0.1 0.1 0.1
0.1 0.1 1 0.1 0.1 0.1 0.1
0.1 0.1 0.1 1 1 1 1
0.1 0.1 0.1 1 1 1 1
0.1 0.1 0.1 1 1 1 1
0.1 0.1 0.1 1 1 1 1
[35-50 Ma Eocene]

1 1 0.1 0.1 0.1 0.1 0.1

1 1 0.1 0.1 0.1 0.1 0.1
0.1 0.1 1 0.1 0.1 0.1 0.1
0.1 0.1 0.1 1 0.1 0.1 0.1
0.1 0.1 0.1 0.1 1 1 1
0.1 0.1 0.1 0.1 1 1 1
0.1 0.1 0.1 0.1 1 1 1
[50-120 Ma Late Cretaceous]

1 1 1 1 0.1 0.1 0.1

1 1 1 0.1 0.1 0.1 0.1
1 1 1 0.1 0.1 0.1 0.1
1 0.1 0.1 1 0.1 0.1 0.1
0.1 0.1 0.1 0.1 1 1 1
0.1 0.1 0.1 0.1 1 1 1
0.1 0.1 0.1 0.1 1 1 1
[120-145 Ma Early Cretaceous]

1 1 1 1 0.1 0.1 0.1

1 1 1 1 0.1 0.1 0.1
1 1 1 1 0.1 0.1 0.1
1 1 1 1 0.1 0.1 0.1
0.1 0.1 0.1 0.1 1 1 1
0.1 0.1 0.1 0.1 1 1 1
0.1 0.1 0.1 0.1 1 1 1
[145-160 Ma Late Jurassic]

1 1 1 1 1 1 1
1 1 1 1 1 1 1
1 1 1 1 1 1 1
1 1 1 1 1 1 1
1 1 1 1 1 1 1
1 1 1 1 1 1 1
1 1 1 1 1 1 1
[160-200 Middle Jurassic-Triassic]
#periods Myr=35.0 50 120 145 160 219
File S2. Total Evidence Matrices for Node dating.[syen12527-sup-0006-AppendixS2]
#NEXUS
[written Wed Nov 18 16:22:02 COT 2020 by Mesquite version 3.6 (build 917) at Leonardos-MacBook-Pro.local/192.168.0.12]

begin data;
dimensions ntax=164 nchar=5658;
format datatype = MIXED(DNA:1-5350,Standard:5351-5658) gap = - missing =? interleave=yes;

MATRIX
Blasticotoma_filiceti ??????????????????????????????????????????????????????CTGGTCTCAT-TTATATTA-ATGATTGAATTTAATTTTTGAGTGAAAAAGCTTAAATTTAATTAAGGGACGAGAAGACCCTATAGAGTTTTATA-A---TT-G-T-----A--GTGTT-TATGTATTTAA-TT--
ATTT-ATTA---A-ATA-A-C-TATATTTATTTGGTTGGGGTGATTGGAAAATTTATTAAACTTTTTTAA-TG-G-G-------CAAA-CATTGATTTATGAT-TTT----ATTGATCCTTAT--T--T-ATAAGATTGTTAGATTAAATTACCTTAGGG-----------------------------------------------
A???????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????
????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????
?????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????
ATCCTTATTCGATTAGAATTAAGAACTCCTAACAGTCTTCTAAGTAATGACCAAACATACAACACTCTAGTAACCTCTCATGCCTTCTTAATAATTTTCTTCATAGTTATACCAATCATAATTGGAGGGTTCGGAAATTGACTAATCCCCCTCATAATTTCAGCTCCTGATATAGCTTTTCCTCGTATAAACAACTCTAGATTCTGACTTCTACCCCCTTCTCT
AATTCTACTAACTTCAAGAAACTTTATTGACTTAGGAACAGGAACAGGATGAACTATCTACCCTCCTCTATCAAGAAACTTAGGTCATGCAAGATCTTCAATAGACCTAACTATTTATTCCCTACATATAGCAGGTATTTCCTCAATTATAGGAGCTATTAATTTTATCTCTACAACAATTAACATACGAAATAAAGGGATATCCTCTGAACGACTAACCTTAT
TCACTTGATCAGTATCAATCACAGCACTACTTTTAATATTATCATTACCCGTTCTAGCAGGGGCTATCACAATATTATTAACAGACCGAAACTTAAATACATCATTTTTTGACCCAGCTGGAGGAGGAGACCCCATCCTATTCCAACACCTATTT?????????????????????????????????????????????????????????????????????
????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????
???????????????????????????????????????????????????????????????????????????????????????????????????????????????????????
GATTCCATAAAAGAAAAGCTGATCCTGCCCTTCCTCGACGTCGAGCTGCACACCTACGACCTCGGCATGGAGAACCGCGACGCGACCCAGGACAAAGTAACGGTGGACTGCGCGGAGGCGATAAAGAAGTACCACGTGGGGATAAAATGCGCGACGATTACGCCGGACGAGAAGAGAGTAGAGGAGTTTAACCTGAAGCAGATGTGGAAGAGTCCCAACGGCAC
CATCCGGAATATCCTGGGCGGAACGGTGTTCCGCGAGGCGATAATCTGCAAGAACATTCCGCGCTTGGTCTCCGGCTGGAACCTGCCCATCATCATCGGTCGGCACGCCCACGCCGATCAGTACAAGGCGACCGACTTCGTCGTCCCCGGCGCCGGAAAGCTCGAGATCACTTGGACGGGCGCTTCGGGAGACAAGATCCATCATACCGTGCACGAATTCAAGG
GAGCGGGCGTCGCGCAGGCTCAGTTCAACACCGACGACAGCATCCAGGCGTTTGCTCACAGCTCGATGCAGTACGCGCTGTCGCGAAGCTATCCGTTGTACTTGTCGACCAAGAACACTATCTTGAAGAAATACGACGGCAGGTTCAAGGACATCTTCCAGGAGATCTACGAGAAGGATTACCGGCCCAAGTTCGAGGCCAAGAAGATACGGTACGAGCATCGC
CTGATCGACGATATGGTCGCCTACGCGATGAAATCCGAAGACATGCAATTAATTTGCGAAGCCTATCACATCATGCGAAACGGTTTGGGACTTAATCCTAAGGAGATGAGCGATGTTTTCGACGAATGGAACAAGGGCGTACTCGATTCCTTTTTAATTGAAATTACCAGAGATATTTTGAAGTACAAGGATGAAAAAGGGTATCTCCTTGAGCGAATCAGGGA
TACGGCTGGGCAGAAGGGTACGGGAAAATGGACAGCGATCGCCGCTTTAGATTACGGAATTCCCGTAACTCTAATCGGAGAATCGGTATTCGCCAGGTGTCTTTCGTCGCTGCAGAGCGAAAGAGTAGAGGCCAGTGTCGTTCTGGCTGGACCGCACGCAGTGGACGGACCGCACGCAACTTACCAGGGCAATAAGAAACAGTTCGTCGAACATCTCGGGAAAG
CGCTCTACGCTTCCAAAATCATTTCCTACGCCCAAGGATTTATGTTGCTAAGAGAAGCTGCAAAGATTCATAATTGGAATTTAAATTACGGCGGCATTGCGCTCAGCAAGAAGGAAATTGACGAAATCGTCGCGTTTCTAAAAGCCGGCCCGCTCGATCCCAGCGTCGGTAAGTTTTCAGAGGTCGTGGTCGGAGTACCCGCTATATACTTGACTTACGCGAAA
AGTATTTTACCCAGCAATGTACAAGTCAGCGGTCAGAATACCTACAAAGTAGCGAAGGGGGCTTTCACCGGCGAGCTCAGTCCTGCCATGCTTTTGGACAACGACGTCCCGTGGGTAATCCTAGGGCACTCCGAGCGCCGCAACGTCTTCGGCGAATCGGACGACTTGATTTCCGAGAAAACAAGCCACGCTCTCGAGGCTGGGCTAAAGGTAGTCATCGCTTG
CATCGGCGAGAAGCTGGACGAGCGCGAGGCTGGAAAGACAGAAGAGGTCGTCTTTAGGCAGACCAAAGCTATCGCGGACAAAGTTAAATCCTGGGACAACGTCGTACTG???????????????????????????????????????????????????????????????????????????????????????????????????????????????????
?????????????????
ATCGACGACGAGGAGCTCGAGAAGCCGACCGACAAGCGAACGTTCGTTCTGGCGGCCGCCCTGAAGTCCGGCTACACGGTGGACCGTCTGTACGAGCTGACGAAGATCGATCGTTGGTTCCTGGAGAAGATGAGGAACATCACCTCGTATTACAGCATCCTCGAGGCCCTGGACCAGACGAAGCTCTCCCACGAGGTTCTCCTGCGGGCGAAGCAAATCGGCTT
CTCGGACAAGCAGATCGCCAAGGCGGTCAAGAGCACCGAACTGGCCGTGCGGAAGCAGCGACAGGAGAGCAACATCCGCCCTATCGTCAAACAGATCGACACCGTCGCGGCCGAGTGGCCGGCGACCACAAACTACCTCTACCTCACCTACAACGGAATCTCCCACGACATCCTCTTCCCCGGCGGCTACACAATGGTCATCGGTAAGGCAGGCTCCGGCGTCT
ACCGCATCGGAAGTTCCGTGGAGTTCGACTGGTGCGCCGTGGGTTGTCTTCGCGAGTTACGCCGCCTGGGGCGCAAGACGATCATGGTCAACTACAACCCGGAGACGGTCAGCACCGACTACGACATGTGCGATCGCCTCTACTTCGAGGAGATCTCCTTCGAGGTCGTCATGGATATTTACGACCTCGAGAACCCGGAGGGC?????????????????????
?????????????????????????
GCAGTTTACAGAAAAACTTCAAATCCTTATTCGGCGAGAAACTCGAGGTTGTTAGAACGCACCAGCAACAAGAAAACCTCAAATTCATGGCGCATTTCAAGCGTAAGTTTATTATTCATCAAGGAAGACGCAAGCAGCCAAAGTCACCCGCCACTAACAAAGTCGAATTTTACCATCTTCGAAGCAACGGTAGCGCCTTGTGTACTAGGCTGATACAAGTGCAG
CCGGACGCCCTTTTATTGAATTCCGCGTTTTGGTACAGCTACATTTTAAACGTACCTTTTAATAACGACGACGAGACCGGAATTGTCTACGTTTGGGTTGGATCGAAAGCTGACAGCGAAGAGGCGAGGCTCGTCGAAGAAATGGCAGAAGAAATGTTCAACAATGTACCGTGGATAAGTCTACAAGTATTAAACGAGGGCGAGGAACCAGACAACTTTTTCTG
GGTCGGACTTGGCGGTAAAAAACCGTACGACACCGTGGCCGAGTACATGAACTACACCAGGCTATTTCGATGCTCCAACGAAAAAGGATACTTCACGATTAGCGAGAAGTGCACAGATTTCTGCCAGGTAGACGATCTGGCCGACGACGATATAATGATACTGGACAACGGCGAACAAGTGTTCCTGTGGCTGGGTGCAAGATGTTCGGAAGTCGAGATTAAGC
TTGCGTACAAATCGGCTCAAGTACGGGTGTATATACAACACTTACGAGCGAAGCAACCAGAAAATCCCCGCAAGTTGTTCCTTACGGCTAAAAGCAAGGAATCTCGACGATTCACA
Abia_spp ATAATAATTAGTCTTTTAATTGGGGGCTGGAATGAAGGATTAGACAAAATAAAAACTGTCTTTA-GTTAATT-TT-AATTGAATTTAATTTTTAAGTTAAAAAGCTTAAATTTTATTAAAGGACGAGAAGACCCTATAGAGTTTTATATT---TA-T--TAT--AATTTAGA-----TTTTTAAT-T--
TA-TT-TTTAA------ATTATTATAAATATTTTATTGGGGTGATAGATAAATTTAATTAACTTTTTTAT-TT-A-T-----A-TTTA-CATTAATTAATGAA-A-T----TTTGATCCTAATTTT--T-T-TAGATTATTAGATTAAATTACCTTAGGGATAACAGCATAATTTTTTTTAAAAGTTCTTATTGATTAAAAAGATTGC????
TCTAAGTGGGTGGTAAACTCCATCTAAGGCTAAATATGACCACGAGACCGATAGCGAACAAGTACCGTGAGGGAAAGTTGAAAAGAACTTTGAAGAGAGAGTTCAAGAGTACGTGAAACCGTTCAGGGGTAAACCTGAGAAACCCGAAAGTTCGA-ACGGGGAGATTCACCGTCAGCGAGGCCGGCTTCGC-CGCGG-TTCGCGATG--C-CGCC-GGTA-
CGT-TCGC----GTACCGCGCGGC-ACGCGTCCGTGG-TGCA----ACGCTCGACAGCGCCGTCGTGCACTTCTCCCCAAGTAGGACGTCGCGACCCGTTGGGTGTCGGTCTAAGGCCCGGTG----GGAGGCCCGTCG--ACGGCGT--TCGC--GC-CGTT--TCGGCGGACC-C-C-CGGT-TGCCCGTCCGGCTGCCCGGCGGTACACGCAC-
GGTATAGAGCCTCATTT-GAA-CTGAGTCGGGCCCGTCGCAAGCCTGGTCAGAGTT----TCCCGGTGGTCGGAC-CTAGCGCCGTCCCCGGGCCTGGCCGGCTGTTGGCAGGCGGTGTCCTCCGACAGGC---------------------------------------------------------------------
TCATAATTATTCGAACTGAATTAGGAACTTCAGGATCATTAATTGGTGATGATCAAATTTATAATGTAGTAGTAACATCACATGCCTTTTTAATAATTTTCTTTATAGTAATACCTATTATAATTGGAGGATTTGGAAACTGACTAGTTCCTTTAATATTAGGAGCCCCTGATATAGCTTTTCCTCGATTAAATAATATAAGATTTTGATTATTACCCCCTTCA
ATTATCTTATTATTATCAAGAAGAATTGTAAATTCAGGTAGAGGAACTGGATGAACAGTATATCCTCCTTTATCCTGA---
ATAGGGCATATAGGGGCCTCAGTAGATTTAACTATCTTCTCACTACATTTAGCAGGAGTTTCTTCTATTCTAGGAGCTGTAAATTTTATTTCTACAACTATTAATATAAAAATTAAAAATATAAAATATGATCAACTTTCACTATTTATTTGAGCTACAATCTTAACTACTATTCTATTATTATTATCACTCCCTGTATTAGCAGGAGCAATTACTATACTATT
AACGGATCGTAATTTAAATACATCATTTTTTGACCCTGCAGGGGGAGGAGACCCTATTTTATATCAACACTTATTTTGATTTTTTGGTCATCCAGAAGTTTATATTCTAATTTTACCAGGATTTGGAGTAATCTCTCATATTATTACACAAGAATCAGGTAAAAAAGAAACCTTTGGAGCCTTAAGAATAATTTATGCAATATCTGCTATTGGATTATTAGGAT
TTGTAGTCTGAGCTCACCATATATTTACTATTGGTATAGATATTGATACACGAGCATACTTTACAGCAGTAACTATAATTATTGCAATCCCTACAGGAATTAAAATTTTTAGTTGAGTAGCCACATTACATGGATCTCAAATTAAGTTTACTCCTTCAATATTATGAGCCTTAGGATTTATATTTTTATTTACAAGAGGGGGTTTAACAGGTATTATATTAGCA
AATTCATCTATTGATATTATTTTACATGATACTTATTATGGACTCCATCAAGGAGAAACTCATTCTTCCATTCCTCGATATCGAGCTCCACATCTACGATCTTGGTATGGAGAATCGAGACGCAACTAATGACCAGGTAACCGTGGACTGCGCCGAAGCGGTAAAGAAGTATAACGTAGGGATAAAATGCGCAACAATAACACCGGATGAGAAGCGAGTCGAAG
AATTCAAATTAAAGGAGATGTGGAAGAGTCCCAACGGTACGATAAGAAACATATTGGGCGGCACTGTTTTCAGAGAGGCGATAATTTGTAAAAATATTCCGCGTTTGGTGACTGGATGGAATCAGCCGATAATAATCGGTCGTCACGCTCATGCCGATCAGTACAAAGCTACAGATTTTGTCGTTCCTGGTGCTGGTAAGCTGGAGATCACCTGGACAGGTGAA
AAGGGGGAAAAGATTCAGCATACCGTCTATCAGTTCAAGGGAGCTGGCGTCGCCCAGGCTCAGTTCAATACTGACGAAAGTATCGAGGCCTTTGCCCACAGCTCTTTCCAGTATGCTTTGTCACGGGCTTACCCTCTCTACCTCTCAACCAAGAATACAATTTTAAAGAAGTACGACGGTAGATTCAAGGACATTTTTCAAAAAATTTATGACTCTGAGTATAA
GTCTAAATTTGAGGCAAAGAATATTTGGTACGAGCACCGTCTTATCGACGATATGGTAGCCTATGCCATGAAATCTGAGGACATGCAGTTGATCTGCGAAGCGTATCACGTAATGAGAAACGGATTGGGTTTGAGCCCGCAGCAGATGAGCGACGTTTTCGGCGAGTGGAACAAGGGCGTACTAGATTCATTCCTTATTGAAATAACGCGAGATATATTGAAGT
ACAAGGACGATAAGGGATACCTTTTGGAAAGAATCAAAGACTCTGCGGGACAGAAGGGGACCGGGAAGTGGACCGCGATCGCCGCCCTCGACTACGGAGTTCCCGTAACTCTAATCGGCGAGTCGGTATTCGCTAGGTGCCTCTCGGCTCTCCAGAGTGAGAGAATAGAGGCCAGTGCGGTTCTCGA------------------
AGGGCCGAACACGGTCTACCAAGGCGACAAGACACAGTTCCTCGAACATTTGAGGAAGGCTCTCTATGCCTCGAAGATAATTTCGTATGCTCAAGGGTTCATGCTGTTGAGAGAAGCGGCAAAAGTTCACAACTGGAATTTGAACTATGGTGGAATTGCGCTCGTCAAGAAAGAGATCGATGAAATCGTTGCATTTCTGAAAACCGGTCCGCTCGATCCAAATG
TCGGTACGTTTTCAGAGGTCGTTGTCGGCGTACCTGCCCTCTATCTGTCCTATGTCAAGAGTATACTACCAGCCAATGTACAAGTCAGCGCTCAAAATTCCTACAAAGTAGCTAAGGGTGCTTTTACTGGTGAAATAAGCCCCGCTATGCTCGTTGACAGCGGAATTCCTTGGGTGATTCTTGGACACTCGGAGCGTCGTAACGTATTCGGTGAATCGGATGAA
CTTATTTCTGAGAAGATAGCTCACGCCCTTGATACTGGCGTCAAGGTAGTAATTGCTTGCATCGGAGAGAAGCTCGACGAACGCGAAGCTGGAAAGACAGAGGAAGTTGTATTCAAGCAGACCAAAGCTATTGCGGACAAAATCAAGTCCTGGGATAATGTCGTACTG????????????????????????????????
CGAGGCGATGGCGATCGGTAGGAAGTTCGAAGAGGCTTTTCAGAAGGCTCTTCGCATGGTTGACGAGAATGTGAACGGTTTTGATCCTTACATAAAACCTATCGATGACGAGGACCTCGAGAGGCCAACCGACAAAAGAATGTTCGTACTCGCAGCAGCCCTCAAAGCCGGCTACACGATCGATCGCTTATACGAATTGACAAAAATCGATCGCTGGTTCCTGG
AAAAAATGAGGAACATTACTTCATACTATACGCTTCTCGAAGATCTGGATCAGATGAAACTCTCGTACGAAGTCCTGTTGCGTGCCAAGCAGATCGGTTTCTCCGAYAAGCAGATAGCAACCGCAGTAAAAAGCACTGAGCTTGCTGTCCGCAAACAAAGACAGGAAAGCAACATCACACCTTTTGTCAAACAGATAGATACGGTCGCCGCCGAGTGGCCAGCT
ACCACAAACTACCTGTATCTCACCTACAATGGAAAAAGTCACGACATTCAATTCCCTGGCGGGTACACGATGGTCATCGGTATGCCAGGTTCCGGAGTTTACCGCATAGGCAGTTCCGTTGAGTTTGATTGGTGTGCTGTGGGGTGCCTCAGAGAGTTGCGTCAACTCAACAGGAAGACGATAATGGTGAACTACAACCCCGAGACAGTGAGCACGGATTACGA
TATGTGCGATCGCTTATACTTTGAGGAAATCTCCTTCGAGGTCGTGATGGACATTTACGACCTCGAGAATCCGGAGGGG??????????????????????????????????????????????
TCAGCTTACAAAAAAAATTCAAATCTTTATTTGGTGAAAAACTGGAGGTTGTCAGGACACATCAACAACAAGAAAACCTGAAGTTTATGTCGCATTTTAAACGCAAGTTTATCATTCGTCAAGGAAGGCGAAAAGACGCAAAAACACCAGCTAATAATAAAGTTGAGTTTTATCATCTCCGAAGCAATGGCAGCGCCTTGTGTACCAGGTTGATACAAGTGAAT
CCGGATGCGATTTTATTAAATTCTGCGTTYTGGCACAGCTACATATTAAATGTGCCATTCAATAACGACGATGAAACTGGTATTGTTTACGTTTGGATTGGGTCAAAGGCCGATAGTGAAGAAGCCAGACTCGTGGAAGAGATAGCAGAGGAGATGTTCAATAACGTTCCATGGATAAGTCTGCAAGTTCTAAACGAGGGTGAAGAACCAGATAACTTTTTCTG
GGTCGGTATTGGAGGAAAAAAACCATACGATACGAATGCGGAATACATGAACTACACAAGACTATTTCGGTGTTCAAATGAAAAAGGGTATTTCACGATCAGCGAGAAATGTACAGATTTTTGTCAGGTGGATGACTTGGCTGACGATGATATTATGATATTAGACAACGGCGAGCAGGTGTTCCTCTGGCTAGGTGCCCGATGTTCAGAAGTTGAGATTAAGC
TTGCCTACAAATCGGCTCAGGTGAGTGTGTACATACAACACTTGCGCGTTAAACAGCCAGAAAGACCTAGGAAACTATTCCTCACTGCCGCAAGCAGCGAGTCTAGAAGATTCTCA
Andeana_farcta ATAATAATTTGTCTTTTTATTGAAGACTAGAATGAAAGATTTGACGAGATAAAAACTGTCTTT-TTTTAATTT-TTTTTTGAATTTAATTTTTAAGTTAAAAAGCTTAAATTATTTTAAAGGACGAGAAGCCCCTATAGAGTTTTATA----ATT-A-TATT--TTTAAAAA-A--AAATATAA-ATT--
T-AA-TTTTT-TAAAT-A-A-ATTAATTATTTTATTGGGGTGATAAAAAAATTTAAAAAACTTTTTTAT-T-TT-A-A------AAA-CAATAATTTTTGAA-T-T----TTTGATCTTAAA--T--T-C-TAGATTAATAGAATA------------------------------------------------------------
ACAGCTCTAAGTGGGTGGTAAACTCCATCTAAGGCTAAATATGACCACGAGACCGATAGCGAACAAGTACCGTGAGGGAAAGTTGAAAAGAACTTTGAAGAGAGAGTTCAAGAGTACGTGAAACCGTTCAGGGGTAAACCTGAGAAACCCGAAAGATCGA-ACGGGGAGATTCATCGCAAGCGACGCTGGCTTCTC-CGTGG-CTCGTGATG---CCG-G-G--
A-C-T-T--C----GCGT-CCACGGC-ACTCGGCCGCGG-TGTT----ACGTCCGGCGGCGCCGGCGTGCACTTCTCCCCCAGTAGGACGTCGCGACCCGTTGGGTGTCGGTCTACGGCCCGGTG----GGACGACTGT-C--CCGGCGC--TCGC--GT-C-----GGGGCAGACC-C-C-CGGT-TGCCCGTCCGACCGCCCGGCGGTACACGCAC-
GGTATAGAGCCGCAT-T-TTA-CTGCGTCGGGCCCGTCGCAAGCGCGGTCAGCGAT----TCCCGGTGGTCGGAC-CTAGTGCCGTCCCCGGGCCTGGCCAGCTGTTGGCAGGCGGTGTCCTCTGACTGGCTC-ATT---T-GAAA-C-----AATAT-
TATACCGGTCGGCGACGCTATTGCTTTGGGTACTTTCAGGACC?????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????
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TTAAATAATATAAGATTTTGATTTTTACCCCCCTCTTTAATTCTTTTAATCACTAGAAGATTAGTTAGTTCAGGATCAGGAACAGGATGAACAGTTTATCCTCCTTTATCAAATAGATTTTCTCATTCAGGACCTTCAGTAGATATAACTATTTTTTCTCTTCATTTAGCAGGAATTTCATCTATTCTTGGAGCAATCAATTTCATTTCAACAATATTAAATAT
ACGAATTAAAATAATAAAATATGAACAAATACCTTTACTTATTTGAGCTGTTTCATTAACAGCTCTTTTATTATTATTATCATTACCTGTATTAGCAGGTGCAATTACTATATTATTAACAGATCGAAATTTAAATACTTCATTTTTTGATCCTTCAGGAGGAGGAGATCCAATTCTATATCAACATTTATTTTGATTTTTTGGACACCCAGAAGTATATATTT
TAATTATTCCTGCTTTTGGTATAATTTCTCATATTATTTCCCAAGAATCAGGAAAAAAAGAAACATTTGGAACTTTAGGAATAATTTATGCAATATTAACAATTGGATTACTTGGATTTATTGTTTGAGCTCACCATATATTTACTATTGGAATAGATGTAGATACTCGAGCATATTTTACATCTGCAACTATAATTATTGCAGTCCCTACTGGAATTAAAATT
TTTAGATGATTAGCAACAATTCATGGATCAAAATTAATTTTTAACCCATCAATAATATGAACATTAGGATTTGTATTTTTATTTACAATTGGAGGATTAACAGGTATTATATTATCAAATTCATCAATTGATATTATTTTACATGATACTTATTATG???????????????????????????????????????????????????????????????????
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GACATGCAGTTGATCTGCGAGGCTTATCACATAATGAGAAATGGTTTGGGATTAAGTCCGCAAGAAATGAGCGATGTTTTCGGAGAATGGAATAAAGGGGTGTTGGATTCATTTCTTATCGAAATAACCAGGGACATATTGAAGTTCAAAGATCACAAGGGATATCTTTTAGAGAGAATCAGGGATACTGCTGGTCAAAAAGGAACTGGAAAGTGGACTGCGAT
ATCTGCCTTAGATTACGGTATTCCTGTTACTCTAATCGGGGAATCAGTATTCGCCAGATGTCTGTCGTCCCTGCAAAACGAGAGGATAGAAGCCAGTACCGTGTTAGA------------------
AGGACCGAATGCTCTCTATCAAGGAGACAAGAAGCAGTTTCTCGAACATCTTAGAAAAGCTCTCTATGCTTCGAAAATCATTTCATACGCTCAGGGTTTCATGTCACTGAGGGAAGCA??????????????????????????????????????????????????????????????????????????????????????????????????????????
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ACGTTACCGGTTTTGATCCTTACGTAAAACCCATAGACGACGAGGATCTAGAAAGGCCAACTGACAAACGAATGTTCGTACTAGCGGCTGCCCTAAAATCTGGCTACACGATCGATCGATTGTACGAATTGACGAAAATTGATCGATGGTTTTTAGAAAAAATGCGTAACATCATCGCGTTTTATACCCTTCTCGAAGATCTTGATCAAACGAAATTATCCCAC
GATGTTCTTCTGCGTGCCAAACAAATCGGTTTCTCCGATAAACAAATCGCCAAAGCTTTCAAGAGCACCGAACTCGCTGTTCGCAAACAAAGACAAGAGAGTAACATCAGACCATTCGTAAAACAAGTCGACACCGTTGCTGCCGAGTGGCCGGCAACCACTAATTACCTTTATCTTACCTACAACGGAAACAGTCACGATCTTCCATTCCCCGGTGGTTACAC
CATGGTTATTGGTTCGTCAGGTTCCGGCGTCTATCGCATCGGAAGTTCCGTGGAGTTCGATTGGTGCGCGGTTGGATGTTTAAGAGAATTACGACGATTAAATCGTAAGGCGATCATGGTGAATTATAATCCTGAAACGGTAAGCACGGACTACGACATGTGCGATCGATTGTACTTCGAGGAAATATCATTCG??????????????????????????????
?????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????
CATTTCAAGCGTAAGTTCATAATTCGGCATGGAAGACGCAAGCAACCGAAATCACCTGCCAATAACAAAGTAGAATTTTATCATCTTCGGAGCAATGGTAGCGCTTTATGTACCAGATTGATTCAAGTGAATCCTGACGCCTGTTTATTGAACTCTGCTTTTTGGTTCAGCTATATTTTGAACGTACCATTTAACAATGACGATGAAACTGGTATTGTCTACGT
TTGGATTGGATCTAAAGCAGACAGTGAAGAAGCGAGACTTACAGAAGAGATAGCGGAGGAAATGTTCAATAATGTTCCATGGATCAGTTTGCAAGTATTAAACGAGGGCGAAGAACCAGACAATTTCTTCTGGGTTGGTATCGGAGGAAAAAAACCGTATGATACGGACGCCGAATACATGAATTTCACAAGACTATTTAGGTGCTCTAACGAAAAGGGGTATT
TTACAATTAGTGAAAAGTGTACTGATTTTTGCCAGGTAGATGACTTAGCCGATGACGATATCATGGTGCTAGACAACGGAGAGCAAGTATTTCTATGGCTTGGTGCACGGTGTTCAGAAGTAGAAATCAAACTTGCTTATAAATCAGCGCAGGTGAAA??????????????????????????????????????????????????????????????????
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Athalia_spp ATAATAATTTGTCTTTTAATTGAAGGCTGGTATGAATGATTTGATGAAGTAAAAACTGTCTCT-ATTAAATTA-TTAATTTAATTTTATTTTTTAGTTAAAAAGCTAAAATATTATTAAAGGACGAGAAGACCCTATAGAGTTTAATA-A---TA-A-A-TT--AATTATT--T--TATTTAAA-TTT--
T-AA-TAAAT-TATTT-T-T-TTTTATTATTTTGTTGGGGTGATAGGAAAATTAATTAAACTTTTTTAA-TTAT-T-T------TTAACATTGATTTATGAA-T-T----TTTGATCTTTAA--A--T-T-AAGATTATTAGAATAAATTACCTTAGGGATAACAGCGTTATTTTTTTT-
AAAGTTCTTATTTATAAAAAAGATTGC?????????????????????????????????????????????????????????????????????????????????TTGAAAAGAACTTTGAAGAGAGAGTTCAAGAGTACGTGAAACCGTTCAGGGGTAAACCTGAGAAACCCGAAAGATCGA-ACGGGGAGATTCATCGTCAGCGAGACCGGCTTCGC-
CGCGG-ATCGTGATG---TCGGG-G--A-C-C-T--C----GCGT-CCTCGGC-ACTCGTCCGCGG-TGTT--C-ATGTCCGGTGACGCCGGCGTGCACTTCTCCCCTAGTAGGACGTCGCGACCCGTTGGGTGTCGGTCTAAGGCCCGGGA----GGACGCCTGT-C--ACGTCGT--TCGC--GT-C-----GTGGCAGACC-C-C-CGGT-
TGCCCGTCCGACTGCCCGGCGGTACCCGCAC-GGTATAGAGCCGCAT-T-TGA-CTGCGTCAAGCCTGCCGCAGGCTCGGTCAGCGAT----TCCCGGTGGTCGGAC-CTAGCGCCGTCATCGGGCCTGGCCAGCTGTTGATCGGCGGTGTTCTCTGACTGGCTC-ATT---C-GAATAT-----TACAT-
TGTACCGGTCGGCGACGCTATTGCTTTGGGTACTTTCAGGACCATAATTATTCGAAGAGAATTAAGATCATCAAATTCACTAATCAATAATGATCAAATTTATAATGTAATTGTTACATCACATGCCTTCTTAATAATTTTTTTTATAGTTATACCCATTATAATTGGGGGATTTGGAAATTGACTAATTCCATTAATATTAGGGGCTCCTGATATAGCATTCC
CTCGAATAAATAATATAAGATTTTGATTTTTACCTCCCTCATTAATTTTACTTATTTCAAGAAGTTTAGTAAACACTGGATCAGGTACGGGTTGAACAGTTTACCCACCATTATCTTCAAATACTTCCCATGCAGGAGCATCAGTTGATTTAACAATTTTTTCATTACATTTAGCTGGAATTTCTTCAATTCTAGGAGCTATCAATTTTATTTCTACTATTTTA
AATATACGAATAAATGGAATAACAAATGAACGAATATCATTATTTACATGATCAGTTTTATTAACAGCCATTTTATTATTATTGTCATTACCAGTATTAGCAGGAGCAATTACAATATTATTAACTGACCGAAACCTTAATACCTCATTTTTTGACCCATCAGGAGGAGGGGACCCTATTTTGTACCAACACTTATTTTGATTTTTTGGGCACCCTGAAGTTTA
TATTTTAATTCTCCCTGCATTTGGAATAATTTCTCATATTATTTCACAAGAATCTGGGAAAAAAGAAACTTTTGGAACATTAGGGATAATTTATGCAATATCTAGAATTGGATTATTAGGATTTATTGTATGAGCACATCATATATTTACAGTAGGAATAGATGTAGACACCCGAGCGTACTTTACAGCAGCAACAATAATTATTGCTATCCCAACAGGAATTA
AAATTTTTAGATGAATAGCAACTATATATGGTAATAAAATAATCTACTCCCCCCCAATATTATGAACACTAGGATTCATTTTTCTATTTACTTTAGGGGGTCTAACAGGAATTATTTTATCTAATTCATCAATTGATATTATTTTACATGATACCTACTATGGACTCTATCAAAGAGAAACTGATCCTTCCATTCCTAGACATCGAACTACATATCTATGATCT
TGGAATGGAGAACCGAGATGCTACCAACGACCAGGTGACTGTCGACTGCGCAGAGGCCGTAAAACGGTACAATGTGGGTATAAAATGTGCAACGATAACCCCTGATGAGTTGAGAGTGAAAGAATTCAATCTCAAAGAAATGTGGAAGAGCCCCAATGGGACCATCCGAAACATTCTGGGTGGTACCGTCTTCAGAGAGGCTATAATCTGCAAAAACATTCCTC
GATTAGTCACCGGTTGGAATCTGCCCATCATAATTGGGCGTCACGCTCACGCTGACCAGTACAAAGCTACCGACTTTGTAGTACCGGGAGCCGGCAAGCTGGAAGTTACTTGGACAGGAGAATCGGGTGAAAAAATTCAGCACACCGTCTATCAGTTCAAGGGAGCAGGCATCGCTCAGGCCCAATTCAACACCGATGAGAGCATTCAAGCTTTTCCCCACAGT
TCTTTCCAGTACGCTCTTGCTAGGGCTTACCCTCTATACTTGTCGACCAAGAATACGATCCTCAAGAAATACGATGGGAGATTCAAGGACATATTCCAGGAAATCTATGAGGCGGAATACAAGTCTCAATTTGAGGCAAAAAATATCTGGTATGAGCATCGTCTCATCGATGACATGATAGCATACGCGATGAAATCCGAGGATATGCAGCTGATTAGCGAAGC
ATATCATATCATGAGAAACGGGTTGGGGCTTAATCCACAAGAAATGAGTGATGTTTTTGGTCAATGGAATAAAGGGGTGCTTGATTCTTTCCTGATTGAAATCACCAGAGATATTTTGAAGTATAAAGATGAAAAAGGATACCTTTTAGAAAGGATCAGAGATACAGCTGGGCAGAAGGGTACAGGAAAATGGACAGCCATCGCTGCTTTGGATTACGGTATTC
CCGTTACTCTAATCGGAGAATCTGTATTCTCTAGGTGCCTTTCCTCCCTACAAAGTGAGAGAATAGAAGCCAGCTCTGTCTTGGA------------------
AGGACCAAGCAGTCTCTATCAAGGTGATAAAAAGCAATTCATCGAACACCTTGGGAAAGCACTCTATGCTTCGAAGATTATCTCCTACGCTCAAGGTTTCATGCTTCTAAGAGAAGCTGCCAAGGTTCATAACTGGAATTTGAACTATGGAGGCATTGCACTTATCAAAAAGGAGATTGATGAAATCGTCGCCTTCCTAAAGGCCGGTCCACTCGATCCAAATG
TTGGTACGTTTTCAGAGGTCGTTGTTGGAGTACCTGCTCTTTACCTAACTTACGCTAAAAGCGTATTGCCACCCAATGTACAAGTCAGTGCACAGAACAGCTACAAGGTAGCAAAAGGTGCTTTCACCGGAGAAATCAGCCCTGCTATGCTCGTAGATAGTGGAATTCCATGGGTTATTTTGGGCCACTCGGAACGTCGTAACGTATTTGGCGAATCTGATGAA
TTAATTTCTGATAAAATTGCCCATGCACTTGAAGCTGGTGTGAAGGTAGTCATTGCTTGCATTGGTGAAAAACTGGACGAACGTGAAGCTGGTAAAACTGAAGAAGTAGTTTTCAAGCAGACTAAAGCTATTGCTGACAAAATCAAGTCTTGGGACAATGTTGTGCTG????????????????????????????????
CGAGGCGATGGCGATTGGTAGGAAATTCGAGGAGGCGTTCCAGAAGGCTCTTCGAATGGTGGATGAAAATGTTAACGGTTTTGACCCTTACGTAAAACCAATCGATGATGAGGACCTGGAGCGACCAACTGACAAGAGAATGTTCGTCCTTGCAGCTGCTCTAAAAGCCGGTTACTCGATCGACCGGCTTTACGAATTGACTAAAATCGATCGGTGGTTTTTAG
AGAAAATGAAAAACATCACCACTTATTATACACTTTTAGAAGATCTTGATCAAACTAAACTATCCCATGAAGTTCTTCTTCGCGCAAAGCAAATTGGATTCTCTGATAAGCAAATTGCTACCGCTGTAAAAAGCACGGAACTTGCTGTTCGCAAACAAAGACACGAAAGTAACATTGGACCATTCGTTAAACAGATT???????????????????????????
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ACAGTTTGCAGAAGAAGTTCAAATCCTTATTCGGTGAAAAATTGGAAGTAGTCAGAACCCATCAACAACAAGAAAATTTAAAATTCATGGCGCATTTCAAACGCAAATTCATTATACGCCATGGAAGACGCAAACAGCCGAAGTCACCAGCTAACAATAAAGTTGAATTTTACCACCTCCGAAGCAACGGCAGTGCCTTGTGCACAAGATTGATTCAAGTGAAT
CCGGATGCCAGTTTATTGAATTCTGCTTTTTGGTACAGCTACATATTAAATGTGCCATTCAATAATGACGACGAAACTGGTATCGTTTACGTTTGGATTGGATCCAAAGCTGACAGCGAAGAAGCCAGACTTACAGAGGAGATAGCAGAAGAAATGTTTAACAACGTACCATGGATAAGTCTGCAGGTATTAAACGAGGGTGAAGAACCAGATAACTTTTTCTG
GGTTGGAATTGGAGGGAAAAAACCATATGATACCACTGCCGAGTACATGAACTTCACTAGACTATTTCGCTGCTCAAATGAAAAAGGTTACTTCACAATCAGCGAGAAATGTACAGATTTTTGTCAGGTAGATGACTTGGCCGATGACGACATTATGGTGCTAGACAATGGTGAGCAAGTATTTCTATGGCTCGGTGCTCGATGTTCAGAAGTGGAGATTAAAC
TTGCCTACAAATCAGCGCAGGTAAGAGTTTATTTACAACACCTGCGTGTGAAGCAGCCTGAAACACCAAGAAAACTTTTTCTTACCGCAAAGGGTAAAGAGTCGCGCAGATTTACG

Cimbex_cuadrimaculata ??????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????
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Cladius_pectinicornis ATAATAATTTGTCTTTTAATTTAAGGCTGGAATGAATGATTGGTCAAGGTATAAACTGTCTTT-ATTTAATTA-ATA-TTGAATTTAATTTTTAAGTCAAAAAGCTTAAATAAATTTAAAGGACGAGAAGACCCTATAGAGTTTAATA----AAA-A-TTAT--TTTT--TT-G--AAATTTAA-TTT--
T-TT-TTAAA-AATTT-A-A-TTAAATTATTTTGTTGGGGCGATAGAAAAATTTAATAAACTTTTTTAT-A-AT-T-T------TTA-CATTAATTAATGAT-T-T----TTTGATCCTATA--T--T-T-TAGATTATTAGAAAAAATTACCTTAGGGATAACAGCGTTATTTTTTTTA-GAGTTCTTATCAATAAAGAAGATTGC????
TCTAAGTGGGTGGTAAACTCCATCTAAGGCTAAATATTACCACGAGACCGATAGCGAACAAGTACCGTGAGGGAAAGTTGAAAAGAACTTTGAAGAGAGAGTTCAAGAGTACGTGAAACCGTTCAGGGGTAAACCTGAGAAACCCGAAAGATCGA-ACGGGGAGATTCATCGTCAGCGACTCCGGCTTCTC-CGTGG-CTCGCGATGCATCCG-G-G--C-A-
C-T--C----GTGT-CCGTGGC-ACGCGGCCGCGG-TGTT----ACGCCCGGTGGCGCCGGCGTGCACTTCTCCCCTAGTAGGACGTCGCGACCCGTTGGGTGTCGGTCTAAGGCCCGGTT----GGACGACTGT-T--CCGGCGT--TCGC--GT-C-----GGTGCAGACC-A-C-CGGT-CGCCCGTCCGGCTGCCCGGCGGTAACCGTAC-
GGTATAGAGCCGCAT-T-GAA-CTGCGTCGGGCCCGCCGCAAGCGCGGTCAGCGTT----TCCCGGTGGTCGGAC-CTAGCGCCGTCCCCGGGCCTGATCAGCTGTTGGCTGGCGGTGTCCTCTGACTGGCTC-ATT---T-GAAA-T-----ATCAA-CATACCGGTCGGCGACGCTATTGCTTTGGGTACTTTCAGGAC?
ATACTTATTCGAACAGAACTAGGAATACCAGGATCCTTAATTGGAGATGATCAAATTTATAATGTAATTGTAACTTCACACGCATTTTTAATAATTTTCTTTATAGTAATACCTATTATAATTGGAGGATTTGGAAACTGACTAGTACCTTTAATATTAGGAGCCCCCGATATAGCATTTCCTCGATTAAATAACATAAGATTTTGATTTTTACCACCATCACT
AATTCTATTACTTTCAAGAAGAATAGTAAATTCAGGTTCTGGAACAGGTTGAACAGTTTATCCTCCTTTATCAAGAAGAATTTCACATGCAGGAGCATCAGTAGATCTTACCATTTTCTCTCTTCATTTAGCAGGAATTTCATCAATTCTTGGAGCAATTAATTTCATTTCAACAATAATTAATATAAAAGTAAAAGGAATAAGATTTGAACGAATACCTCTAT
TTGTTTGAGCAGTATCATTAACAGCTTTATTACTTCTTCTTTCACTACCTGTACTAGCCGGAGCAATTACTATACTTCTCACAGATCGAAACTTAAATACATCATTTTTTGACCCATCAGGAGGAGGAGACCCAATTTTATACCAACACTTATTT?????????????????????????????????????????????????????????????????????
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GTGAAGAAGGAGATTGACGAAATCGTAGCCTTCTTAAAAGCCGGTCCACTGGATCCCAACGTCGGTACGTTTTCAGAAATTGTTGTCGGCGTACCGGCTCTTTATTTGACCTACGCTAAGAGCGTACTGCCTCCCAATGTCCAAGTCAGCGCACAGAACAGCTACAAAGTTGCCAAGGGTGCTTTCACGGGTGAAATTAGTCCAGCGATGCTGTTAGACAGTGG
AATTCCGTGGGTCATTCTTGGCCATTCCGAACGTCGCAATGTTTTCGGAGAATCGGATGAACTTATTTCAGAAAAAATTGCTCATGCTCTGGAGGCTGGCTTAAAGGTGGTGATAGCCTGTATCGGAGAGAAATTGGACGAACGTGAGGCTGGAAAGACTGAAGAAGTTGTGTTCAAGCAAACCAAGGCGATCGCCGACAAAATCAAGTCATGGGATAACGTTG
TTTTG???????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????
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 Corynis_crassicornis ATAATAATTTGTCTTTTAATTAAAGGCTAGAATGAATGATTGGACGAGATAAAATCTGTCTCT-AATTAATTT-TTATTTGAATTTAATTTTTAAGTTAAAAAGCTTAAATTTTTTTAAAGGACGAGAAGACCCTATAGAGTTTTATA----ATA-A-TATT--TTTTATTT-A--ATTT-TAA-TTT--
T-T--TTAAT---AAT-A-A-ATTTATTATTTTATTGGGGTGATAGGAATATTAATAGAACTTTTTTAA-AAAA-T-T------TTA-CATTTATTTATGAA-T-T----TTTGATCCTAAA--A--T-T-TAGATTATTAGATTAAATTACCTTAGGGATAACAGCGTAATTTTTTTTA-
GAGTTCTTATCGAAAAAGAAGATTGCCAGCTCTAAGTGGGTGGTAAACTCCATCTAAGGCTAAATATGACCACGAGACCGATAGCGAACAAGTACCGTGAGGGAAAGTTGAAAAGAACTTTGAAGAGAGAGTTCAAGAGTACGTGAAACCGTTCAGGGGTAAACCTGAGAAACCCGAAAGGTCGA-ACGGGGAGATTCATCGTCAGCGTCGCTGGCTTCGC-
CGCGG-TTCGTGATG--C-CG-G-G--A-C-C-C--C----GCGTC-CACGGC-ACTCGACCGCGG-TGCA----ACGCCCGGTGACGCCGGCGTGCACTTCTCCCCTAGTAGGACGTCGCGACCCGTTGGGTGTCGGTCTAAGGCCCGGGTT---GGGTGCCTGT-C--TCGGCGT--TCGC--GT-C-----GGGGCAAACC-C-C-CGGT-
TGCCCGGCCGGCCGCCCGGCGGTAAACGCAC-GGTACAGAGCCGCAT-C-TAA-CTGCGTCGGGCCCGCCGCAAGCGCGGTCAGCGAT----TCCCGGTGGTCGGAC-CTAGCGCCGTCCCCGGGCCTGGCCAGCTGTTGGCAGGCGGTGTCCTCTGACGGGCTC-AAT---C-GAAT-C---AATTTTA-
CGTACCGGTCGGCGACGCTATTGCTTTGGGTACTTTCAGGACCATRTTAATTCGTAGAGAATTAGGTACACCAAATTCTTTAATTGGWGATGATCAAATYTATAAYACAGTAGTWACTTCYCATGCATTTTTAATAATTTTTTTTATAGTAATACCAGTWATAATCGGAGGATTTGGAAAYTGATTAATTCCTTYAATACTAGGAGCACCAGATATAGCATTTC
CTCGTTTAAATAATATAAGATTTTGATTATTACCTCCATCATTAWTAYTATTATTAATAAGAAGAATTATTGATTCAGGAKCAGGAACTGGATGAACARTTTATCCACCATTATCAAGAAATTTAGCWCATGCAGGAGCTTCTGTAGAYATAACAATTTTYTCACTACATATAGCAGGAATCTCTTCAATTTTAGGAGCYATTAATTTTATTTCTACTATTATA
AATATAAAAATTAATGGRATAAARTTTGATCAATTACCTTTATTTGTTTGAGCTGTAATTTTAACTACTATTYTATTATTATTATCMTTACCTGTTTTAGCRGGAGCTATTACAATATTATTAACAGATCGAAATTTAAATACATCATTTTTTGATCCATCMGGAGGAGGAGATCCAATYTTATAYCAACATTTATTCTGATTTTTTGGCCACCCAGAAGTATA
TATTYTAATTTTACCAGCATTTGGTGTTATYTCTCATATTATTTCTAACGAATCAGGAAAAAAAGAAACATTTGGAACATTAGGWATAATTTATGCYATATTATCAATTGGTTTAYTAGGTTTYGTAGTTTGAGCTCATCATATATTTACTGTTGGTATAGATGTTGATACACGAGCATATTTTACTGCAGTAACWATAATAATTGCAATTCCTACAGGAATTA
AAATTTTTAGATGATTAGCAACTCTMTTYGGARCAATTATAAAATCAATTCCTTCAATATTATGAACWATAGGATTTGTATTCTTATTTACAATAGGAGGATTAACAGGTATCATTTTATCTAATTCATCAATTGATGTAATTTTACATGATACATATTATGGATTCAATTAAGGAAAAACTGATACTTCCTTTCCTCGACATTGAACTCCATATCTATGATTT
GGGGATGGAAAATCGAGATAAAACCAATGACCAAGTAACAGTAGATTGCGCTGAAGCTATTAAGAAATACAATGTGGGTATAAAATGTGCGACAATAACACCTGATGAGAAGCGAGTCGAAGAATTTAAACTTAAACAAATGTGGAAAAGCCCCAATGGCACAATCAGGAACATATTAGGTGGTACTGTCTTCAGGGAAGCAATTATTTGCAAAAATATTCCAC
GCTTGGTTACTGGGTGGAATCAGCCTATAATTATTGGTCGTCATGCACATGCCGATCAGTATAAAGCTACAGATTTTGTAGTTCCTGGTGCTGGCAAATTGGAAATCACTTGGACAGGTGAATGTGGTGACAAGATCCACCATACCGTTTACCAATTTAAAGGACCAGGTGTTGCCCAGGCTCAATTCAATACCGACGAAAGTATTCAGGCTTTTGCTCACAGC
TCCTTCCAGTACGCCTTATCTCGAAGCTATCCACTTTATCTTTCAACCAAGAACACCATTTTGAAGAGATATGATGGTAGATTCAAAGATATTTTCCAAGCTATTTACGAATCTGATTACAAGTCTAAGTTTGAGGCCAAGAATATCTGGTACGAACATCGTCTGAT??????????????????????????????????
GATATGCAGCTGATCTGTGAAGCATATCATATTATGAGAAATGGACTAGGACTCAATCCAAAAGAAATGAGTGATGTTTTTGGTGAATGGAACAAGGGGGTACTTGATTCTTTCCTGATAGAAATCACCAGAGATATTTTGAAGTATAAGGATGATAAAGGATATCTGCTAGAGCGAATTAGAGATACAGCTGGGCAAAAGGGTACTGGAAAGTGGACAGCAAT
TGCTGCTCTAGATTACGGCGTTCCTGTTACTCTAATCGGAGAGTCAGTATTTGCCAGGTGTCTTTCTTCGCTTCAGAGCGAGAGGATAGAAGCCAGCACTGTTTTAGA------------------
TGGTCCAAATACCATCTATCAAGGTGACAAAAAACAGTTCCTCGAACATCTTAGAAAAGCACTCTATGCTTCGAAAATTATTTCTTATGCTCAAGGATTCATGCTACTAAGAGAAGCTGCTAAAATTCATAACTGGAATTTAAATTACGGCGGAATCGCACTTGTCAAAAAGGAAATTGACGAAATCGTCGCCTTCCTAAAAACAGGTCCACTTGATCCGAATG
TTGGTACGTTTTCAGAAGTTGTCGTTGGTGTGCCTGCTTTGTACCTGACTTATGTCAAAAGCATATTACCACCAAATGTACAAGTTAGTGCTCAAAATTGCTACAAAGTGCCCAAAGGTGCTTTCACTGGTGAAATCAGTCCTGCTATGCTTTTGGACAATGGAATTTCTTGGGTAATTCTAGGACATTCAGAACGTCGTAACGTTTTTGGCGAATCAGACCAA
CTCATTTCTGAAAAAGTAGCTCATGCCCTTGATGCTGGTTTAAAGGTAGTGATTGCCTGTATTGGAGAGAAACTGGATGAACGCGAAGCTGGAAAGACCGAGGAAGTTGTATTCAAGCAGACTAAAGCTATAGCTGACAAAATCAAGTCTTGGGATAATGTAGTACTGTTTCAGATTGGTAGCTCAATGAAAAGCGTGGGAGAGGCGATGGCGATCGGTAGAAA
ATTTGAGGAGGCTTTTCAGAAAGCTCTTCGTATGGTCGACGAAAATGTCAACGGTTTTGATCCTTACATAAAATCGATCGACGATGAAGAGCTGGAGAGACCAACCGACAAAAGGATGTTCGTCCTTGCAGCAGCACTGAAGGCTGGTTACACAATTGACCGACTGTATGAATTAACAAAAATTGATCGTTGGTTTTTAGAAAAAATGAAAAATATTACTTCGT
ATTATACAATTCTCGAAGGTCTCGATCAAACGAAACTTTCTCACGAAGTTCTATTGTGTGCGAAACAAATTGGTTTTTCTGACAAACAAATTGCAACAGCCGTGAAAAGTACGGAACTTGCTGTTCGTAAACAAAGACAGGAAAGCAATATCACACCTTTCGTCAAACAGATAGATACAGTAGCTGCTGAATGGCCAGCCACTACAAATTACCTTTACCTCACT
TACAACGGAAAAAGTCACGATCTTCAATTTCCTGGAGGTTACACGATGGTTATAGGTACGGCAGGTTCTGGCGTTTATCGCATAGGAAGTTCCGTGGAGTTTGATTGGTGTGCGGTAGGATGTCTCAGAGAATTACGTCAACTTAATAAAAAAACGATAATGGTAAATTATAATCCAGAAACAGTGAGCACAGATTATGACATGTGCGATCGTCTATATTTCGA
AGAAATTTCCTTTGAAGTCGTAATGGATATTTACGATCTCGAAAACCCGGAAGGT??????????????????????????????????????????????
TTAGCTTACAGAAAAAGTTTAAATCTTTATTTGGTGAAAAATTGGAGGTCGTTAGGACGCATCAGCAACAGGAAAACCTTAAGTTCATGGCACATTTTAAGCGCAAGTTTATCATTCGTCATGGAAAACGAAAAGACTCAAAATCACCTACTAATAACAAGGTTGAATTTTACCATCTTCGAAGTAATGGAAGTGCCTTATGTACCAGATTGATCCAAGTGAAT
CCAGATGCTGCTTTATTGAATTCTGCTTTCTGGTACAGCTATATATTAAATGTGCCATTTAATAACGACGATGAAACTGGTATCGTCTATGTTTGGATTGGATCTAAAGCTGACAGTGAAGAAGCTAGACTCGTGGAAGAGATAGCTGAGGATATGTTCAATAACGTTCCGTGGATTAGTCTGCAAATATTAAATGAAGGTGAAGAACCAGACAACTTTTTCTG
GGTTGGTCTTGGTGGTAAAAAACCATATGATACAAATGCCGATTACATGAATTACACAAGACTATTTCGATGTTCAAACGAAAAAGGTTACTTTACTATCAGTGAAAAATGTACAGATTTTTGTCAGGTAGATGATTTAGCTGACGATGATATCATGGTGCTAGATAATGGAGAACAAGTGTTTCTTTGGCTGGGTGCCAGATGCTCAGAGGTTGAAATTAAGC
TCGCCTATAAATCAGCTCAGGTAAAAGTGTACATACAGCATTTACGTGTGAAGCAGCCAGAGAGACCACGTAAACTGTTCCTTACTGCCAAAAGTAAAGAATCCCGAAGATTTACA

Euura_spp ??????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????
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CAGCTCTAAGTGGGTGGTAAACTCCATCTAAGGCTAAATATTACCACGAGACCGATAGCGAACAAGTACCGTGAGGGAAAGTTGAAAAGAACTTTGAAGAGAGAGTTCAAGAGTACGTGAAACCGTTCAGGGGTAAACCTGAGAAACCCGAAAGATCGA-ACGGGGAGATTCATCGTCAGCGACTCCGGCTTCTC-AGCGG-TTCGCGATG---TCG-G-G--
ACA-C-T--T----GTGT-CAACGGC-ACGCGGCCGCGG-TGTC----ACGCCCGGTGGCGCCGGCGTGCACTTCTCCCCTAGTAGGACGTCGCGACCCGTTGGGTGTCGGTCTACGGCCCGGGT----GGTCGCCTGT-C--CCGGCGT--TAGC--GT-C-----GGGGCAGACC-C-C-CGGT-TGCCCGTCCGGCTGCCCGGCGGTAACCGCAC-
GGTATTGAGCCGCAT-T-GAA-CTGCGTCGGGCCCGCCGCAAGCGCGGTCAGCGAT----TCCCGGTGGTCGGAC-CTAGCGCCGTCCCCGGGCCTGGTCAGCTGTTGGCTGGCGGTGTCCTCTGACTGGCTC-ATA---T------------AT------
TACCGGTCGGCGACGCTATTGCTTTGGGTACTTTCAGGACC???????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????
ATAGCATTTCCCCGAATGAATAATATAAGATTTTGATTTTTACCACCATCATTAATTTTATTATTATCAAGAAGAATAGTAAATTCAGGATCAGGAACAGGATGAACTGTTTATCCACCTTTATCAAGAAGAATTTCACATTCAGGAGCATCTGTAGATTTAACAATTTTTTCTCTTCATTTAGCCGGAATTTCATCAATCTTAGGAGCCATTAATTTTATCTC
TACAATAATTAATATAAAATTAAAGGGAATAAGATTTGAACAAATACCTTTATTTGTATGAGCAGTATCACTAACAGCACTATTATTATTATTATCACTACCAGTTTTAGCGGGAGCTATTACTATACTTCTTACAGATCGTAATCTAAATACATCATTCTTTGATCCATCTGGAGGAGGAGATCCAATTTTATATCAACATTTATTTTGATTTTTTGGTCACC
CAGAAGTTTATATTTTAATTATCCCAGCTTTTGGTATAATTTCACATATGATTTCACAAGATTCAGGAAAAAAGGAAACTTTTGGAACTCTTGGAATAATTTATGCTATAATAACAATTGGACTTTTAGGATTTGTTGTATGAGCACATCATATATTTACAGTAGGTATAGATGTTGATACACGAGCTTATTTTACTGCAGCTACAATAATTATTGCAATTCCA
ACAGGTATTAAAATTTTTAGATGATTAGCCACATTATATGGATCACAAATTATATTTAACCCATCAGTAATATGAACTCTTGGGTTTGTATTTTTATTTACATCGGGAGGATTAACAGGAATCATCCTATCAAATTCATCAATTGATATCATTCTACATGATACA???????????????????????????????????????????????????????????
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GACATGCAGTTGATCTGCGAAGCCTATCACATCATGAGGAACGGATTGGGTCTCAGTCCCAAGGAGATGAGCGACGTATTTGGAGAATGGAACAAGGGAGTGCTCGACTCTTTCCTCATCGAAATAACCAGAGACATCCTCAAGTACCAAGACGACAAGGGATACCTGTTGGAAAGAATCAGGGACACCGCTGGTCAAAAGGGAACCGGAAAGTGGACCGCGAT
AGCTGCCCTGGACTACGGAATCCCCGTTACTCTAATTGGAGAGTCGGTATTCGCCAGGTGTCTGTCCTCCCTTCAGAGCGAGAGGATAGAAGCCAGCGCTGTATTGGA------------------
GGGACCCAGCGGTATCTACCAAGGGGACAAGAAACAGTTCCTCGAACACCTTAGGAAAGCCCTCTACGTCGCTAAGATCATTTCCTACGCTCAGGGTTTCATGCTGCTCAGAGAAGCGGCCAAAATCCATAAGTGGAACTTGAACTACGGAGGCATCGCTCTCGTTAAGAAGGAGATCGACGAAATCGTCGCTTTCCTAAAAGCCGGTCCCCTCGATCCCAACG
TCGGTACGTCCTCAGAGGTCGTGGTTGGAGTCCCGGCTCTATATCTCACCTACGCCAAGAGCGTCCTTCCCCCCAACGTTCAAGTCAGCGCACAAAACAGCTACAAAGTAGCCAAGGGTGCTTTCACGGGCGAAATCAGCCCCGCTATGCTACTGGACAGCGGGATCCCTTGGGTCATCCTTGGTCACTCTGAGCGACGAAACGTTTTCGGAGAAACGGACGAA
CTTATCTCTGAAAAAGTCGCCCACGCTCTGGAAGCCGGTTTGAAGGTAGTAATCGCCTGCATCGGAGAGAAGCTTGACGAACGCGAGGCGGGAAAGACCGAAGAAGTCGTGTTTAGACAGACCAAGGCGATCGCTGACAAAGTCAAATCTTGGGACAACGTTGTTTTGTTCCAGATCGGTAGTTCGATGAAGAGCGTCGGCGAGGCGATGGCGATCGGTAGGAA
GTTCGAAGAGGCTATCCAGAAGGCTCTGAGGATGGTCGACGAGAACGTGAACGGTTTCGACCCGTACATAAAGCCGATAGACGACGAGGACCTTGAGAGGCCAACGGACAAACGGATGTTCGTATTGGCGGCTGCTCTGAAGGCAGGATACACGATAGACCGACTGTACGAGCTGACGAAAATCGATCGTTGGTTTTTGGAAAAGATGAGAAACATAACGTCGT
ATTATACTCTCCTTGAGGATCTAGACCAGACAAAACTGTCCCACGAGGTTTTACTGCGTGCGAAGCAAATCGGTTTCTCCGACAAGCAGATCGCGACTGCGGTCAAGAGCACGGAACTCGCGGTGAGGAAACATCGCCAGGAGAGCAACATCAGACCGATAGTCAAACAAATCGATACCGTTGCCGCCGAGTGGCCGGCCACCACCAACTATCTCTACCTCACG
TACAACGGAAATAGCCACGATATTCTCTTTCCTGGTGGATACACGATGGTCATAGGTTCGTCAGGTTCCGGCGTCTACCGGATAGGCAG???????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????
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Heptamelus_dahlbomi ??????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????
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ATAATTATTCGAACAGAATTAGGAACACCCGGATCATTAATTGGTAATGATCAAATTTATAATACAGTAGTAACTTCACACGCTTTCTTAATAATTTTTTTTTTAGTTATACCAGTAATAATTGGAGGATTTGGTAATTGATTAATTCCATTAATATTAAGAGCACCTGATATAGCATTCCCACGAATAAATAATATAAGATTTTGATTATTACCTCCATCCCT
AACAATCCTTTTATTCAGAAGAATTGTTGACTCAGGATCAGGGACAGGATGAACTGTTTATCCACCACTATCAAGAAGAATTGCACATGCAGGATCATCAGTAGATTTAACTATTTTTTCACTTCATTTAGCAGGAATTTCATCAATTTTAGGAGCAATTAATTTTATTTCAACTATTATTAATATACGAACTCTTGGAATAAAATTTGAACAAATACCACTAT
TTATTTGAGCCGTATTATTAACAACTATTTTATTATTATTATCCTTACCTGTTCTAGCTGGAGCCATTACCATATTATTAACCGATCGAAACTTAAATACATCTTTCTTTGATCCATCAGGAGGAGGAGATCCAATTTTATATCAACACTTATTT?????????????????????????????????????????????????????????????????????
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Leptocimbex_yorofui ??????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????
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Monoctenus_juniperi ATAATAATTTGTCTTTTAATTTAGGACTAGAATGAATGATTTAACGAAGTTTAAGCTGTCTTA-AATTTAT-A-ATATTTGAATTTAATTTTTAAGTTAAAAAGCTTAAATTTTTTTATAGGACGAGAAGACCCTATAGAGTTTTATT-AAAAT--A-TAAT--AATTTTTT-T--AAATAATA-ATT--
T----TTATT-AAAAT-T-T-TATTTTAATTTTATTGGGGTGATAATAAAATTTTAATAACTTTTATAT-T-TC-A-A------TAA-CATAAATTAATGAA-T-T----ATTGATCCTTTT--T--T-T-AAGATTATTAGATTAAATTACCTTAGGGATAACAGCGTAATTTTTTTTA-
AAGTTCTTATTAACATTAAAGATTGCCAGCTCTAAGTGGGTGGTAAACTCCATCTAAGGCTAAATATGACCACGAGACCGATAGCGAACAAGTACCGTGAGGGAAAGTTGAAAAGAACTTTGAAGAGAGAGTTCAAGAGTACGTGAAACCGTTCAGGGGTAAACCTGAGAAACCCGAAAGATCGA-ACGGGGAGATTCACCGTCAGCGGGGCTGGCTTCGC-
CGCGG-CTCGTGATG--T-CG-G-G--A-C-C-T--C----GCGT-CCACGGC-ACTCGGTCGCGG-TGCC----ATGTCCGGCCTCGCCGGCGTGCACTTCTCCCCTAGTAGGACGTCGCGACCCGTTGGGTGTCGGTCTAAGGCCCGGGA----GGCTGTCTGT-C--TCGGCGT--TCGC--GT-C-----GTGGCAGACC-C-C-CGGT-
TGCCCGTCCGGCTGCCCGGCGGTACCCGCAC-GGTATAGAGCCGCAT-C-GAA-CTGCGCCGGGCCCGCCGCAAGCGCGGTCAGCGAT----TCCCGGTGGTCGGAC-CTAGCGCCGTCCCCGGGCCTGGCCAGCTGTTGGCAGGCGGTGTCCTCTGGCCGGCTC-AAT---C-GAAT-------ATTCA-
TATACCGGTCGGCGACGCTATCGCTTTGGGTACTTTCAGGGAGATAATTATTCGGATAGAATTAGGAATACCAGGTTCATTAATTGGAGATGATCAAATTTATAATGTAATTGTAACTTCACATGCATTTTTAATAATTTTTTTTTTAGTTATACCTATTATAATTGGGGGGTTTGGTAATTGGTTAGTTCCTCTTATATTAGGAGCTCCAGATATAGCTTTCC
CACGGATAAATAATATAAGATTTTGATTTTTACCTCCATCTTTAATTTTAATAATTACAAGAATAATAGTAGAATCAGGATCAGGAACGGGGTGAACAGTTTATCCCCCGTTATCTAGAAGATTGGGGCATTCTGGTAGATCTGTAGATTTAACTATTTTTTCACTTCATTTAGCAGGAATTTCTTCTATTTTAGGGGCTATTAATTTTATTTCAACAATAATT
AATATACGAATAAATGGTATAAATTTTGATCAACTTTCTCTTTTTGTATGATCAGTTAAAATTACAGCAATTTTATTATTATTATCTTTACCTGTTTTAGCTGGCGCTATTACTATATTACTTACAGATCGAAATTTAAATACAACATTTTTTGATCCTGCTGGAGGAGGGGACCCAATTTTATATCAACATTTATTTTGATTTTTTGGGCATCCGGAGGTTTA
TATTTTAATTTTACCTGCTTTTGGTATAATTTCTCATATTATTTCTCAAGAATCAGGTAAAAAGGAAACATTTGGAACATTGGGGATAATTTATGCAATAATAACTATTGGATTATTAGGATTTATTGTATGGGCACATCATATATTTACTGTAGGAATAGATGTTGATACTCGAGCATATTTTACAGCGGCAACTATAATTATTGCAATTCCTACGGGGATTA
AAATTTTTAGATGATTAGCTACATTATATGGATCAAATATAAAATTTAATTCTGTTACATTATGAGCATTAGGTTTTATTTTTTTATTTACAATAGGGGGATTAACTGGTATTATATTATCAAATTCTTCTATTGATATTATTTTACATGATACATATTATG??????????????????????????????????????????????????????????????
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GACATGCAGCTGATCTGCGAGGCTTATCACATCATGAGAAACGGACTGGGTCTCAGCCCGAAAGAAATGAGCGATGTTTTCGACGAGTGGAACAAGGGGGTGCTCGATTCTTTCCTGATTGAAATCACCCGGGATATATTGAAGTACAAAGACGAAAAGGGCTATCTGCTGGAGAGAATCAGAGACACGGCTGGGCAGAAGGGCACAGGAAAGTGGACGGCCAT
CGCGGCCCTAGACTACGGGATTCCTGTCACTCTAATTGGAGAATCAGTCTTCGCCAGGTGTCTCTCGTCGCTCCAGAGCGAGAGGATAGAAGCCAGCACAGTCTTGGA------------------
GGGTCCAAACACCCTCTACCAAGGGGACAAGAAACAGTTCCTCGAACATCTCAGGAAAGCCCTCTACGCCTCGAAAATCATTTCGTATGCCCAAGGATTCATGCTGCTGAGGGAAGCGGCCAAAATCCACAACTGGAATTTGAATTATGGTGGAATTGCACTCGTCAAAAAGGAAGTTGACGAAATCGTCGCTTTCCTTAAGGCCGGTCCTCTCGATCCCAATG
TCGGTACGTTTTCAGAGGTCGTTGTCGGGGTTCCGGCTCTCTACCTGAACTACGTCAAAAGTATATTGCCCCCGAACGTACAAGTCAGCGCTCAGAACAGCTACAAGGTAGCGAAAGGCGCTTTCACTGGAGAAATCAGCCCTGCGATGCTCGTAGACAACGGTATCCCTTGGGTGATCCTGGGACACTCGGAACGTCGTAACGTTTTTGGCGAATCTGACGAA
CTGATCTCTGAAAAGATTGCTCATGCCCTTGAATCCGGCGTTAAGGTAGTGATCGCATGTATCGGAGAGAAACTGGACGAGCGTGAGGCTGGGAAAACCGAGGAGGTAGTGTTCAGGCAAACCAAAGCGATCGCGGACAAAATTAAGTCCTGGGACAACGTAGTACTC????????????????????????????????????????????????????????
??????????????????????????????????????????????????????????????????
TCTGAAAGCGATCGATGACGAGGACCTTGAGAGACCAACGGACAAGAGGATGTTCGTGCTGGCAGCAGCATTGAAGGCCGGCTACACGATCGATCGTTTATACGAGCTAACAAAAATCGACCGTTGGTTTTTGGAGAAGATGCGAAACATCACATCGTATTACACGCTTCTCGAAGATCTCGATCAGACGAAGCTCTCCCACGAAGTTCTGCTGCGGGCCAAGC
AAATCGGCTTTTCTGATAAGCAGATCGCTACTGCGGTGAAAAGCACCGAACTCGCTGTCCGCAAGCAACGACAGGAAAGCAGTATCAGACCTTTTGTCAAGCAGATAGACACTGTCGCTGCCGAGTGGCCAGCGACCACGAACTACCTTTACCTCACCTACAATGGAAACAGTCACGACCTTCAGTTCCCTGGCGGCTACACCATGGTCATAGGTGCGTCAGGT
TCTGGCGTGTATCGCATTGGAAGTTCGGTTGAGTTCGACTGGTGCGCCGTGGGCTGTCTCAGGGAATTGCGTTGTCTCAACCGTAAGACGATAATGGTGAACTACAACCCGGAGACAGTGAGCACCGACTACGATATGTGCGATCGCTTATACTTCGAGGAAATATCCTTCGAGGTCGTGATGGATATATACGATCTCGAGAACCCGGAAGGC???????????
???????????????????????????????????
TTAGTTTACAGAAAAAGTTTAAGTCCTTATTTGGCGAAAAACTAGAGGTTGTCAGGACACATCAACAGCAAGAAAACCTAAAGTTCATGGCTCACTTTAAACGCAAGTTTATCATTCATCATGGAAGGCGAAAAGAAGCAAAATCACCTAGCAATAACAAGGTCGAATTTTACCATCTTCGAAGCAATGGTAGCGCCTTGTGTACCAGGTTGATACAAGTGAAT
CCAGATGCTTGTTTATTGAATTCTGCCTTCTGGTACAGCTACATCTTAAATGTACCTTTCAACAACGATGACGAAACTGGTATCGTTTATGTATGGATCGGGTCCAAGGCTGACAGCGAAGAGGCAAGACTTGTGGAAGAAATAGCAGAGGAAATGTTCAACAATGTTCCATGGATAAGTTTGCAAGTTTTGAACGAGGGTGAAGAACCAGACAACTTCTTCTG
GGTCGGCATTGGGGGAAAGAAACCGTATGACACTGACGCGGACTACATGAACTATACAAGATTATTTAGGTGCTCAAATGAAAAGGGTTACTTCACGATCAGTGAAAAATGTACAGATTTTTGTCAGGTAGATGATTTAGCTGACGACGACATTATGGTGCTAGACAACGGGGAGCAAGTATTCCTCTGGCTTGGTGCTCGATGTTCAGAAGTAGAGATTAAGC
TTGCTTACAAATCAGCTCAGGTAAAAGTTTACATACAGCATTTGCGGGTTAAGCAGCCTGAAAGACCTCGGAAATTGTTTCTTACCGCGAAAAGCAAA??????????????????
Neodiprion_sertifer ATAATAATTTGTTCTTTAATTTAGGACTAGAATGAATGATTTGACGAGATTAAAGCTGTCTTA-TAATTATTA-ATTATTGAATTTAATTTTTAAGTTAAAAAGCTTAAATTTTATTAGAGGACGAGAAGACCCTATAGAGTTTTATA----ATT-A-AATT--AATTTAAT-T--AATTAATT-ATT--
T----TTATA-AAATT-A-A-TATAATTATTTTGTTGGGGTGATAATTAAATTTAATAAACTTTTTATT-T-AA-T-T------TTA-CATTAATTAATGAA-T-T----TTTGATCCTATT--T--T-T-TAGATTAATAGATTAAATTACCTTAGGGATAACAGCGTTATTTTTTTTA-
AAGTTCTTATTGACATAAAAAATTGC???????????????????????????????????????????????????????????????????????????????????????????????GAAGAGAGAGTTCAAGAGTACGTTAAACCGTTCAGGGGTAAACCTGAGAAACCCGAAAGATCGA-ACTGAAAGATTCATCGTCAGCGACGCAGGCTTCAC-
CGCGG-CTTGTGATG---TCT-G-G--A-C-C-T--C----GCGT-CCACGGC-ACTCGGTCGCGA-TGCA----ATGTCTGGCGGCGCTGGCGTGCACTTTTCCCCTGGTGGGACACCGCGACCCGTTGGGTGTCGGTCTAAGGCCCGGTC----GGTCGCCTGT-C--TCGGCGT--TTGC--GT-C-----TGGGCGGACC-C-C-TGGT-
TGCCCGTTCAGCTGCCCGGGGGTACCCGCAA-GGTAAAGAGCCGCAT-T-TAA-CTGCGTCGGGCCCGCCGCAAGCGCGGTCAGCGAT----TCCCGGTGGTCGGAC-CTAGCGCCGTTCCCGGGCCTGGCCAGCTGTTGGCTGGCGATGTCCTCTGGCTGGCTT-GTG---C-AAA--T-----TATCA-
GATACCGGTCGGCGACGCTATTGCTTTGGGTACTTTCAGGACCATAATTATTCGTATAGAACTTGGTACACCATCATCACTTATTGGTAATGATCAAATTTATAACATAATTGTTACATCACACGCTTTCTTAATAATTTTTTTTATAGTTATACCTATCATAATTGGAGGATTTGGAAATTGATTAATCCCTCTAATATTAGGAGCACCAGACATAGCTTTTC
CTCGAATAAACAATATAAGATTTTGACTTCTACCTCCTTCATTAATTTTACTTATTTCAAGAAGATTAGTAGATTCAGGAACAGGAACTGGTTGAACAGTTTATCCACCTTTATCAAGAAATATAGGTCATTCAGGACCATCTGTTGATTACACAATTTTCTCTTTACATATAGCAGGAATTTCATCTATTCTTGGAGCAATTAATTTTATTTCTACAATAATT
AATATACGACCAAAAGGAATAAAATTTGAACAAATACCTCTATTTGTTTGAGCAGTAAAACTAACAGCTATTCTTCTATTATTATCTCTTCCCGTATTAGCAGGAGCAATTACAATATTATTAACTGATCGAAATATAAATACAACTTTTTTTGATCCTTCAGGAGGGGGAGACCCAATCTTATACCAACACTTATTTTTTTT?????????????????????
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GACATGCAACTGATCTGCGAAGCCTATCACATCATGAGAAACGGGCTTGGTCTGAACCCACAGGAAATGAGCGACGTTTTTGGCGAGTGGAACAAGGGGGTGCTTGATTCTTTCCTAATTGAAATTACCCGAGATATCTTGAAGTACAAGGATGGGAAGGGCTATCTTCTCGAAAGAATCAGAGACACAGCTGGGCAGAAGGGTACAGGAAAGTGGACGGCTAT
CGCTGCCTTGGATTACGGAATTCCTGTCACGCTAATTGGAGAATCAGTGTTTGCAAGGTGTCTCTCCTCGCTCCAAGGTGAGAGGATTGAAGCTAGCACCGTTTTGGA------------------
GGGTCCAAACGCTGTTTACCAAGGAGATAAAAAGCAATTCCTCGAACATCTTAGGAAAGCGCTGTATGCTTCGAAAATCATTTCGTATGCTCAAGGATTCATGCTGCTAAGAGAAGCTGCTAAAATTCACAACTGGAATTTGAATTATGGTGGAATTGCGCTCGTGAAAAAGGAAATTGACGAAATCGTTGCATTTCTAAAGACCGGCCCCCTCGACCCGAATG
TCGGTACGTTTCCAGAGGTCGTTGTTGGTGTACCTGCCATTTACCTGAACTATGCCAAGAGTATATTACCTGCCAACGTACAAGTCAGTGCACAGAACAGTTACAAGGTAGCTAAAGGTGCTTTTACTGGAGAAATTAGCCCTGCGATGCTTGTGGACAATGAGATCCCTTGGGTAATCCTAGGACACTCTGAACGTCGAAACGTGTTTGGCGAGTCAGATGAA
CTCATTTCTGAAAAGATTGCTCATGCCCTTGAAGCTGGTGTAAAGGTAGTGATTGCCTGTATCGGAGAGAAACTGGACGAACGTGAAGCTGGAAAAACTGAAGAAGTTGTGTTCAGGCAGACTAAGGCGATTGCGGATAAAATCAAATCCTGGGACAATGTTGTACTTTTACAGATCGGAAGTTCGATGAAGAGCGTCGGCGAGGCGATGGCGATCGGTAGAAA
ATTTGAGGAGGCTTTCCAGAAGGCGCTTCGCATGGTGGATGAAAACGTGAACGGTTTCGACCCGTATCTGAAAAAGATTGATGACGAGGACCTGGAGAGACCGACAGATAAGAGGATGTTCGTTTTGGCAGCAGCATTGAAGGCTGGCTATACGATCGATCGACTGTACCAGTTGACGAAAATCGATCGTTGGTTTTTGGAAAAAATGAAAAATATCACATCTT
ACTACACACTTCTCGAAGATCTCGACCAGACGAAGCTCTCCTACGAAGTTCTACTACGTGCCAAACAAATCGGTTTTTCTGACAAGCAGATTGCGACAGCGGTGAAAAGCACCGAACTTGCTGTTCGTAAGCAACGACAGGAAGGCAACATTAGACCGTTTGTCAAACAGATCGATACTGTCGCTGCCGAGTGGCCGGCGACCACAAACTACCTGTACCTGACC
TACAATGGAAATAGCCACGACCTTCAATTCCCCGGGGGCTACACCATGGTTATTGGTACGCTAGGTTCCGGCGTGTACCGGATTGGAAGTTCGGTTGAATTTGACTGGTGTGCTGTTGGCTGCCTTAGGGAATTGCGTTGTCTCAACCGAAAGACTATAATGGTGAATTACAACCCGGAGACAGTGAGCACAGATTACGATATGTGCGATCGTTTGTACTTTGA
GGAAATATCCTT?????????????????????????????????????????????????????????????????????????????????????????
TTAGTTTACAGAAAAAGTTTAAATCTTTATTTGGTGAAAAATTGGAGGTCGTCAGGACTCATCAGCAGCAAGAAAATCTGAAATTCATGGCACATTTTAAACGCAAGTTTATCATTCGCCAAGGAAAACGAAAAGAAGTGAAATCACCTAGTAATAATAAGGTTGAGTTTTACCATCTCCGAAGCAATGGTAGTGCCTTATGTACTAGATTAATACAAGTTAAT
CCAGATCCTTTTTTATTAAATTCTGCTTTCTGGTGCAGCTATATTTTAAATGTGCCTTTCAACAACGACGATGAAACTGGTATCGTTTACGTATGGATCGGGTCAAAAGCTGACAGTGAAGAAGCAAGACTTGTGGAAGAAATAGCAGAGGAAATGTTTAACAACGTTCCATGGATCAGCCTGCAAGTCCTAAACGAGGGTGAAGAACCGGATAACTTTTTCTG
GGTCGGCCTTGGGGGGAAAAAACCATATGATACCAATGCGGAATACATGAATTACACAAGACTATTTAGGTGCTCAAACGAAAAGGGCTACTTCACAATCAGTGAAAAATGCACAGATTTCTGTCAGGTAGATGATTTAGCCGACGATGATATTATGGTGCTAGATAATGGGGAGCAAGTATTCCTTTGGCTGGGGGCCCGATGCTCGGAAGTGGAGATTAAAC
TTGCTTATAAGTCAGCTCAGGTAAGGGTCTACATACAGCATTTGCGTGTTAAGCAGCCGGATAGGCCTCGAAAATTGTTCCTCACTGCCAAAAGCAAGGAGTCTCGAAGATTTA??

Pachylosticta_albiventris ??????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????
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Runaria_reducta ATAATCATTAGTCTTTTAATTGAAGACTTGTATGAAGGATTGAATGAAATATATACTGTCTCAT-TAAAATTT-AAAATTGAATTTAATTTTTGAGTTAAAAAGCTTAAATATTATTAGAGGACGAGAAGACCCTATAGAGTTTTATA-T---TT-A-TTAA--A--ATATT----AATTAAAT-GT--
AT-AAATTAAT-A-TTG-A-T-GATATTTATTTTGTTGGGGTGATGGGAAAATTTGTTAAACTTTTTTATGTA-T-T-------TATA-CATTAATATATGAT-TTT----ATTGATCCTTAA--T--T-A-
AAGATTAATAGATTAAATTACCTTAGGGATAACAGCGTAATTTTTTTTTAAAGTTCATATTTGAAGAAAAGATTGCCAGCTCTAAGTGGGTGGTAAACTCCATCTAAGGCTAAATACGACCACGAGACCGATAGCGAACAAGTACCGTGAGGGAAAGTTGAAAAGAACTTTGAAGAGAGAGTTCAAGAGTACGTGAAACCGTTCAGGGGTAAACCTGAGAAACC
CGAAAGATCGA-ACGGGGAGATTCATCGTCAGCGACGCCGGCTTCGC-CGCGG-CTCGTGATG---CCG-G-G--A-C-C-A--C----GCGT-CCACGGC-ACTCGGTCGCGG-TGTC----ACGTCCGGTGGCGCCGGCGTGCACTTCTCCCCTAGTAGGACGTCGCGACCCGTTGGGTGTCGGTCTACGGCCCGGGTTTG-GTTCGCCTGT-C--
TCGGCGT--TCGC--GT-C-----GGGGCAGACC-C-C-CGGTTTGCCCAACCGGCCGCCCGGCGGTACTCGCAC-GGTATTGAGCCGCAT-T-GAA-TTGCGTCGGGCCCGCCGCGAGCGCGGCCAGCGAT----ACCCGGTGGTCGGAC-CTAGCGCCGTCCCCGGGCCTGGCCAGCTGTTGGCTGGCGGTGTCCTTTGACGGGCTC-GCT---T-
CGA----------------
TACCGGTCGGCGACGCTATTGCTTTGGGTACTTTCAGGACCGTATTAATCCGATTAGAACTAAGAACACCTACTAGATTTTTAGGAAGAGATCAAACTTATAATACTTTAGTAACCTCTCATGCATTCTTAATGATTTTTTTTATAGTTATACCTATTATAATTGGAGGATTCGGTAATTGATTAGTACCCTTAATAATCTCTGCCCCTGACATAGCCTTCCCT
CGAATAAATAATTCAAGATTTTGATTATTGCCCCCCTCTCTTATTTTATTAACTTCAAGAAGATTTATTGATCTAGGATCTGGTACAGGATGAACAATTTACCCCCCTCTATCAAGAAATTTAGGACATGCTAGATCATCTATAGACTTAACAATTTTTTCCTTACATCTAGCTGGAATTTCATCAATTATAGGTGCTATTAACTTTATTTCAACAACTATTAA
TATACGAAATAAAGGAATAACATCTGAACGACTTACATTATTCACATGATCTGTAGCAATTACTGCATTATTGTTAATACTATCATTACCTGTATTAGCAGGAGCCATCACAATATTACTGACAGACCGAAATTTAAATACATCATTTTTCGACCCTTCTGGGGGTGGAGACCCGATTCTATTCCAACACTTATTTTGATTCTTTGGACACCCTGAAGTATACA
TTTTAATTTTACCTGGATTTGGAATTATTTCCCATATTATTTATCAAGAAGCAGGAAAAAAAGAAACATTTGGAACTTTAGGAATAATCTATGCTATATTAACAATTGGCATTCTAGGATTTGTTGTTTGAGCTCATCATATATTTACAGTAGGAATAGATGTTGATACTCGAGCATACTTTACAGCCGCTACAATAATTATTGCCGTTCCCACTGGAATTAAA
ATTTTTAGATGATTAGCAACAATCCACGGAACCCAACTAATTTATACCCCTTCAATTTTATGAACACTAGGATTTATTTTTCTATTTACTTTAGGAGGACTAACAGGTATTATATTATCTAACTCATCATTAGATATTATTTTACATGATACATATTATGGATTCTATAAAGGAAAAGCTGATCCTGCCGTTCCTCGACGTCGAGCTACACACCTACGATCTCG
GTATGGAGAACCGCGATGCGACCGACGACAAAGTAACGGTGGACTGCGCGGAGGCGATAAAGAAGTATCACGTAGGGATAAAATGCGCGACGATAACTCCGGACGAGAAGAGAGTGGAGGAGTTTGGCCTGAAACAGATGTGGAAGAGTCCGAACGGTACGATCAGGAATATCCTGGGCGGAACCGTGTTCCGCGAGGCGATAATCTGCAAGAACATTCCGCGC
TTAGTCTCTGGCTGGAACCTGCCGATCATCATTGGTCGGCACGCCCATGCCGATCAGTACAAAGCCACCGACTTTGTCGTCCCCGGGGCTGGAAAGCTCGAAATTATGTGGACCGGCACTTCGGGGGATAAAATCCATCATACCGTGCACGAATTTAAGGGAGCGGGCGTCGCGCAGGCCCAATTCAACACGGACGATAGCATCCGGGCGTTCGCTCGCAGCTC
TATGCAATACGCGCTGTCGCGAAGCTACCCGCTGTACTTGTCGACCAAGAACACTATCTTGAAGAAGTACGACGGCAGGTTCAAGGACATCTTCCAGGAGATCTACGAGAAGGATTACCGGTCCGAGTTCGAGGCCAAGAAGATATGGTACGAGCATCGCTTGATCGACGACATGGTCGCTTACGCGATGAAATCGGAGGACATGCAATTAATTTGCGAAGCCT
ATCACATCATGCGAAACGGTTTGGGACTTAATCCCAAGGAGATGAGCGATGTTTTCGACGAATGGAACAAGGGCGTACTCGATTCCTTTTTAATTGAAATTACCAGAGATATTTTGAAGTACAAGGATGAAAAGGGGTACCTCCTGGAGCGAATCAGGGATACGGCTGGGCAGAAGGGTACAGGAAAATGGACAGCGATCGCCGCTTTAGATTACGGAATTCCC
GTAACTCTAATTGGTGAATCGGTATTCGCCAGGTGCCTTTCGTCGTTGCAGAGCGAAAGAATAGAGGCCAGTGTCGTTTTGGA------------------
CGGACCGCACGCAACTTACCAGGGCGATAAGAAACAGTTCCTCGAACATCTAGGGAAAGCACTCTACGCT?????????????????????????????????????????????????????????????????????????????????????????????
AGCAAGAACGAAGTCGACGTACTCGTCGCGTTTCTGAAAGCCGGCCCGCTCGATCCCAACGTCGGTAAGTTTTCAGAGGTCGTGGTCGGAGTACCCGCTATATACTTGACTTACGCGAAAAGTATTTTGCCCGGTAATGTGCAAGTTAGCGGTCAGAATACTTACAAAGTAGCGAAGGGTGCTTTCACCGGCGAGCTCAGTCCCGCCATGCTTCTGGACAACGA
CATCCCTTGGGTGATCCTCGGGCACTCCGAGCGTCGCAACGTGTTTGGCGAATCGGACGACTTGATTTCCGAGAAAACAAGCCACGCTCTCGAAGCCGGACTAAAGGTAGTCATCGCTTGCATCGGCGAGAAGCTAGACGAGCGCGAGGCTGGCAAGACAGAAGAGGTCGTCTTTAGGCAGACCAAAGCTATCGCGGACAAAATTAAATCCTGGGACAACGTCG
TACTG???????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????
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ACAGTTTACAGAAAAACTTCAAATCCTTATTCGGCGAGAAACTGGAGGTTGTTAGAACGCACCAGCAACAAGAAAACCTCAAATTCATGGCACATTTTAAGCGTAAATTTATTATTCATCAAGGAAGACGCAAGCAGCCAAAGTCACCCGCCACTAACAAAGTCGAATTTTACCATCTTCGAAGCAACGGCAGCGCCTTGTGTACTAGGCTGATACAAGTGCAG
CCAGAAGCCACTTTATTGAATTCCGCGTTTTGGTACAGCTACATTTTAAATGTGCCATTTAATAACGACGACGAGACTGGAATCGTCTACGTTTGGGTCGGATCGAAAGCCGACAGCGAAGAGGCGAGGCTCGTCGAAGAAATGGCAGAAGAAATGTTCAACAACGTACCGTGGATAAGTCTACAAGTATTAAACGAGGGCGAGGAACCGGACAATTTCTTCTG
GGTCGGACTCGGCGGTAAGAAACCGTACGACACCACGGCCGAGTACATGAACTACACCAGGCTATTCCGATGCTCCAACGAAAAAGGATACTTCACGATTAGCGAGAAGTGCACAGATTTCTGCCAGGTAGACGATCTGGCCGATGACGATATAATGATACTGGATAACGGCGAACAAGTGTTCCTGTGGCTGGGTGCCAGATGTTCGGAAGTCGAGATTAAGC
TCGCGTACAAATCGGCTCAGGTACGGGTGTATATACAACACTTACGAGTGAAGCAACCAGAGAAACCCCGAAAAT?????????????????????????????????????????
Siobla_spp ??????????????????????????????????????????????????????ACTGTCTCTT-TTTAATTA-TTAATTTAATTTAATTTTTAAGTTAAAAAGCTTAAATTTTATTAAAGGACGAGAAGACCCTATAGAGTTTTATA-C---TT-T--TAT--A--TTTA-TA--AAATTTAA-TT--
AA-AT-TT-TA-T-AAA-T-A-TAAAAGTATTTTATTGGGGCGATAAAAAAATTTATTAAACTTTTTTAT-TA-T-------T-ATTA-CATTGATGAGTGAA-T-T----ATTGATCCTTT----ATT------------------------------------------------------------------------------
T?????????????????????????????????????????????????????????????????????????????????TTGAAAAGAACTTTGAAGAGAGAGTTCAAGAGTACGTGAAACCGTTCAGGGGTAAACCTGAGAAACCCGAAAGATCGA-ACGGGGAGATTCATCGTCAGCGGGGTTGGCTTCTC-CGCGG-CTCGTGATG---TCG-G-G--
A-C-C-A--C----GCGT-CCACGGC-ACTCGGCCGCGG-TGTT--C-ACGTCCGACCACGCCGGCGTGCACTTCTCCCCTAGTAGGACGTCGCGACCCGTTGGGTGTCGGTCTACGGCCCGGGT----GGACGACTGT-C--TCGGCGT--TCGC--GT-C-----GGGGCAGACC-C-C-CGGT-TGCCCGTCCGGCTGCCCGGCGGTACACGCAC-
GGTATAGAGCCGCAT-T-GAA-CTGCGTCGGGCCCGCCGCAAGCGCGGTCAGCGAT----TCCCGGTGGTCGGAC-CTAGCGCCGTCCCCGGGCCTGATCAGCTGTTGGCAGGCGGTGTCCTCTGACAGGCTC-ATT---T-GAAA-T-----ATTAT-
TATACCGGTCGGCGACGCTATTGCTTTGGGTACTTTCAGGACC????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????
CCTCGACTAAATAATATAAGATTTTGATTTCTCCCACCTTCTATTACTCTATTACTTTCAAGAAGAATAGTAAATTCTGGATCAGGAACAGGATGAACTGTATATCCACCATTATCAAGAAACATATCACATGCTGGAGCATCAGTAGATTTAACAATTTTCTCATTACACCTCGCAGGAATTTCATCAATTTTAGGAGCTATTAACTTCATTACAACTATAAT
TAATATACGAACATCAGGAATAACTTTAGAACGAATACCTTTATTTATTTGAGCAGTATCATTAACTGCTTTACTATTACTTTTATCATTACCAGTTTTAGCTGGAGCTATTACAATATTATTAACAGATCGAAATTTAAATACATCATTTTTTGATCCTTCAGGAGGAGGAGATCCTATTTTATATCAACATTTATTTTGATTTTTTGGACATCCCGAAGTTT
ATATTCTAATTATTCCAGCATTTGGAATAATTTCTCATATTATTTCACAAGAATCAGGAAAAAAAGAAACATTTGGAACATTAGGAATAATTTATGCTATACTAACTATTGGATTATTAGGATTTGTTGTATGAGCTCACCATATATTTACTGTTGGTATAGATGTTGATACACGAGCTTATTTTACAGCTGCTACCATAATTATTGCTATTCCTACAGGAATT
AAAATTTTTAGATGATTAGCAACATTATATGGGTCTCAATTAATTTTTAATCCTTCAATAATATGAACTTTAGGATTTGTATTCCTATTCACTATAGGAGGATTAACAGGAATTATCTTATCTAATTCCTCAATTGAT?????????????????????????
GACTCTATTAAGGAGAAATTGATCCTCCCGTTCCTAGAGATCGAGCTGCACATCTACGACCTTGGTATGGAAAGCCGAGACGCGACCAATGACCAAGTGACCGTCGATTGCGCGGAAGCTGTAAAGAAATACAACGTAGGAATAAAATGCGCGACTATAACTCCCGACGAGAATAGAGTCGAGGAGTTCAAGCTAAAAGAAATGTGGAAGAGTCCCAACGGCAC
CATCAGAAACATCCKGGGTGGTACCGTCTTCAGGGAGGCAATAATCTGCAAAAATATTCCGCGACTGGTCACTGGCTGGAACCAGCCGATCATCATAGGCCGTCACGCTCATGCCGATCAATACAAAGCTACCGACTTTGTTGTCCCTGGGGCTGGTAAACTGGAGATTACTTGGACAGGGGAATCTGGCGATAAGATAAATCACACCGTGTACCAGTTCAAGG
GGGCAGGGGTTGCTCAGGCTCAGTTCAATACGGATGAAAGTATCGAGGCCTTTGCACACAGCTCTTTCCAGTACGCCTTGTCTCGCGCCTATCCCCTTTATCTCTCCACCAAGAACACTATCTTGAAGAAATATGACGGTAGGTTCAAGGATATCTTCCAAGCTATCTACGAAGCTGAGTACAAGGCAAAGTTCGAAGCAAAGAAGATCTGGTACGAGCATCGT
CTGATCGACGATATGGTCGCCTATGCCATGAAATCGGAGGACATGCAGTTGATTTGCGAGGCTTATCACATAATGAGAAACGGATTAGGGCTAAGCCCCAAAGAAATGAGCGATGTTTTCGGGGAGTGGAATAAGGGGGTGCTGGATTCATTCCTGATTGAAATAACCAGAGACATATTGAAGTACAAAGACGAGAAGGGATATCTGTTGGAAAGGATCAGAGA
TACAGCCGGACAAAAGGGTACAGGAAAGTGGACAGCCATAGCTGCCCTAGATTACGGAATTCCCGTAACTCTAATCGGGGAATCGGTCTTTGCCAGATGTCTCTCGTCGTTGCAGAGTGAGCGGATAGAAGCGAGCAGCGTTTTGGA------------------
AGGACCAAATGCTCTCTATCAAGGCGACAAAAAGCAGTTCCTTGAACATCTCAGGAAAGCCCTTTACGTGTCGAAGATCATTTCATATGCTCAGGGATTTATGCTGCTGAGAGAAGCGGCCAAAATTCATAAATGGAATCTGAATTACGGCGGTATCGCACTCGTGAAAAAGGAAATTGACGAAATCGTCGCATTCCTGAAAGCCGGTCCTCTCGATCCAAATG
TCGGTACGTTTACAGAAATCGTTGTCGGAGTACCAGCTCTTTACCTGACCTATGCCAAAAGTATATTGCCACCGAACGTACAAGTCAGCGCTCAGAATAGCTACAAAGCAGCCAAAGGTGCTTTCACTGGCGAAATTAGCCCTGCTATGCTTTTAGACAATGAAATTCCTTGGGTAATCCTCGGGCATTCGGAACGTCGTAACGTATTCGGCGAATCTGATGAA
CTCATTTCTGAAAAAATCGCCCACGCCCTTGAGGCTGGTTTGAAGGTAGTGATTGCCTGCATCGGAGAGAAGCTGGATGAGCGTGAGGCTGGAAAAACTGAGGAAGTAGTATTCAGACAGACTAAAGCAATCGCGGACAAGATAAAGTCATGGGACAATGTTGTGCTGTTACAGATTGGAAGTTCGATGAAGAGCGTAGGGGAGGCAATGGCGATCGGTAGAAA
GTTTGAAGAGGCTTTTCAGAAGGCGCTTAGGATGGTCGATGAAAATGTGAACGGTTTTGATCCCTACATAAAACCAATCGACGACGAAGACCTGGAGAGGCCTACCGACAAACGAATGTTCGTACTGGCAGCTGCCCTCAAGGCTGGCTACACGATCGATCGCTTGTACGAATTGACTAAAATAGATCGTTGGTTTTTGGAAAAGATGAGAAACATCACCTCGT
ATTATACCCTCCTCGAAGATCTCGACCAGACCAAACTCTCGCACGAAGTATTGCTCCGTGCCAAACAAATCGGTTTCTCTGACAAGCAAATAGCGACTGCGGTAAAAAGCACCGAACTTGCCGTACGCAAACAAAGACAAGAAAGCAACATCAGACCCTTCGTAAAACAGATAGATACTGTCGCTGCCGAGTGGCCGGCCACCACGAACTACCTCTACCTCACC
TACAACGGAAACAGTCACGATGTTCAATTCCCTGGAGGCTACACCATGGTTATAGGTTCGGTAGGCTCGGGTGTATATCGCATAGGAAGTTCCGTGGAGTTTGACTGGTGTGCGGTAGGATGTCTTAGAGAATTACGGCGATTGAACCGTAAGACGATAATGGTAAACTACAACCCGGAGACAGTCAGTACGGATTATGACATGTGCGATCGTTTATACTTCGA
GGAAATTTCCTTCGAGGTCGTGATGGACATTTACGATTTTGAGAATCCGGAGGGG??????????????????????????????????????????????
TTAGTTTACAGAAAAAGTTCAAATCCTTATTCGGCGAGAAGCTTGAGGTGGTCAGAACACATCAACAACAAGAAAATCTAAAATTCATGGCCCATTTCAAGCGCAAGTTTATCATCCGTCACGGAAGACGCAAGCAGCCAAAGTCACCCGCTAATAACAAAGTCGAATTTTATCATCTTCGAAGCAATGGTAGCGCTTTGTGTACAAGACTAATTCAAGTGAAT
CCAGATGCCTGTTTGTTGAATTCTACCTTTTGGCGCAGCTACATATTGAATGTACCATTCAACAACGACGACGAAACTGGGATAGTTTATGTTTGGATCGGATCTAAAGCAGACAGCGAAGAAGCGAGACTCGCGGAAGAGATAGCGGAGGAAATGTTCAACAACGTACCTTGGATCAGTCTGCAAGTATTAAACGAGGGCGAAGAACCTGACAACTTTTTCTG
GGTAGGAATCGGAGGGAAAAAACCCTACGATACCAACGCCGAGTACATGAACTACACCAGACTATTTAGGTGCTCTAACGAAAAGGGGTACTTCACGATAAGTGAAAAATGCACAGACTTTTGCCAGGTAGATGATTTAGCCGACGATGATATTATGGTGCTAGACAACGGCGAGCAAGTCTTTCTCTGGCTCGGTGCCCGATGTTCAGAAGTGGAGATTAAGC
TTGCGTTTAAATCGGCTCAGGTAATAGTCTACATTCAGCACCTGCGAGTTAAGCAGCCAGAACGACCTAGAAAATTATTCCTCACAGCTAAAAGCAAGGAGT??????????????
Waldheimia_amazonica ATAATAATTAGTCTTTTAATTAGAGACTGGAATGAAAGATTGAATGAAATATAAACTGTCTTAA-AATTATTA-AAGATTTAATTTAATTTTTAAGTTAAAAAGCTTAAATAACTTTAAAGGACGAGAAGACCCTATAGAGTTTTATA-G---TTTT--TTT--ATTAAAAT-A--GGTTTAAA-TT--
TT-CA-TATT--T-T-A-T-AAGAGGATTATTTTGTTGGGGCGACAGGAAAATTAGATTAACTTTTTTAT-TA-T-------A-TAAA-CACTTATTTGTGAG-T-T----TTTGATCCTTAG--T--T-A-AAGATTATTAGATTAAATTACCTTAGGGATAACAGCGTTATTTTTTTT-
AGAGTTCTTATCGATATAAAAGATTGCCAGCTCTAAGTGGGTGGTAAACTCCATCTAAGGCTAAATATGACCACGAGACCGATAGCGAACAAGTACCGTGAGGGAAAGTTGAAAAGAACTTTGAAGAGAGAGTTCAAGAGTACGTGAAACCGTTCAGGGGTAAACCTGAGAAACCCGAAAGATCGA-ACGGGGAGATTCATCGTCAGCGGAGTAGGCTTCTC-
CGTGG-TTCGTGATG---TCG-G-G--A-C-C-C--C----GTGT-CCACGGC-ACACGGCCGCGG-TGTT--T-ATGCCCAACGACGCCGGCGTGCACTTCTCCCCTAGTAGGACGTCGCGACCCGTTGGGTGTCGTACTACGGCCCGTTG----GGCTCTCTGT-C--TCGGCGT--CCGC--GT-C-----GGGGCCGATC-T-CACGTT-
GGTCCGTACGGCTGCCCGGCGGTACACACAC-GGTATCGAGCCGCAT-T-GAA-TTGCGTCGGGCCCGTCGCAAGCGCGGTCAGCGTT----TCCCGGTGGTCGGAC-TCAGCGCCGTCCCTGGGCCTGGCCAGCTGTTGGCTGACGGTGTCCTCTGACCGGCTC-ATT---C-GAAT-TATTTTTTTCA-
TATACCGGTCGGCGACGCTATTGCTTTGGGTACTTTCAGGACC???????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????
TAATGGGGCCCCCGATATGGCTTTCCCACGTCTTAATAATATAAGATTCTGATTTCTACCCCCATCATTAATATTACTAATATTAAGAAGAATAATCAATTCAGGTTCAGGAACAGGGTGAACTGTATACCCCCCACTATCAAGAAATATAGCTCATGCAGGACCATCAGTAGATTTAACAATTTTTTCCCTACATTTAGCAGGAATTTCATCAATCCTAGGGG
CAATTAACTTCATCTCTACTATATTAAATATAAATGTCAAAGGAATAAGATTAGAACGATTACCTCTTTTTGTTTGAGCTGTCTCTTTAACTGCAATTTTACTATTACTATCCCTACCAGTACTAGCAGGAGCCATCACAATACTATTAACTGACCGAAATATTAATACATCTTTCTTTGACCCTACAGGAGGAGGTGACCCTATTCTATATCAACACCTATTT
TGATTTTTTGGTCACCCTGAAGTATACATTTTAATTATCCCAGCCTTCGGAATAATCTCTCACATTATTTCAAATGAATCCGGCAAAAAAGAAACCTTTGGTGTCTTAGGGATAATCTACGCAATATCAACAATTGGAATCCTAGGTTTTGTAGTCTGAGCCCATCATATATTTACTGTAGGAATAGATGTAGATACACGAGCTTACTTTACTGCAGCAACTAT
AATTATTGCTATTCCTACAGGAATTAAAGTTTTCAGATGGTTAGCAACTATCTATGGATCAAAAATTAACTATAGACCCTCAATATTATGAGCATTAGGATTTGTATTCTTATTTACTGTAGGAGGATTAACCGGAATTTTATTATCAAATTCTTCAATTGATATTATTCTCCATGATACATACTACG????????????????????????????????????
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AGGCGATGGCGATTGGCAGGAAGTTCGAGGAGGCTTTCCAGAAGGCTCTGAGGATGGTCGACGAGAATGTTAACGGCTTCGACCCCTACATCAAGTCCATAGACGACGAGGAACTGGAGAAACCAACTGACAAGCGGATGTTCGTGTTGGCGGCGGCCCTGAAGGCTGGCTACACGATCGACCGCTTGTACGAACTGACAAAAATCGATCGGTGGTTTTTGGAA
AAAATGCGAAACATCACGTCGTACTACACCGTTCTCGAGGATCTGGATCAGACCAAGCTTTCCCACGAGGTTCTGCTCCGCGCCAAGCAAATCGGTTTCTCTGACAAGCAAATCGCCACTGCTGTGAAAAGCACCGAACTGGCTGTCCGCAATCAGAGACAGGAAAGCAACATCAAACCCTGCGTCAAACAAATCGATACCGTCGCCGCCGAGTGGCCGGCTAC
GACGAACTATCTCTATCTCACTTACAACGGAAATAGCCACGATGTCCTGTTCCCTGGGGGATACGCAATGGTTATCGGTTGGCCAGGCTCCGGCGTTTACCGGATCGGCAGTTCCGTGGAGTTCGATTGGTGCGCCGTGGGATGTCTCAGAGAATTACGGAGACTGAATCGGAAGACAATCATGGTCAATTACAACCCCGAGACGGTCAGCACGGACTACGACA
TGTGCGATCGTTTGTACTTCGAGGAGATATCCTTCGAGGTGGTGATGGATATTTACGATCTCGAGAACCCTGAAGGT???????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????
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Zenarge_turneri ??????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????
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ATAATTATTCGTACAGAATTATGTTCAACAGGCTCTTTAATTGGAGATGATCAAATTTATAACACAATTGTTACATCACATGCATTTTTAATAATTTTTTTTATAGTTATACCAATTATAATTGGAGGGTTTGGAAATTGATTAATTCCCTTAATATTAGGAGCCCCAGACATAGCTTTCCCTCGATTAAATAATATAAGATTTTGATTCTTACCCCCCTCTTT
ATTTATATTAACAATAAGAAGATTAATTGATTCAGGAACAGGAACTGGGTGAACTATTTACCCTCCCCTTTCAAATAATATTGCTCATGCTGGAGCATCTGTAGATTTAACTATTTTTTCTTTACATCTAGCAGGTATTTCATCAATCCTTGGAGCAATTAATTTTATTTCCACAGTTATTAATATACGAACTATAGGAATAAATTTTGATAAAATACCTTTAT
TAATTTGAGCCGTATCAATTACTGCAATTCTTTTACTTCTATCCTTACCAGTTTTAGCAGGAGCAATTACTATATTATTAACTGACCGAAATTTAAATACATCATTCTTTGATCCTTCAGGAGGAGGAGACCCAATTTTATACCAACATTTATTTTGATTTTTTGGTCATCCAGAAGTTTACATTTTAATTCTTCCAGCATTTGGAATTATTTCTCATATTATT
TCTCAAGAATCAGGAAAAAAAGAAACCTTTGGAACCCTAGGAATAATTTATGCAATATCAGCTATTGGACTCCTTGGGTTTATTGTTTGAGCCCATCATATATTCACAATTGGAATAGATGTTGATACACGAGCTTATTTTACTGCAGCAACAATAATCATTGCGGTACCCACAGGAATTAAAATTTTTAGATGATTAGCAACAATTCATGGATCCAAATTAAT
TTATACTCCTAGAATACTATGAACGTTAGGTTTTGTATTCTTATTTACTATTGGAGGATTAACAGGAATTATCCTAGCTAATTCTTCTATTGACATTATTCTTCATGATACTTATTACGGACTCGATAAAGGAAAAGCTGATTCTTCCCTTCCTCGACATCGACCTGCACATCTACGATCTTGGAATGGAGAACCGAGACGCGACGAACGATCAAGTTACGATC
GACTGCGCGGAAGCGGTTAAGAAGTACAACGTTGGTATAAAGTGCGCGACGATAACTCCGGACGAGAAACGAGTGGAAGAGTTCAAGCTGAAGCAGATGTGGAAGAGTCCTAACGGCACGATCCGTAACATCCTCGGCGGGACCGTCTTCAGAGAGGCGATAATTTGCAAGAACATTCCGCGCCTCGTAACTGGGTGGAACCAACCGATCATAATCGGTCGTCA
CGCGCACGCGGATCAATACAAAGCGACCGACTTCCTCGTCCCGGGCGCAGGCAAGCTCGAGATAACGTGGCGCGGAGTCGACGGCAAGAAGATCGAACACACCGTCTACGAGTTCAAAGGCGCCGGCGTGGCGCAAGCACAGTTCAATACGGACGAGAGTATCGAGGCCTTCGCGCACAGCTCGTTCCAGTACGCGCTATCGAGGACCTATCCGCTTTACCTGT
CCACGAAAAACACGATCCTCAAGAAGTACGACGGCAGATTCAAGGACATTTTCCAGGAGATCTACGAGAAAGAATACCGCAAGCAGTTCGAGGCGAAGAACATCTGGTACGAGCATCGC???????????????????????????????????????
GACATGCAGTTGATTTGCGAAGCCTATCACATCATGCGAGATGGATTAGGTCTCAGTCCTCAGGAGATGAGCGACGTTTTCGGCGAATGGAACAAGGGTGTCCTCGACTCGTTCCTAATCGAAATCACTAGAGATATCCTGAAGTACAAGGACGACAAGGGTTACCTACTGGAACGAATCCGCGACACAGCTGGACAGAAGGGCACAGGAAAATGGACGGCGAT
CGCCGCCTTGGACTACGGAATTCCCGTAACTCTAATTGGAGAATCAGTATTTGCCAGGTGTCTTTCCGCGCTGCAGAGCGAAAGGATAGAAGCCAGCACAGTTTTGAC------------------
CGGGCCAAACACTCGCTACCAGGGTGACAAGAAACAATTCCTCGAGCATCTTAGAAAGGCTCTCTACATTTCGAAAATTATATCTTACGCCCAAGGATTCATGCTTCTGCGCGAGGCCGCAAAAATACACAAATGGAATCTGAACTACGGTGGCATCGCGCTCAGTAAGAAGGAAATCGACGAAATTGTCGCTTTCCTGAAGGCCGGTCCTCTGGATCCAAATG
TTGGTAGGCTTGTAGAGATTGTTGTCGGAGTACCGGCTATCTACTTGAATTATGCCAAGAGTATTTTGCCACCGAACGTGCAAGTCAGCGGCCAAAACAGCTACAAAGTAGCAAAGGGCGCTTTCACCGGCGAGATCAGTCCCGCCATGCTTTTGGACAACGGAGTTTCCTGGGTGATCTTAGGCCACTCGGAACGACGCAACGTATTCGGAGAATCTGATGAC
CTGATCTCAGATAAAATCAGCCACGCGCTGGAAGCTGGTTTGAAGGTGGTGATTGCTTGCATCGGAGAAAAGTTGGACGAACGCGAAGCCGGGAAAACGGAGGAGGTCGTTTTCAGGCAGACCAAAGCTATTGCGGACAAAATTAAGTCCTGGGACAACGTTGTGCTG????????????????????????????????????????????????????????
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AAACAAATCGGATTCTCCGACAAACAGATCGCAAAGGCTGTTAAGAGCACGGAATTGGCTGTTCGCAAGCAACGCCGAGAGAGTAATATACGGCCCTTCGTCAAGCAAATTGATACTGTCGCTGCAGAATGGCCAGCGACGACAAACTACCTCTACCTGACTTACAACGGAAACTGCTACGACGTGCAATTTCCCGGTGGCTACACGATGGTTATAGGTAAGCC
AGGCTCCGGTGTATATAGGATAGGAAGCTCCGTGGAATTTGACTGGTGCGCCGTTGGCTGTATCAGAGAATTACGTCGACTGGGTCGTAAGACTATAATGGTCAATTATAACCCGGAGACTGTCAGTACCGACTATGATATGTGCGATCGTCTCTACTTCGAAGAAATATCGTT?
GAGGTGGT????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????
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TCGTCGAAGAAATAGCTGAAGACATGTTTAACAATGTTCCATGGATCAGTCTGCAGGTATTAAACGAGGGTGAAGAACCCGATAACTTTTTCTGGGTTGGAATCGGAGGAAAGAAACCCTACGATACCGATGCGGAGTACATGAATTATACTAGGCTTTTCCGATGCTCCAATGAGAAGGGGTACTTTACAATTAGTGAAAAATGTACAGATTTCTGTCAGGTA
GATGATTTGGCTGACGATGATATTATGGTGCTCGACAACGGCGAGCAAGTATTTCTATGGCTAGGTGCCAGGTGTTCGGAAGTGGAAATTAAGCTAGCATACAAATCGGCTCAAGTACGGGTGTACATACAGCATTTGCGCGTGAAGCAACCAGAAAGACCACGCAAACTCTTCCTTACGGCCAAAAGCAAGGAATCCCGAAGATTC???

Acordulecera_spp ??????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????
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CAGCTCTAAGTGGGTGGTAAACTCCATCTAAGGCTAAATATGACCACGAGACCGATAGCGAACAAGTACCGTGAGGGAAAGTTGAAAAGAACTTTGAAGAGAGAGTTCAAGAGTACGTGAAACCGTTCAGGGGTAAACCTGAGAAACCCGAAAGATCGA-ACGGGGAGATTCATCGTCAACGATTACGGCTTCGC-CGAGG-CTCGTGATG---CTGTG-G--
GC--C-T--C----GTGCT-CGCGGC-ACGCGGTCGCGC--GCG--TCATGTCCGGCGTCGTCGGCGTGCACTTCTCCCCCAGTAGGACGTCGCGACCCGTTGGGCGTCGGCGTAAGGCCCGTT---C-GGTAGCCTGTC---TCGGCGC----TCGCGT-C-G---G-GGCAGACC-C-G-CGGT-TGCCCGGCCGGCTGCCCGGCGGTACTCGCAC-
GGTATCGAGCCGCAT-CGAAC-ATGCGTCCAGCTCGTCGCAAGCGCGGTCAGCGAT----TCCCGGTGGTCGGAC-CTTGTGCCGTCCCCGGGCCTGGCCAGCTGTTGGCAGACGATGCCCTCGGACCGGCTC-GCT---C-G--------------T-
TTTACCGGTCGGCGACGCTACTGCTTTGGGTACTTTCAGGACC?????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????
ACTAATATTAGGAGCACCTGATATAGCATTCCCACGAATAAATAACCTAAGATTTTGACTTCTCCCACCATCATTAATTTTATTAACATTCAGAAGATTTATTGACTCAGGGTCAGGAACAGGATGAACAGTTTATCCACCACTATCAAGAAACATTGCTCATGCAGGATCATCTGTTGATATAACTATCTTTTCACTTCATATAGCAGGAATCTCCTCAATTT
TAGGTGCAATTAACTTTATTTCAACAATAATTAATATACGAACCCATGGAATAAGATTTGATAAAATACCATTATTAATTTGAGCAATTATAATTACAGCTATTCTTTTAGTTATTTCATTACCAGTATTAGCAGGGGCTATTACTATATTATTAACAGATCGAAACTTAAATACATCATTTTTTGACCCATCTGGAGGAGGAGATCCTATCTTATATCAACAT
TTATTTTGATTTTTTGGACACCCAGAAGTTTACATTTTAATTTTACCAGCCTTTGGAATTATCTCTCATATTATTTCTCAAGAATCAGGTAAAAAAGAAACATTTGGAACACTAGGAATAATTTATGCAATATCAGCAATTGGATTACTAGGATTTGTAGTATGAGCTCATCATATATTTACTGTAGGAATAGATGTAGATACTCGAGCCTACTTTACTGCAGC
AACTATAATCATTGCTGTACCAACAGGAATTAAAATTTTTAGTTGATTAGCAACAATTCACGGATCAAAAATAATTTACACGCCTAATATATTATGAACCCTAGGATTTGTATTTCTATTCACTATAGGAGGATTAACAGGAATTATCTTATCAAATTCATCAATTGATATTATCATACATGATACTTACTACG??????????????????????????????
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CGATCAAGAAATACAACGTCGGCATAAAGTGCGCGACGATAACTCCGGATGAGAATCGAGTGGRGGAATTCAAGTTGAAACAGATGTGGAAGAGTCCGAACGGGACGATCCGAAACATCTTGGGCGGTACAGTGTTCCGCGAGGCTATAATCTGCAAAAATATTCCCCGCTTGGTGACCGGCTGGAACAAGCCGATAATAATTGGTCGCCACGCATACGGAGAC
CAGTATAGAGCGACAGATTTCCTAGTTCCCGGCCCCGGCACGCTGGAAATCAGTTGGGTAGGAAAAGATGGACAAAAAATCCAGCACACCGTCAACGAATTCAAGGGACCTGGCATCGCGCAGGCGCAGTTCAACACCGACGAGAGTATCAAGGACTTTGCGCACAGTTCGTTCCAGTACGCGCTTGCTCGAGAGGTGCCTTTGTATCTGTCGACGAAGAACAC
TATTCTCAAGAAGTACGACGGCGGATTCAAGGACATTTTCCAAGAGATATACGAGAAGCAGTACAAGTCCAAGTACGAAGCGGCGAACATTTGGTACGAGCACCGTCGAATCGACGATATGGTTGCCTATGCCATGAAGTCCGAGGATATGCAGTTGATCTGCGAGGCGTATCACATAATGCGGGATGGCTTAGGTTTGAATCCGCAAGAAATGAGCGACGTCT
TCGACGAATGGAACAAAGGAGTCCTGGATTCTTTCCTGATCGAGATCACCAGAGACATTTTGAAGTACAAGGACGAGAAAGGCTATCTCCTGGAGCGAATTAGGGACACGGCAGGGCAAAAAGGCACTGGGAAATGGACTGCGATCTCTGCGTTGGATTACGGTATCCCTGTCACGCTAATCGGCGAATCAGTCTTCTCGAGGTGCTTGTCCGCACTCCAAGCC
GAAAGAGTCGAGGCGAGCACCGTCCTCAC------------------
CGGACCCAGCGCTCTCTATCAAGGAAACAAAAAACAGTTCCTCGAACATCTCAGGAAAGCTTTATACCTGTCGAAAATCATTTCCTACGCCCAAGGGTTCATGCTGCTCCGAGAAGCCGCCAAAATTCATAACTGGAATCTGAATTATGGAGGTATAGCACTCAGCAAGAAAGAAATCGACGAGATCGTTGCATTTCTGAAAACTGGCCCCCTAGATCCSAACG
TTGGTACGTTTTCAGAGATTGTCGTTGGAGTACCAGCGATCTACTTGACYTACGCCAAGAGCACTTTGCCAGCAAACGTGCAGGTGAGCGGCCAGAATACTTATAAAGTAGCCAAGGGCGCTTTCACCGGCGAACTRAGTCCGGCTATGCTTTTGGACAACGACGTGCACTGGGTAATTCTTGGTCATTCTGAGCGTCGCAACGTTTTCGGAGAATCCGACGAA
TTGATCGCCGATAAAGTGAGCCACGCGCTCGAAGCTGGCTTGAAGGTAGTGATCGCTTGCATCGGCGAGAAATTGGAGGAACGCGAGGCTGGCAAGACAGAGGAAGTAGTCTACAGGCAGACTAAAGCTATCGCGGACAAAATCAAGTCCTGGGACAACGTCGTATTATTGCAGATCGGTAGTTCGATGAAGAGCGTCGGCGAAGTGATGGCGATCGGGCGCAG
GTTCGAGGAGGCGTTTCAAAAAGCGCTTAGAATGGTAGATGAGAACGTGAACGGATTCGACCCATACATCAAGTCCATCGACGACGAGGAGCTGGAGAGGCCAACCGACAAGCGGATGTTCGTYTTAGCTGCAGCCCTCAAGGCTGGCTACACGACAGAACGACTGTACGAACTCACCAAGATCGATAGATGGTTCTTGGAGAAGATGAGGAACATCACCGCTT
ACTACACCCTACTCGAAGGACTCGAGCAAACCAAGCTCTCCTACGAAGTCCTTCTGCGTGCCAAGCAGATCGGCTTTTCGGATAAACAGATCGCGAAGGCTGTTAAGAGCACGGAACTAGCTGTACGCAAACAACGCCAGGAGAGCAGTATCCGACCTATTATCAAACAGATCGACACTGTCGCTGCTGAATGGCCAGCAACGACCAACTATCTTTACCTAACC
TATAACGGAGACCGCCACGATATTCAATTCCCTGGCGGATACACCATGGTCATTGGTACG????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????
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Anathulea_sp ??????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????
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CAGCTCTAAGTGGGTGGTAAACTCCATCTAAGGCTAAATATGACCACGAGACCGATAGCGAACAAGTACCGTGAGGGAAAGTTGAAAAGAACTTTGAAGAGAGAGTTCAAGAGTACGTGAAACCGTTCAGGGGTAAACCTGAGAAACCCGAAAGATCGA-ACGGGGAGATTCATCGTCAGCGGCTGCGGCTTCGC-CGGGG-CCCGTGATG---TCGCG-G--
AC--C-T--C----GCGTT-CGCGGC-ACACGGTCTCGC--GCG--TCATGTCCGTGGTCGTCGGCGTGCACTTCTCCCCCAGTAGGACGTCGCGACCCGTTGAGCGTCGGCGTAAGGCTCGAT---T-GGTAGACTGTC---TCGGCGT----TCGCGC-C-G---G-GGCAGACC-A-T-CGGT-TGCCCGGCCGGCTGCTCGGCGGTACTCGCAC-
GGTATCGAGCCGCAT-CGAAC-ATGCGTCCAGCCCGTCGCAAGCGCGGTCAGTGAT----TCCCGGTGGTCGGAC-CTAGTGCCGTCCCCGGGCCTGGCCAGCTGTTGGCAGACGGTGTCCTCGGACCGGCTC-GCT---C-G--------------T-
TTTACCGGTCGGCGACGCTATTGCTTTGGGTACTTTCAGGACC?????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????
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GATATGCAGTTGATCTGCGAGGCGTATCACATAATGCGGGATGGTCTAGGTCTGAATCCACAAGAAATGAGCGACGTCTTCGACGAATGGAACAAAGGAGTGCTCGATTCATTCCTCATCGAGATCACTAGAGATATTTTGAAGTACAAGGACGACAAAGGGTATCTTCTGGAGAGAATTCGAGACACAGCGGGACAAAAGGGCACTGGCAAATGGACTGCCAT
ATCTGCATTGGACTACGGAATCCCTGTCACTCTAATTGGCGAATCAGTTTTCTCTAGGTGCTTGTCCGCGCTTCAAACGGAACGAGTCGAAGCAAGCGCTGTCCTTAC------------------
TGGGCCGAACGCTCTCTATCAAGGAAACAAAAAACAATTCCTTGAACATCTCAAGAAAGCTTTATACCTGTCGAAGATCATATCCTATGCTCAAGGGTTCATGCTTCTCCGAGAAGCAGCCAAAATTCACAATTGGAATTTGAATTACGGAGGTATCGCTCTG?????????????????????????????????????????????????????????????
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 Aulacomerus_spp ATAATAATTTGTCTTTTAATTGGAGGCTTGAATGAAAGATTGGACGAAATAAAATCTGTCTCA-TAAAAATT--TATTTTGAATTTAATTTTTAAGTTAAAAAGCTTAAATTTTATTAAAGGACGAGAAGACCCTATAGAATTTTATA-A---TT-A----A--AAATTTAT-T--TAATAAAA-TTT--
T-TT-TTTA--A-AGT-T-T-TTAAATTATTTTGTTGGGGCGACAGTAAAATTAATTAAACTTTTATAA-AT-AATAAAAAA-TTAA-CATTAATTAATGAA-T-ATATTTATGATCCTGTA--A--T-A-
TAGATTATTAGAATAAGTTACCTTAGGGATAACAGCGTTATTTTTTTTTAGAGTTCTTATTAATAAAAAAGATTGC??????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????
CCCGAAAGATCGA-ACGGGGAGATTCATCGCAAGCGGGGTCGGCTTCGC-CGCGG-CCCGTGATG--C-CGCG-G--AC-C-T---C--GC--GCC-CGTGGC-ACGCGGCCGCG-TCCC-GC--ATGCCCGACCTCGTAGGCGTGCACTTCTCCTCCAGTAGGACGTCGCGACCCGTCGGGCGTCGGCATGAGGCCCGAATG---TGTCGACTGT-AC--
CGGCCCT--A--GAG------CCGGTGCAGACCTT---CGG-CCGCCCGGCCGGCTGCCCGGCGGTACACGCAC-GGTATGGAGCCGCA-CA-GAACTTGCGTCCAGCCCGTCGCAAGCGCGGCCAGCGCAT---CACCGGTGGTCGGAC-AAAGCGCCGTCCCCGGGCCTGGCCAGCTGTTGGCAGACGGTGACCTCGGACCGGCTC-GCG------
G----------------ATACCGGTCGGCGACGCTACCGCTTTGGGTACTTTC-----
AATAATTATTCGAACTGAATTAAGTCATCCAGGATCTTTTATTGGAGATGACCAAATTTATAATGTATTAGTTACATCTCATGCATTTTTAATAATTTTTTTTATAGTTATACCAATAATAATAGGAGGATTTGGTAATTGATTAATTCCCCTAATATTAGGAGCTCCTGATATAGCATTTCCTCGAATGAATAATTTAAGATTTTGATTTTTACCCCCATCAT
TAATTTTATTAACTTTCAGAAGATTTGTTGATTCAGGATCAGGAACTGGATGAACTGTTTACCCCCCATTAGCAAGAAATATTGCCCATAGAGGGACATCTGTAGATTTAACTATTTTTTCTCTTCATATAGCAGGAATTTCTTCTATTTTAGGAGCAATTAATTTTATTTCTACATTAATCAATATACGAACTTTAGGGATAAAATTTGATAAAATACCTCTT
ATAATTTGAGCAATTAACATTACTGCACTTTTATTAATACTTTCATTACCTGTATTAGCAGGGGCAATTACAATACTTTTAACGGATCGAAACTTAAATACTTCATTTTTTGACCCATCGGGCGGAGGAGACCCTATTCTTTATCAACATTTATTTT???????????????????????????????????????????????????????????????????
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Decameria_spp ATAATAATTTGTTTTTTAATTGAAAACTAGAATGAAAGATTGGACGAAAAATTAACTGTCTCA-AATTTATTA-GTATTTAAATTTAATTTTTAAGTTAAAAAGCTTAAATTTAATTATAGGACGAGAAGACCCTATAGAATTTTATA-A---TT-A-AATT--A--TATTT-A--TTT--ATA-AAT-
AA-AT-TTTA--A-ATA-T-A-ATAAATTATTATATTGGGGTGATAAAAAAATTAAATAAATTTTTTTTT-TA-T-A-A------AAA-CAATAATTTTTGAA-T-T----TTTGATCCTAAA--T--T-T-
TAGATTATTAGAAAAAATTACCTTAGGGATAACAGCGTTATTTTTTTTAAAAGTTCATATTAATAAAAAAGATTGC??????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????
CCCGAAAGATCGA-ACGGGGAGATTCATCGTCAGCGGCGTCGGCTTCGC-CGCGG-TCAGCGATG--C-CTCG-G--GC-C-TT--CG--GGTCTC-CGTGGC-ACGCGGCCGCG-TCGCG--TTATGTCCGGCGACGTGGGCGTGCACTTCTCCCCTGGTAGAACGTCGCGACCCGTTGGGCGTCGGCATAAGGCACAA-TC-GGCACAGACTGT-CC--
CGGCGTT--C--GCG----T-C-GGGGCAAACCGAG--AGTTTCGCCCGGTCGGCTGCCCGACGGTACACGCAC-GGTATTTGGCCGCA-TCGAAC-GTGCGCCCGGCCCGTCGCAAGCGCGGTCAGCGAT----TCCCGGTGGTCGGAC-CTAGCGCCGTCCCCGGGCCTGGCCAGCTGTTGGCGATCGGTGTCCTCGGGCCGGCCA-T------------
T---ATA------
ATACCGGTCGGCGACGCTATTGCTTTGGGTACTTTCA???????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????
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GACTCCATAAAGGAGAAGTTAATTCTTCCATTCCTTGACATAGACTTGCATATCTATGATCTCGGCATAGAGAACAGAAATGCTACAGAGGACAGGGTGACTGTGGACTGTGCAGAGGCAGTGAAGAAATATAGYGTAGGCATAAAGTGTGCCACAATTACGCCAGACGAGAACCGTGTGAAGGAGTTTGATCTGAAGGAAATGTGGAAGAGCCCCAATGGCAC
CATTCGTAATATTCTAGGAGGCACTGTGTTCAGAGAGGCAATTATTTGCAAAAATATACCAAGGCTGGTAACTGGTTGGAACCAGCCTATCATAATTGGGCGCCATGCGCATGCTGATCAGTACAAAGCCACAGACTTTGTCGTACCTGGTGCTGGAACACTTGAGATCACCTTTACAGGCAAGGACGGGACGAAAATACAGCATACTGTTCATCAATTCAAGG
GCGCAGGTATCGCTCAAGCACAGTTCAATACCGACGAGAGCATCGAAGCGTTCGCACACAGTTCGTTCCAGTATGCACTATCCAGATCATACCCACTTTACTTGTCGACAAAGAACACTATATTGAAGAGATACGACGGCAGATTCAAGGATGTCTTCCAGGAGATCGATGACAGGGAATACAAGCCTAAGTTC?
AGGCTAAAAACATRTGGTACGAGCAGCGCTTGATCGACGATATGGTGGCCTACGCCATGAAATCCGAAGATATGCAGTTGATCTGTGAGGCGTACCATGTGATGAGAGACGGCTTGGGGCTGAGTCCGCAGGAGATGAGCCACGTCTTCGACGAGTGGAATAAGGGGGTGCTCGATTCATTCCTGATCGAAATCACTCGGGACATTCTGAAGTACAAAGACGAC
AAGGGATACTTACTGGAACGTATCCGAGACACGGCTGGTCAGAAGGGCACGGGAAAGTGGACGSCGATCGCCGCCCTGGAGTACGGAGTGCCTGTGACTCTAATCGGGGAATCGGTCTTCTCCAGGTGTCTGTCCGCGTTGCAGAGTGAAAGGGTTGAAGCAAGTGCAGTACTCAC------------------
TGGTCCGAACGCTCGTTATCAGGGCGATAAGAAGCAATTCCTCGAACATCTCAAGAAAGCTCTATATCTGTCGAAAATCATCTCCTACGCCCAGGGATTCATGCTTCTGCGAGAGGCCGCAAAAAYACACAATTGGAATTTGAATTACGGTGGAATTGCGTTG?????????????????????????????????????????????????????????????
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TTTACAGAAATTGTTGTCGGAGTACCAGCTATCTACTTGAATTATACCAAGAGTATTTTACCACCTAATGTTCAAATCAGCGGCCAAAACAGCTACAAAGTAGCTAAGGGTGCGTTTACTGGTGAGATTAGTCCAGCTATGCTACTCGATAATGAAGTTGCCTGGGTGATTCTTGGACATTCTGAACGCCGCAACGTATTTGGAGAAGATGATGCACTAATTGC
AGAAAAAATAAGTCACGCACTGAATGCTGGTTTGAAGGTGGTGATTGCCTGCATTGGGGAAAAGTTGAATGATCGTGAATCTGGTAAAACAGAGGAGGTAGTGTACAAACAGGTTAAAGCTATTGC??????????????????????????????????????????????????????????????????????????????????????????????????
?????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????ACAGGTGGTTTTTGGAGAAAATGAA?
AACATCACCTCGTACTACAGCGTCTTGGAGGCTCTGGACCAGACGAACCTCTCCTACGATGTGCTGCTGCACSCGAAGCAGATCGGATTCTCCGACAAGCAGATCGCTAAAGCTGTCAAGAGCACGGATCTCGCTGTACGCAAGCAACGTCAGGAGAGCAACATACGACCCTTCGTTAAACAAATCGACACTGTTGCCGCCGAGTGGCCGGCAACGACGAATTA
TTTGTACATGACTTACAACGGCAACAGCCATGACTTGCAATTCCCCGGCGGTTACACCATGGTTATAGGTGAG???????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????
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CCAGTTTGCAAAAGAAGTTCAAATCTTTGTTCGGCGAGAAGCTGGAGGTGGTTAGGACGCACCAGCAGCAAGAGAACCTCAAGTTTATGGCTCACTTCAAACGGAAGTTCATCATTCACCAAGGCCGTCGCAAGCAGCCAAAGTCGTCACC---
CAGCAAAGTGGAGTTTTACCATCTTCGYAGCAATGGCAGTGCTCTCTGCACGAGGTTAATTCAAGTACAGCCCGACGCTTTTCTACTGAATTCTGCCTTTTGGTCTAGCTACATTTTGAACGTGCCGTTCAACAATGATGACGAGAGCAGTATCRTCTACGTGTGGATAGGATCCAAGGCTGACAACGAGGAGGCTCGAATCGTCGAAGAAATAGCAGAAGAAA
TGTTTAACAACGTTCCATGGATAAGTCTACAAGTGTTAAACGAGGGCGAAGAACCCGACAACTTCTTCTGGGTTGGGCTTGGCGGCAAGAAACCATACGACACTGATGCTGACTACATGAACTATACCAGACTGTTTCGATGCTCCAACGAAAAGGGGTACTTCACCATCAGCGAGAAGTGCACTGATTTCTGTCAGGTA????????????????????????
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Heteroperreyia_spp ??????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????
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ATAATTATTCGAACAGAATTAAATACTGCAGGATCATTCATTGGAAATGATCAAACTTATAATACTATTGTTACATCACATGCATTCCTAATAATTTTTTTTATAGTAATACCTGTAATAATAGGAGGATTTGGAAATTGATTACTTCCTTTAATATTAGGAGCCCCTGATATAGCATTTCCTCGACTAAATAATTTAAGATTTTGATTACTCCCTCCTTCTTT
AATTCTTTTAACTATAAGAAGATTTACTAATCAAGGAGCTGGAACAGGATGAACTATTTACCCCCCTCTTTCTAGAAACATAGGACACTCTGGAAGATCTGTTGATTTAACAATTTTCTCCTTACATATAGCAGGAATCTCCTCTATTCTTGGAGCTATTAATTTTATTTCTACAATAATTAATATACGATGCTCAAATATAAAATTAGATAAAATATCTTTAA
TAATTTGAGCAATTATTATTACAGCAATCTTATTAGTAATCTCTTTACCTGTATTAGCAGGGGCAATTACTATACTTTTAACTGATCGAAACCTAAATACATCATTTTTTGACCCTTCAGGAGGAGGAGACCCTATTTTATATCAACATTTATTTTGATTTTTTGG??????????????????????????????????????????????????????????
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Parasyzygonia_cyanoptera ??????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????
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Pergagrapta_spp ATAATAATTAGTCTTTTAATTGGAGGCTGGTATGAAGGATTGGACGAGATAAATACTGTCTCAA-TTAAATTA-AGA-TTTAATTTAATTTTTAAGTAAAAAAGCTTAAATTTTATTATAGGACGAGAAGACCCCATAGAGTTTTATA-T---TA-A--GTT--T--TATATTT--TTTATAAA-AT--
TA-TA--TTTA-A-TTA-T-G-ATTTAGTATTTTATTGGGGCGATAGAAAAATTTAATAAACTTTTTTAA-TA-T-------TA-ATA-CAAAGATTATTGAA-A-T----TTTGATCCTAAA--T--T-T-
TAGATTATTAGATTAAGTTACCTTGGGGATAACAGCGTTATTTTTTTTAGAAGTTCATATTAATAAAAAAGATTGC??????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????
CCCGAAAGATCGA-ACGGGGAGATTCACCGTCAACGTGGTCGGCTTCGC-CGCGG-CCCGCGATG--A-GCTG-T--AC-C-A---C--GC--GTT-CAGCGC-ACGCGGTCGCG-TCGCGTTC-ATGCCCGTTCACGTAGGCGTGCACTTCTCCCCTAGTAGGACGTCGCGACCCGTTGGGCGTCGGCATAAGGCCCGT-GA---GATAGCCTGTATA--
CGACGCT--C--GCG----T-CGTGCACAGATCCA---CGG-TAGCCCGGCCTGCTTCCCGGCGGTACACGCAC-GGTATTGAGCCGCA-TCGAAC-ATGCGTCAGATACGTCGCAAGCGCGGTCAGCGTA----ACCCGGTGGTCGGAC-CTAGCGCCGTCCCCGGGACTGGCCAGCTGTTCGCGGACGATATTCTCGGACGGGCTC-GCA---C-
GCAT--------AA----TTACCGGTCGGCGACGCTACTGCTTTGGGTACTTTC-----
AATAATTATTCGAACAGAAATAATAACAACAAGGTCTTTTATTGGAGATGATCAAATCTATAATGTAATTGTTACATCTCACGCATTTTTAATAATTTTTTTTATAGTTATACCTATTATAATAGGAGGATTTGGAAACTGATTACTACCTCTAATATTAGGAGCTCCAGATATAGCATTTCCCCGACTTAATAATTTAAGATTTTGATTATTACCACCTTCAT
TAATTTTATTAACATTTAGTAGATTTATTAATTCAGGATCTGGAACAGGATGAACTGTATATCCGCCCCTATCAAATAATATTGCTCATGCAGGAACCTCTGTTGATATAACCATCTTCTCATTACATATAGCAGGAATCTCATCAATTTTAGGGGCTATTAATTTTGTATCAACAATAATTAATATACGAGCATCAGAAATAAGATTAGATAAAATACCCTTA
TTTATTTGAGCTATTACTATTACTGCTATTTTACTATTATT-
TCATTACCAGTTCTAGCAGGAGCTATTACTATATTATTAACAGATCGAAACTTAAATACATCATTTTTTGATCCCTCAGGGGGAGGAGACCCAATTTTATACCAACATTTATTCTTTTT?????????????????????????????????????????????????????????????????????????????????????????????????????????
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GACTCYATTAAGGAGAAATTAATTCTTCCATTTCTCGACATAGACTTGCACGTGTATGATCTTGGTATGGAGAATAGGGATGCAACCAATGACCAAGTGACRRTAGAYTGTGCMGAGGCTATYAAGAAATACAATGTTGGCATTAAATGTGCAACCATAACGCCTGACGARAAACGTGTGGARGAGTTCAAGCTSAAGCARATGTGGAAGAGYCCAAACGGYAC
TATTCGTAATATWCTTGGSGGAACAGTYTTTCGAGAAGCKATTATYTGTAAAAATATCCCAAGACTGGTGACTGGGTGGGAYAARCCCATCATAATTGGRCGTCACGCTCACGCTGAYCAATACAAAGCTACCGACTTYGTCGTTCCTGGYGCYGGAAAAYTGGAGATCAGYTGGACYGGWACAGATGGAAAKAARATWCAGCATACCGTTTAYGAATTTCGMG
GYGCTGGAGTGGCACAAGCACARTTCAACACCGACGAGAGCATCGAGGCYTTCGCTCACAGYTCATTMCAGTAYGCKCTGRASMGARAATATCCCCTKTACTTGTCCACCAAGAAYACCATCTTGAAGAARTAYGATGGCAGATTCAAAGAYATATTCCAAGATATATACGAGAGAGAATACGAGGCCAAATTCGAAGCRCTSAACATCTGGTAYGAGCATCGY
CTCATCGACGATATGGTTGCCTAYGCCATGAAATCCGAGGACATGCAACTGATTTGCGAGGCGTATCACATAATGCGAGACGGATTGGGCCTAAGTCCGCAGGAAATGAGCTCAGTGTTCGACGAGTGGAACAAAGGTGTCCTCGATTCCTTCCTCATCGAGATCACTCGAGACATTCTGAAGTACAGAGACGAGAAGGGATATCTCCTGGAGCGAATCCGAGA
TACAGCCGGGCAAAAGGGGACTGGGAAGTGGACGGCTATCTCCGCACTGGACTACGGGATTCCAGTAACCCTCATCGGAGAGTCTGTATTCTCCAGATGTCTGTCAGCTCTGCAGAGCGAAAGGATGGAAGCGAGTAGAGTCTTGAC------------------
AGGACCCAACGCTCTGTACGAGGGCGACAAGAAACAATTCCTCCAGCACCTCATGAAAGCTCTCTATCTATCCAAGATCATATCCTACGCTCAAGGATTTATGCTTCTGCGCGAAGCAGCCAAAATTCACAAGTGGAATTTGAATTACGGTGGAATTGCGCTGAATAAAAAGGAAATCGACGAAATTGTCGCTTTCTTGAAGGCTGGTCCTTTAGATCCGAATG
TCGGTATGACTATAGAGGTTGTCGTCGGAGTACCGGCTATCTATTTGAGTTACGTGAAAAACATTCTACCACCTAATGTACAAGTTAGTGGTCAGAATAGTTACAAGGTAGCTAAGGGAGCTTTTACGGGGGAGCTTAGTCCTGCTATGCTCTTGGACAACGGCATTCCCTGGGTAATCTTGGGGCACTCGGAGCGGCGTAATGTTTTTGGTGAATCAGATGAC
TTGATTTCTGACAAAATTAGCCACGCACTCGAAACTGGCTTAAAGGTGGTGATTGCCTGTATCGGTGAGAAACTGGAGGAGCGTGAGTCAGGTGCAACTGAAGAAGTTGTCTTCAAGCAGACTAAAGCTATTGCAGATAAAATTAAATCTTGGGACAATGTTGTGTTG????????????????????????????????????????????????????
GCAAGTTAGAGGAAGCGTTTCAAAAAGCCCTCAGAATGGTAGACGAGAACGTGAATGGATTCGATCCGTACCTCAAGGGTATAGACGAGGAGGAGCTGGAGAGGCCGACCGACAAACGGATGTTCGTTCTGGCGGCAGCCCTGAAGGCCGGTTACTCGGTAGACCGTCTGTACGAACTGACCAAAATCGACAGATGGTTTTTGGAGAAAATGAAGAACATCACC
TGCTTCTATACCGTTCTCGAAACCCTCGACCAATCGAAGCTCCGCCACCACCTGCTCCGGCAGGCCAAGCAAATAGGGTTCTCCGACAAACAGATCGCCAAGGCTGTCCAAAGCACGGAACTGGCTGTGCGCAAGCAACGCTACGAAAACAACATTCGACCATACGTTAAGCAAATCGACACTGTCGCCGCTGAATGGCCGGCGACGACGAATTACCTCTATTT
GACCTACAACGGAACCGTTCACGACGTCCAATTTCCTGGGTGGTATACCATGGTCATAGGTAAGCCAGGTTCAGGTGTATACAGAATTGGCAGTTCCGTTGAGTTCGACTGGTGCGCGGTTGGGTGTCTTCGGGAGCTGCGCCATCTCGGCCGGAAGACCATCATGGTCAATTATAACCCGGAAACGGTCAGCACCGACTACGACATGTGCGATCGCCTGTACT
TTGAAGAGATATCATTCGAAGTCGTCATGGACATTTATGACTTGGAAAATCCCGAGGGT??????????????????????????????????????????????
TCAGTTTGCAGAAGAAGTTCAAATCCCTATTTGGCGAAAAGTTGGAGGTCGTTAGAACGCATCAGCAGCAAGAAAATCTCAAATTCATGGCGCACTTCAAACGCAAATTTATCATTCACCAAGGACGACGTAAACAGCCCAAGTCTTCACC---
CAGTAAAGTGGAATTTTATCATCTGCGAAGTAATGGCAGCGCCTTGTGTACCAGATTAATTCAAGTACAGCCCGACGCCTTTCTATTAAATTCTGAATTTTGGTTTAGCTATATTCTGAATGTGCCATTTAATAATGACGATGAAACTGGAATCGTTTACGTATGGATTGGATCGAAAGCTGACAGCGAGGAAGCTAGACTCGTCGAAGAGATAGCCGAAGACA
TGTTCAACAATGTTCCATGGATAAGTCTGCAGGTGTTGAATGAGGGCGAGGAACCAGACAATTTCTTCTGGGTTGGCATCGGTGGAAAGAAAYCTTACGATACAAATGCAGATTATATGAATTACACTCGACTATTTCGATGCTCCAACGAGAAGGGCTATTTCACCATCAGCGAAAAGTGTACAGATTTTTGTCAGGTAGATGATTTAGCTGATGATGATATT
ATGATCCTTGACAACGGAGAGCAAGTATTTTTGTGGCTTGGTGCTCGATGCTCGGAGGTGGAAATTAAGCTCGCTTACAAATCAGCTCAAGTAAGGGTGTACATACAGCACACGCGCGTGAAGCAGCCGGAGAAACCGCGGAAATTATGTCTGACGGCCAAAAGCCAGGAATCACGAAGAATCACG

Perga_sp ??????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????
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Perreyia_picea ATAATAAATTGTTTTTTAATTAAGAACTAGAATGAATGATTGGACGAGAAATTAACTGTCTCATAATTAATAT-TTATTTGAATTTAATTTTTAAGTTAAAAAGCTTAAATTTTATTAGAGGACGAGAAGACCCTATAGAATTTTATA-A---TT-A-A-TT--AATTTTTA-A--AAT--TAA-AAT-
AG-AA-TTTT--A-TAT-T-A-AGTAATTATTATATTGGGGTGATGGAAAAATTATTTTAATTTTTTTAT-TA-T-T-A------TAA-CAATAATGATTGAA-T-T----TTTGATCCTATT--A--T-A-
TAGATTATTAGAATAAATTACCTTAGGGATAACAGCGTTATTTTTTTTAGAAGTTCATATTAATAAAAAAGATTGC??????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????
CCCGAAAGATCGA-ACGGGGAGATTCATCGTCGGCGGCGTCGGCTTCGC-CGCGG-CTCGCGATG--T-CGCG-T--AC-C-TC--TC--GGGGTA-CGGGGC-ACGTGGTCGCG-TCGCG--TTATGTCCGGCGACGTGGGCGTGCACTTCTCCCCCGGTAGAACGTCGCGACCCGTTGGGCGTCGGCATAAGGCCCGC-AC-GGCACAGACTGT-CC--
CGGCGTT-AACCGCG----T-CCGGGGCAGACCGTCG-CGGTTCGCCCGGCCGGCTGCCCGACGGTACACGCAC-GGTATTTGGCCGCA-TCGAAC-GTGCGCCCGGCCCGCCGCAAGCGCGGCCAGCGAT----TCCCGGTGGTCGGAC-CCAGCGCCGTCCCCGGGTCTGGTCAGCTGTTGGCGCGCGGTGTCCTCGGGCTGGCCA-ATC---G-AATT-
T---ACACAGA-
AATACCGGTCGGCGACGCTATTGCTTTGGGTACTTTCA??????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????
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GATATGCAGCTGATCTGTGAGGCGTACCACGTGATGCGGGACGGGTTAGGACTCAATCCAAAGGAGATGAGCGAGGTGTTCGACGAGTGGAACAAGGGGGTGCTGGATTCGTTCTTAATTGAAATCACCCGCGACATCCTGAAGTACAGGGATGATAAGGGGTATCTGCTGGAGCGTATCCGAGACACGGCTGGTCAGAAGGGCACGGGAAAATGGACGGCGAT
CGCCGCTCTGGATTACGGAATCCCTGTGACCCTGATCGGGGAGTCGGTGTTCTCCAGGTGTCTGTCCGCGCTACAAGCCGAAAGGATAGAAGCCAGCGCCGTTCTCAC------------------
TGGGCCAAGCGCTCGGTATCAGGGCGACAAGAAGCAGTTTCTCGAGCACCTCAAGAAAGCTCTTTATCTGTCCAAAATCATCTCCTACGCCCAGGGATTCATGCTACTGCGAGAGGCCGCGAAAATACACAACTGGAATCTGAATTACGGTGGAATTGCGCTG?????????????????????????????????????????????????????????????
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Perreyia_tropica ATAATAAATTGTTTTTTAATTAAAAACTAGAATGAATGATTGGACGAGAAATTAACTGTCTCATAATTAATTG-TAATTTGAATTTAATTTTTAAGTTAAAAAGCTTAAATATAATTGAAGGACGAGAAGACCCTATAGAATTTTATA-G---TT-A-AATT--ACTTTTTT-A--AAA--AAA-AAT--
A-AT-TTTT--A-TAG-T-A-ATAAATTATTATATAAGGGTGATAAAAAAATTAATTAAATTTTTTTTT-TA-T-A-A------AAA-CAATGATTATTGAG-T-T----TTTGATCCTATA--A--T-A-
TAGATTAATAGAATAAATTACCTTAGGGATAACAGCGTTATTTTTTTTTAAAGTTCATATTAGTAAAAAAGATTGCCAGCTCTAAGTGGGTGGTAAACTCCATCTAAGGCTAAATATTACCACGAGACCGATAGCGAACAAGTACCGTGAGGGAAAGTTGAAAAGAACTTTGAAGAGAGAGTTCAAGAGTACGTGAAACCGTTCAGGGGTAAACCTGAGAAACC
CGAAAGATCGA-ACGGGGAGATTCATCGTCGGCGGCGTCGGCTTCGC-CGCGG-CTCGCGATG--T-CGCG-T--AC-C-TC--TCGCGGGGTA-CGGGGC-ACGTGGTCGCG-TCGCG--TTATGTCCGGCGACGTGGGCGTGCACTTCTCCCCCGGTAGAACGTCGCGACCCGTTGGGCGTCGGCATAAGGCCCGC-AC-GGCACAGACTGT-CC--
CGGCGTTTAACCGCG----C-CCGGGGCAGACCGTCG-CGGTTCGCCCGGCCGGCTGCCCGACGGTACACGCAC-GGTATTTGGCCGCA-TCGAAC-GTGCGCCCGGCCCGCCGCAAGCGCGGCCAGCGAT----TCCCGGTGGTCGGAC-CTAGCGCCGTCCCCGGGCCTGGTCAGCTGTTGGCGCGCGGTGTCCTCGGGCTGGCCA-ATC---G-AATT-
T---ACAAAGA-
AATACCGGTCGGCGACGCTGTTGCTTTGGGTACTTTCAGGACCATAATCATCCGTACCCAATTAAGAACCTCTGGAACATTAATTTCTAATGATCAAATTTATAATACTATTGTAACATCCCACGCTTTTCTAATAATTTTCTTTATAGTTATACCAATTATAATAGGAGGATTTGGAAATTGATTAATCCCTTTAATAATTGGAGCCCCTGATATAGCTTTTC
CTCGAATAAATAATTTAAGATTTTGATTATTACCTCCTTCACTAATTCTTCTTTTAATAAGAAGAATAACTGACTCAGGGTCTGGGACAGGATGAACAGTTTACCCCCCCCTATCCAATAATTTAGCTCATGCAGGAGCTTCAGTTGATATAACTATTTTTTCTCTCCACATAGCAGGTATTTCTTCAATTTTAGGGGCAATTAATTTTGTATCAACCATAATT
AATATACGAGCATCAGAAATAAACTTTGATAAACTTCCCTTATTAATTTGATCAATTGCAATCACAGCTATCCTTTTAATTATTTCATTACCAGTATTAGCAGGTGCTATCACTATATTACTTACTGACCGTAATCTAAACACATCATTTTTCGACCCCTCTGGAGGAGGAGACCCTATTTTGTACCAACATTTATTTTGATTTTTTGGGCACCCTGAAGTCTA
TATTTTAATTTTACCTGCTTTTGGTATAATCTCACATGTAATTTCTCAAGAATCAGGAAAAAAGGAAACCTTTGGATCATTAGGAATAATTTACGCTATAATAGCTATTGGATTATTAGGATTTATTGTTTGGGCCCATCATATATTTACAGTTGGTATAGATGTTGACACTCGAGCTTATTTCACAGCTGCCACCATAATTATTGCTATCCCAACAGGAATTA
AAATTTTTAGATGACTAGCTACATTTCATGGTTCAAAATTTATTTATTCCCCTAATTTAATATGAGCTATAGGTTTTGTATTCTTATTCACTATAGGAGGATTAACAGGAATTATACTATCTAATTCTTCAA????????????????????????????????????????????????????????????????????????????????????????????
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GATATGCAGCTGATCTGTGAGGCGTACCACGTGATGCGGGACGGGTTGGGACTCAATCCAAAGGAGATGAGCGAGGTGTTCGACGAGTGGAACAAGGGGGTGCTGGATTCGTTCTTGATCGAAATCACCCGCGACATCCTGAAGTACAGGGATGACAAGGGGTACCTGCTGGAGCGTATCCGAGACACGGCAGGTCAGAAGGGCACGGGAAAATGGACGGCGAT
CGCCGCTCTAGATTACGGAATTCCTGTGACCTTGATCGGGGAGTCGGTGTTCTCCAGGTGTCTGTCCGCACTACAGGCCGAAAGGATAGAAGCCAGCGCCGTTCTTAC------------------
TGGGCCGAGCGCTCGGTATCAGGGCGACAAGAAGCAGTTCCTCGAGCACCTCAAGAAAGCTCTTTATCTGTCCAAAATCATCTCCTACGCCCAGGGATTCATGTTACTGCGAGAGGCCGCGAAAATCCACAACTGGAATCTGAATTACGGTGGAATTGCGCTG?????????????????????????????????????????????????????????????
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CCAGCTACATCCTGAACGTGCCGTTCAACAACGCCGACGAGACTGGCATCGTCTACGTGTGGATCGGGTCGAAGGCCGACCACGAGGAGGCCCGCATCGCCGAGGAGATAGCGGAGGACATGTTCAACAACGTTCCCTGGATCAGTCTGCAGGTGCTCAACGAGGGCGAGGAGCCCGACAACTTCTTCTGGGTCGGCCTCGGCGGGAAGAAGCCCTACGACACC
GACGCGGAGTACATGAACTACACCAGGCTGTTCCGCTGCTCCAACGAGAAGGGGTACTTCACCATCAGCGAGAAGTGCACCGACTTCTGTCAGGTAGACGACCTGGCCGACGACGACATAATGGTGCTCGACAACGGCGAGCAAGTGTTCCTCTGGTTGGGCGCCAGATGCTCGGAAGTGGAAATTAAGCTAGCCTACAAATCAGCACAAGTAAGTGTGTACAT
ACAGCACCTGCGCGTGAAGCAGCCGGAAAAGCC?????????????????????????????????????????????????

Philomastix_xanthophilax ??????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????
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Phylacteophaga_froggatti ATAATAATTAGTCTATTAATTGTAGGCTGGTATGAAAGATTGAACGAGGTAAATACTGTCTCAT-TAAAATTT-TTATTAGAATTTAATTTTTGAGTTAAAAAGCTTAAATTTTATTAGAGGACGAGAAGACCCTATAGAACTTTATA-A---TT-T--TAT--A--TTTTTTT--GAATTAAA-TT--
TA-TT--TGAT-A-ATA-T-T-TAAAATTATTTTATTGGGGCGATAGGAAAATTTAATAAACTTTTTTAA-TT-A-------ATATTA-CAATGATGTTTGAA-T-T----TTTGATCCTAGT--T--T-T-
TAGATTAATAGATTAAGTTACCTTAGGGATAACAGCGTTATTTTTTTTAGAAGTTCATATTAATAAAAAAGATTGC??????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????
CCCGAAAGATCGA-ACGGGGAGATTCATCGTCAGCGTCGTCGGCTTCGC-CGCGG-CTCGAGATG--T-CGTG-G--AC-C-A---C--GC--GTC-CGCGGT-ACACGGTCGCG-GCGCTC--TATGCCCGGCGGCGTCGGCGTGCACTTCTCCCCTAGTAGGACGTCGCGACCCGTTGGGCGTCGGCATACGGCCCGC-GC---GGTAGCCTGT-CT--
CGGCGCT--C--GCG----C-CG-GGGCAGACCCG---CGG-TAGCCCGGCCGGCTGCCCGGCGGTACTCGCAC-GGTACAGAGCCGCA-TCGAAC-TTGCGTCCAGCCCGTCGCAAGCGCGGTCAGCGAT----TCCCGGTGGTCGGAC-CCAGAGCCGTCCCCGGGCCTGGCCAACTGTTGGTTGACGGTGCCCTCGGACCGGCTC-
TTA------------------------TACCGGTCGGCGACGCTACTGCTTTGGGTACTTTC-----
A???????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????
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GACTCCATCAAGGAGAAATTAATTCTTCCATTCCTTGATATCGATTTACATATATATGATCTTGGCATGGAAAATCGGGATGCCACTGATGATAAGGTAACGATTGATTGTGCTGAGGCTGTGAAGAAATACAACGTTGGTATTAAATGCGCTACAATAACTCCTGATGAGAAGCGTGTTGAAGAATTCAAATTGAAGCAAATGTGGAAAAGCCCTAATGGTAC
CATTCGTAATATACTTGGTGGCACAGTCTTTCGTGAAGCTATTATATGCAAGAACATACCAAGATTGGTGACTGGCTGGGACAAACCTATCATCATTGGACGTCATGCATACGCTGATCAGTACAGAGCCACTGACTTTGTGGTTCCTGGTGCTGGAAAATTGGAAATCAGCTGGACTGGAACTGATGGCAAGAAGATTCAACACACTGTCTTTGAATTTAAAG
GTGCGGGAGTCGCCCAGGCACAGTTCAACACTGATGAGAGTATCCAAGCATTTGCTCATAGCTCTTTCCAATATGCGTTGTCTCGAGAGTACCCACTTTACTTGTCCACCAAGAACACCATCTTGAAAAAGTATGATGGTAGATTTAAGGACATCTTCCAAGAAATCTATGAAAAAGATTACCAATCTAAGTTTGAGGCAAAGAAGATCTGGTATGAACATCGT
CTCATCGATGACATGGTAGCTTATGCAATG?????????
GAYATGCAGTTGATCTGCGAAGCATATCACATGATGAGAGATGGACTGGGACTTAATCCTCAGGAGATGAGTGACGTGTTTGGAGAATGGAACAAAGGWGTGCTGGATTCCTTCCTCATCGAGATCACTAGAGACATTCTGAAGTACAAGGACGAKAAGGGRTAYCTACTGGAAMGAATCAGRGAYACGGCAGGGCAAAAAGGCACTGGGAAATGGACAGCMAT
YGCTGCYCTGGATTATGGAATCCCWGTRACTCTAATTGGSGARTCWGTGTTCTCTAGATGYCTATCTGCYYTRCARRCTGAGAGRRTMGARGCAAGYAAAGTKTTRAC------------------
KGGACCMAAYRCTCTCTAYCARGGMRACAARAAGGAGTTTCTCGAACAYYTGARRAAAGCTCTTTAYYTATCMAARATTATMTCYTACGCWCAAGGRTTCATGCTKYTMCGAGAAGCTGCTAARATYCACAAYTGGAATTTAAAYTATGGKGGAATWGCKCTWAGTAAAAAGGAAATTGACGAAATTGTCGCATTCCTGAAAGCCGGTCCACTTGACCCGAACG
TTGGTAAGGTTGCAGAAATTGTTGTCGGAGTCCCTGCAATTTATTTGAATTATGCTAGTGGTATTTTGCCACCCAACGTACAAGTGAGTGGACAAAATTCTTATAAAGTCGCAAAGGGTGCTTTCACTGGAGAAATTAGCCCTGCGATGCTTTTGGACAATGGTGTTCCATGGGTGATCCTGGGACACTCTGAGCGACGCAATGTCTTCGGCGAGTCTGATGAT
TTGATCTCCGATAAAATTCAACACGCACTTGAGACTGGTGTGAAAGTGGTGATTGCTTGCATCGGAGAAAAATTGGAAGAACGTGAGTCTGGAAAAACAGAGGAAGTCGTTTTTAAGCAGACTAAAGCCATCGCCGATAAGATCAAGTCGTGGGACAACGTTGTACTG????????????????????????????????
TGAAGTGATGGCGATTGGGAGAAAGTTTGAAGAAGCATTTCAAAAAGCATTGAGAATGGTAGACGAGAATGTGAACGGATTCGATCCCTACATCAAGCCCATCGACGACGAGGACCTGGAAAGACCAACGGACAAAAGGATGTTCGTCCTGGCCGCAGCTCTCAAAGCTGGTTACACGATTGACCGTTTGTACCAACTGACCAAGATCGATAGATGGTTTTTGG
AGAAGATGAGGAACATCACGACTTACTACACCATCCTGGAAGGCCTGAGGGAGACGAAACTCTCGTACGAAGTGTTGCTTCGCGCCAAACAAATCGGTTTCTCGGACAAACAGATCGCGAATGCGGTCAAGAGTACGGAATTAGCTGTACGAAAGCAGCGCAAGGAAAGCAATATTCGCCCTTTCGTTAAGCAGATCGACACTGTCGCTGCAGAATGGCCAGCT
ACGACGAACTACCTTTACCTTACTTACAACGGAACCAGCCACGACATCCAGTTCCCTGGTGGATACACGATGGTCATAGGTGAGCCAGGTTCCGGTGTATACAGAATCGGTAGTTCCGTGGAGTTCGACTGGTGCGCGGTTGGCTGTCTTCGAGAGCTTCGCAATTTGGGTCGCAAGACGATCATGGTGAACTACAATCCAGAAACGGTCAGCACAGATTACGA
CATGTGCGATCGCCTGTACTTTGAGGAAATTTCTTTCGAAGTCGTAATGGACATCTACGACCTGGAGAACCCGGAGGGA?????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????
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TAACGAAGAAGCAAGACTTGTCGAAGAGATTGCAGAAGAAATGTTTAATAACGTTCCTTGGATAAGTCTGCAGGTTTTGAATGAAGGCGAAGAACCAGACAATTTCTTCTGGGTTGGTATCGGTGGTAAAAAGCCCTACGATACTGATGCAGAATTTATGAATTATACAAGACTATTTCGATGCTCGAATGAGAAGGGTTATTTCACAATCAGTGAAAAGTGTA
CAGATTTTTGTCAGGTAGAAGATTTGGCTGACGATGATATTATGATACTTGACAATGGTGAACAAGTATTTTTGTGGCTCGGTGCTCGATGTTCCGAAGTGGAAATTAAGCTGGCGTACAAGTCAGCGCAAGTAAGAGTATACATACAGCACTTGCGTGTAAAGCAGCCTGAGAAACCTCGAAAATTGTTTCTCACTGCAAAAGGCAAGGAATCTCGACGATTC
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Polyclonus_atratus ATAATAATTTGTTTTTTAAATGAAAGCTAGAATGAATGATTGAACGAGGTAAAATCTGTCTCA-AATAAAATA-AAATTTAAAATTAATTTTTAAGTAAAAAAGCTTAAATTTAGTTATAGGACGAGAAGACCCTATAGAATTTTATA-A---T--T---TT--ATTTTTTG-T--AAA--T-T-
ATTTTT-TA-AATA--A-GAA-A-T-AAATATTATTTTATTGGGGCGATATAAAAATATATAAAACTTTTTTTT-TA-T-T-A------TAA-CATTTATTAATGAATT-T----TTTGATCTTAAA--T--T-T-
TAGAAAAATAGATTAAATTACCTTAGGGATAACAGCATAATTTTTTTTAAAAGTTCATATTAATAAAAAAGATTAT??????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????
CCCGAAAGATCGA-ACGGGGAGATTCATCGTCAGCGGCGTCGGCTTCGC-CGTGG-CTCGTGATG--T-CGCG-T--AC-CCA---C--GG--GTT-CGCGGC-ACACGGTCGCGGTCGTT--TTATGCCCGGCGTCGTAGGCGTGCACTTCTCCCCCAGTAGGACGTCGCGACCCGTTGGGCGTCGGCATAAGGACCGAATT---GGTTGACTGT-CT--
CGGCCGC--TTTGCGA---C-CG-GGGCAGACCGT---CGG-TTGCCCGGCCGGCTGCCCGGCGGTACTCGCAC-GGTATTGAGCCGCAATCGAAC-TTGCGCCCGGCCCGTCGCAAGCTCGGTCAGCGTT----TCCCGGTGGTCGGAC-CTAGCGCCGTCCCCGGGCCTGGCCAGCTGTTAGCGGACGGTGTCCTCGGGCGGGCTC-GCA---A-
ATCG-------------AATACCGGTCGGCGACGCTATTGCTTTGGGTACTTTC-----
AATAATTATCCGAACAGAACTAATAACATCAGGAACATTCATTGAAAATGATCAAATTTATAATACATTTGTAACATCTCACGCATTTTTAATAATTTTTTTTATAGTTATACCCCTAATAATAGGAGGATTTAGAAACTGATTAATTCCTTTAATATTAGGATCTCCTGATATAGCTTTTCCTCGAATAAATAATTTAAGATTTTGACTCCTTCCCCCATCCT
TAATTTTACTTCTTATAAGAAATTTAATTGATTCAGGAACAGGTACTGGATGAACTGTATATCCACCACTTTCAAGAAATATAGCCCATGCAGGATCATCAGTTGATTTAACTATCTTTTCCCTTCATATAGCAGGAATTTCATCAATTATAGGAGCAATTAACTTTGTATCAACTACAATAAATATACGATCATCTAACATAAATTTTGATAAAATTCCTCTA
ATAATCTGAGCTATTATAATTACAGCTATTCTTCTATTATT-
TCTCTTCCAGTATTAGCAGGAGCAATTACTATACTCTTAACTGATCGAAACCTAAATACATCATTCTTTGACCCTTCTGGAGGTGGAGATCCAATTTTATACCAACATTTATTTTTTTT?????????????????????????????????????????????????????????????????????????????????????????????????????????
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GACTCCATCAAGGAGAAATTGATTCTTCCATTCCTCGACATAGATTTGCACGTCTATGATCTTGGCATGGAGAACAGAGATGCAACCGACGACAAGGTGACTGTAGACTGTGCAGAAGCTGTTAAAAAATACAGTGTCGGTATAAAATGCGCAACAATAACGCCCGACGAGAAACGCGTAGAGGAATTCAAATTGAAGCAAATGTGGAAGAGTCCGAATGGTAC
CATTCGTAATATACTTGGCGGAACCGTATTCCGGGAAGCAATTATTTGCAAAAATATACCGAGGCTGGTAACAGGGTGGAATGAGCCCATTATAATTGGTCGTCATGCGCATGCCGACCAGTACAAAGCTACGGACTTTGTAGTCCCCGGCGCTGGAAAACTGGAGATCAGCTGGTCTGGAACAGATGGCAAGAAAATTCAACATACCGTCTACGAATTCAAGG
GAGCGGGTATTGCCCAAGCACAATTTAACACCGATGAAAGTATCGAAGCGTTCGCTCACAGCTCCTTTCAATACGCGTTGTCCCGATCTTATCCGCTGTATCTGTCGACGAAAAACACCATACTGAAGAAATACGATGGCAGATTCAAGGATATATTCCAAGACATCTACGAGAAGGAGTACAAGTCTAAGTTTGAAGCCAAAAACATTTGGTATGAGCATCGC
CTGATCGATGACATGGTCGCCTATGCCATG?????????
GACATGCAGCTGATCTGCGAGGCATACCACGTGATGCGAGATGGTCTCGGGCTCGACCCGAAGGAGATGAGCGACGTGTTCGACGAGTGGAACAAGGGCGTCCTCGACTCCTTCCTCATCGAGATCACCCGCGACATCCTCAAGTACAAAGATAACAAGGGATATTTACTGGAGCGAATCCGTGATACAGCCGGGCAAAAGGGCACAGGAAAATGGACGGCGAT
CTCCGCTCTGGACTACGGAATTCCCGTGACTCTAATCGGGGAGTCTGTGTTCTCCAGGTGTTTGTCCGCGTTGCAGAGCGAGAGGATAGAAGCCAGCGCTGTTCTGAG------------------
TGGACCGAACGAGCGGTACCAGGGTGACAAGAAACAGTTCCTCGAACACCTCAAAAAAGCTCTCTATTTGTCGAAAATCATCTCCTACGCCCAGGGATTCATGCTTCTGCGAGAGGCAGCGAAAATACACAACTGGAATTTGAATTACGGCGGAATTGCGCTTAGCAAGAAGGAAATCGACGAAATCGTTGCATTCTTGAAAGCCGGTCCTCTCGATCCCAACG
TTGGTATGTATACAGAAATTGTTGTCGGAGTACCAGCGATCTACTTGAACTACACCAAGAGTGTTTTGCCACCAAATGTTCAGATTAGCGGCCAAAACAGCTTCAAAGTAGCCAAAGGTGCATTCACCGGCGAGATTAGTCCAGCTATGCTGCTGGACAATGACATTCCTTGGGTGATCTTAGGACATTCGGAACGCCGCAACGTTTTTGGAGAAGACGACGAG
TTGACAGCTGACAAAATAAGCCACGCACTTGAAGCTGGTTTAAAGGTGGTGATCGCATGCATTGGTGAGAAGCTGGACGAGCGTGAAGCTGGCAAAACCGAAGAGGTAGTCTTCAGACAGACCAAAGCTATTGTGGATAAGATTAAATCGTGGGACAACGTTGTCCTG????????????????????????????????
TGAAGTCATGGCGATTGGTCGCAAGTTTGAAGAAGCGTTTCAAAAAGCACTCAGAATGGTGGATGAGAACGTGAATGGCTTCGACCCATACATCAAGTCCATCGACGACGAGGAACTGGAGAAGCCCACCGACAAACGGATGTTTGTCCTGGCTGCAGCACTCAAGTCCGGCTACACCGTGGACCGACTGTACGAGCTGACCAAAATCGACAGATGGTTTTTGG
AAAAGATGAGAAACATCACCGTCTATTACAACGTGCTGGAGGACCTGGACCAAACCAAGCTCTCCTACGAGGTGCTCCTGCGCGCGAAACAGATCGGCTTCTCAGACAAACAGATTGCCAAGGCTGTCAAGAGCACTGAGCTCGCTGTGCGCAAGCAACGACACGAGAGCGACATACGGCCCTTCGTCAAACAAATC???????????????????????????
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CCAGCTTGCAGAAGAAATTCAAATCGTTGTTTGGCGAGAAGCTAGAGGTAGTCATAACACACCACCAACAGGAGAACCTCAAGTTTATGGCCCATTTCAAGCGAAAGTTTATCATTCATCTTGGGCGACGTAAACAGCCCAAAGCATCGCC---
TAGCAGAGTGGAGTTTTATCATCTTCGAAGCAACGGTAGCGCTTTGTGCACCAGATTGATTCAAGTGCAACCGGACGCCTTCTTATTGAATTCTGCTTTCTGGTCCAGCTATATTTTGAACGTGCCCTTCAATAACGACGATGAGACTGGAATCGTGTATGTTTGGATAGGATCGAAAGCTGACAACGAAGAGGCGCGGCTCGTCGAGGAAATAGCAGATGACA
TGTTCAACAACGTTCCGTGGATAAGCCTGCAGGTGTTGAACGAGGGCGAGGAACCTGATAACTTTTTCTGGGTTGGCCTTGGTGGGAAAAAACCATACGATACCGATGCGGAGTACATGAATTACACTAGACTGTTTCGATGCTCCAACGAAAAAGGATACTTCACGATTAGTGAGAAATGCACTGACTTTTGTCAGGTAGATGACTTAGCAGACGACGATATA
ATGGTACTGGACAACGGCGAGCAAGTGTTTCTGCGGCTGGGCACCCGATGCTCGGAAGTGGAAATTAAGCTAGCATACAAATCAGCTCAAGTAAGA??????????????????????????????????????????????????????????????????????????????????????????
Pseudoperga_spp ATAATAATTAGTCTTTTAATTGGAGGCTGGTATGAAAGATTGAACGAGATAAATACTGTCTCAT-TAAAATTA-AAG-TTTAACTTAATATTTAAGTAAAAAAGCTTAAATGTTATTATAGGACGAGAAGACCCCATAGAGTTTAATA-T---TT-A--ATT--A--TAAT-TT--TTTAAAAA-AT--
TA-TA--TTAA-A-TTA-T-T-ATTAAATATTTTGTTGGGGCGACAGGAAAATTTAATAAACTTTTTTAA-TA-T-------TATTTA-CAAAAATTATTGGA-A-T----TTTGATCCTAAA--T--T-T-
TAGATTATTAGATTAAGTTACCTTGGGGATAACAGCGTTATTTTTTTTAGAAGTTCATATTAATAAAAAAGATTGC??????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????
CCCGAAAGATCGA-ACGGGGAGATTCACCGTCAGCGAGATCGGCTTCGC-CGCGG-CCCGCGATG--C-GCCG-T--AC-C-A---C--GC--GTT-CGGCGC-ACGCGGTCGCG-TCGCGTTCTATGCCCGGTCACGTAGGCGTGCACTTCTCCCCTAGTAGGACGTCGCGACCCGTTGGGCGTAGGCATAAGGCCCGT-GA---GGTAGCCTGT-AG--
CGGCGCT--C--GCG----C-CG-TTGCAGATCCA---CGG-TCGCCCGGCCTGCTGCCCGGCGGTACACGCAC-GGTATTGAGCCGCA-TCGAAC-ATGCGTCCAGTTCGTCGCAAGCGGGGTCAGCGAT----ACCCGGTGGTCGGAC-CAAGCGCCGTCACCGGGCCTGATCAGCTGCTGGCAGACGATGTCCTCGGACGGGCTC-GCT---C-
TCAT--------AAAT-ATTACCGGTCGGCGACGCTACTGCTTTGGGTACTTTC-----
AATAATTATCCGAACAGAAATAATAACCACAGGATCATTTATTGGAGATGACCAAATTTATAACGTAATTGTAACATCCCATGCATTCCTAATAATTTTTTTTATAGTTATACCAATTATAATAGGAGGATTTGGAAACTGACTTCTACCCCTTATACTAGGGGCCCCTGATATAGCATTCCCCCGACTTAATAATTTAAGATTTTGACTATTACCTCCATCTT
TAATCCTACTAACATTCAGAAGATTTATTAATTCAGGATCAGGAACAGGATGAACAGTATATCCCCCCCTATCAAGTAATATTGCTCATGCTGGGGCATCAGTAGATATAACTATCTTTTCTCTACATATAGCTGGAATCTCCTCAATTTTAGGGGCTATTAATTTTGTTTCAACAGTTATTAACATACGAGCCTCAGAAATAAGATTAGATAAAATACCCTTA
TTAGTATGAGCTATCACTATCACTGCAATTTTATTAATCATTTCTCTCCCTGTACTAGCAGGTGCTATCACCATATTATTAACGGATCGAAATCTAAATACATCATTCTTTGACCCATCAGGGGGAGGGGACCCTATTTTATACCAACATTTATTCTGATTCTTCGGACATCCAGAAGTTTATATTCTTATTTTACCAGCCTTCGGAATTATTTCCCACACCAT
CTCCCAAGAATCAGGAAAAAAAGAAACATTCGGAACACTAGGAATAATTTATGCCATATCCGCCATTGGATTATTAGGATTCATTGTTTGAGCCCACCACATATTTACAGTAGGAATAGATGTTGACACTCGAGCCTATTTTACAGCTGCAACAATAATTATTGCTGTACCAACAGGAATTAAAATTTTTAGTTGATTAGCAACAATTCATGGATCTAAAATTT
CTTATACTCCCAATATAATATGAACTTTAGGATTTGTATTTTTATTTACAATAGGGGGATTAACAGGAATCATACTATCTAACTCATCTATTGACATCATTATACATGACACATACTATGGACTCTATCAAGGAGAAACGGATTCTTCCATTTCTCGACATAGAATTGCACGTGTATGATCTTGGTATGGAGAATAGGGATGCAACTAATGACCAAGTGACGAT
AGAYTGTGCAGAGGCTGTCAAGAAATACAATGTTGGCATTAAATGTGCAACGATAACGCCYGACGAGATACGTGTGGAGSAATTCAAACTGAAGCAAATGTGGAAGAGYCCAAATGGTACTATTCGTAATATWCTTGGCGGAACAGTCTTYCGAGAAGCTATTATTTGTAAAAATATCCCAAGACTGGTGACTGGGTGGAACAAGCCCATCATAATTGGGCGTC
ACGCTCACGCTGACCARTACAAAGCTACCGATTTTGTCGTTCCTGGYGCTGGWAAATTGGAGATCAGCTGGACYGGAACAGAYGGAAACAAAATWCACCATACCGTTTAYGAATTCCGAGGCGCCGGAGTGGCACAAGCACAATTYAACACTGACGAGAGCATCGCGGCYTTCGCTCACAGYTCATTCCAGTACGCTTTGRACMGAGCRTATCCCCTGTACTTG
TCCACCAAGAACACCATCTTGAAGAAATACGATGGCAGATTCAAAGAYATATTCCAAGATATMTACGAGAAAGAMTACAGTGCGAAATTCGAGGCGCTCAACATTTGGTATGAGCATCGCCTGATCGACGATATGGTTGCCTATGCCATGAAATCTGAG?????????????????????????????????
ATGCGCGACGGATTGGGTCTAACTCCGCAGGAAATGAGCTCAGTGTTCGACGAGTGGAACAAAGGTGTCCTCGATTCCTTCCTCATCGAGATCACTCGAGACATTCTGAAGTACAGGGACGAGAAGGGATATCTCCTGGAGCGAATCCGAGATACAGCCGGGCAAAAGGGGACTGGGAAATGGACGGCTATTTCCGCACTGGATTACGGGATTCCGGTAACCCT
CATCGGAGAGTCCGTATTCTCCAGATGTTTGTCAGCTCTGCAGAGCGAAAGGATAGAAGCGAGTAGAGTGTTAAC------------------
AGGACCCAACGCTCTGTACCAGGGCGACAAGAAACAGTTCCTCGAGCACCTCATGAAAGCTCTCTATCTATCCAAGATCATATCCTACGCTCAAGGATTTATGCTTCTGCGAGAAGCGGCTAAAATTCACAAGTGGAATTTGAACTATGGCGGTATTGCGCTG?????????????????????????????????????????????????????????????
????????TTCTTGAAGGCCGGTCCTTTRGATCCGAATGTCGGTATG????????????????????????????????????????????????????????????????????????????????????????????????????TG------
GATAATGGTGTTTCATGGGTTATCCTGGGACACTCTGAGCCTCGCAATGTTTTCGGAGAATCGGATGACCTAATTGCGGACAAGGTCAACCACGCGCTCGAAGCTGGCTTGCAGGGGGTCATCGCTTGCATTGGTGGGAAATTGGATGAACGTGAAGGTGGTAAGACAGAGGAAGTGGTCTACAAACAGACCAAAGCTATTGCAGATAAAATCAAATCATGGGA
CAATGTCGTTTT?
TTACAGATCGGCAGTTCCATGAAGAGCGTCGGCGAAGTGATGGCGATAGGGCGAAAGTTTGAGGAAGCATTTCAAAAAGCCCTCAGAATGGTGGACGAGAACGTGAATGGATTCGATCCGTACATCAAGGATATAGACGAGGAGGAGCTGGAGAAGCCGACCGACAAACGGATGTTCGTCCTGGCCGCAGCCCTAAAGGCCGGTTACTCGATAGACCGTCTCTA
CGAGCTCACCAAAATCGACAGATGGTTTTTGGAGAAAATGAAGAACATCACCTCCTTCTACACCATTCTCGAAGCCCTCGACCATACGAACGTGTCCCACGAAGAGGTCCTGCAGGCAAAGCAGATCGGTTTCTCCGACAAACAGATCGCCAAGGCTATCCGAAGCACGGAACTGGCTGTGCGCAAGYTACGCCACGAAAACAACATTCGACCATTCGTAAAGC
AAATCGACACCGTTGCCGCTGAATGGCCAGCGACGACGAATTACCTCTATTTGACCTACAACGGAACCGTTCACGACGTCCAATTTCCCGGGTGGTACACCATGGTCATAGGTAAGTCAGGTTCCGGTGTATACAGAATTGGCAGTTCCGTGGAGTTCGACTGGTGCGCGGTTGGGTGTCTTCGGGAACTGCGACATCTCGGCCGGAAGACCATCATGGTCAAT
TATAACCCGGAAACGGTCAGCACCGACTACGACATGTGCGATCGCCTGTACTTCGAAGAGATATCCTTCGAAGTTGTCATGAACATTTACGACTTGGAAAATCCCGAGGGT??????????????????????????????????????????????
TCAGTTTGCAGAAGAAGTTTAAATCCCTATTTGGCGAAAAGTTGGAGGTTGTGAGAACGCATCAGCAGCAAGAAAATCTCAAATTCATGGCGCACTTTAAACGCAAATTTATCATTCACCAAGGACGACGTAAACAGCCCAAGTCTTCRCC---
CAGTAAAGTTGAATTTTATCATCTGCGAAGTAATGGCAGYGCCTTGTGTACCAGATTAATTCAAGTACAGCCCGACGCCTTTCTACTGAATTCTGAATTCTGGTTTAGCTATATTCTGAATGTGCCATTTAACAACGATGACGAAACTGGAATTGTTTACGTATGGATCGGATCGAAAGCTGACAGCGAGGAAGCTAGACTCGTCGAAGAAATAGCTGAAGACA
TGTTCAACAATGTTCCATGGATAAGTCTGCAGGTGTTGAATGAGGGCGAGGAACCAGACAATTTCTTCTGGGTTGGCATCGGTGGAAAGAAACCTTACGATACGCATGCAGATTATATGAATTTCACTCGACTATTTCGATGCTCCAACGAGAAGGGGTATTTCACCATCAGCGAAAAGTGTACAGATTTTTGCCAGGTAGATGATTTAGCTGACGATGATATT
ATGATCCTTGACAACGGAGAGCAAGTATTTTTGTGGCTTGGTGCTCGATGCTCGGAGGTGGAAATTAAGCTTGCTTACAAATCAGCTCAAGTAAGGGTGTACATTCAGCATTTGCGCGTGAAGCAGCCAGAGAAACCGCGAAAATTATTTTTAACCGCGAAAAGCAAGGAATCGCGAAGATTCACG
Pterygophorus_spp ATAATAATTAGTCTTTTAATTGGAGGCTGGAATGAAAGATTGAACGAAATAATTTCTGTCTCAA-TTAAGTTAAAAATTTGAATTTTATTTTTAAGTTAAAAAGCTTAAATTTGATTATAGGACGAGAAGACCCTATAGAGTTTTATA-T---TA-T--TTA--GAATAATT-T---ATTTTAATTT--
AA-TT-ATTAT---T-ATATTTA---AATATTTTGTTGGGGTGATAAAAAAATTTAATTAACTTTTTTAG-TT-T-A-----A-TTTA-CATTAATAAATGAA-T-A----ATTGATCCTAAT--T--T-T-
TAGATTGTTAGATTAAATTACCTTAGGGATAACAGCGTAATTTTTTTATTAAGTTCTTATTGATAAATAAGTTTGC??????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????
CCCGAAAGGTCGA-ACGGGGAGATTCATCGTCAGCGACGCCGGCTTCGC-CGTTA-CTCGCGATG--T-CGCC-G--AC-C-C---C--GC--GTC-GACGGC-ACGCGGTAGCG-TCGCG--TCATGCCCGGCGGCGTCGGCGTGCACTTCTCCCCCAGTAGGACGTCGCGACCCGTTGGGCGTCGGCATAAGGCCCGA-TT---GGTAGCCTGT-CT--
CGGCGTT--C--GCG----C-CG-GGGCAGACCGT---CGG-TCGCCCGGCCGGCTGCCCGGCGGTACTCGCAC-GGTATTGAGCCGCA-TCGAAC-TTGCGCCCGGCCCGTCGCAAGCTCGGTCAGCGTA----TCCCGGTGGTCGGAC-CTAGCGCCGTCCCCGGGCCTGGCCAGCTGTTCGCAGACGGTGCCCTAGGGCGGGCTC-GCA---C-
AAAA-------------ATTACCGGTCGGCGACGCTAATGCTTTGGGTACTTTC-----
AATAATTATTCGAACAGAATTAATAACTTTAACAACATTTATTGGAGATGATCAAATTTATAATGTTTTAGTAACCTCACATGCATTTCTAATAATTTTTTTTATAGTTATACCAATTATAATAGGAGGGTTTAGAAATTGATTACTACCATTAATATTAGGAGCCCCAGATATGGCTTTTCCACGTTTAAATAATTTAAGGTTTTGACTATTACCCCCTTCTT
TAATTCTTCTAACAATAAGAAGATTAATTGAATCAGGTAGAGGAACAGGATGAACAGTTTACCCTCCATTATCAAGTAATATTTCTCATTCTGGATCATCTGTAGACTTAACAATTTTTTCATTACATATAGCAGGAATTTCATCAATTTTAGGGGCTATTAACTTTGTATCAACTATAATTAATATACGAACAAAAGGAATAAGATTTGATAAAATATCATTA
ATAATTTGAGCAATTACAATTACAGCTATTTTATTAATTATTTCTTTACCAGTTTTAGCAGGAGCAATTACTATATTATTAACTGATCGAAATTTAAACACATCATTTTTTGACCCATCTGGAGGAGGTGACCCAATTCTTTACCAACATCTATTTTGATTTTTTGGACACCCTGAAGTTTACATTTTAATTTTACCAGCATTTGGTATTATTTCTCATGTAAT
TTCTCAAGAATCAGGGAAAAAAGAAACTTTTGGTACACTAGGAATAATTTATGCTATATCTACAATTGGATTATTAGGATTTATTGTTTGAGCTCATCATATATTTACTGTTGGTATAGATGTAGACACACGAGCTTATTTTACAGCAGCAACTATAATTATTGCTATTCCTACAGGCATTAAAATTTTTAGATGATTAGCAACTATTCACGGATCTAAAATAA
TTTATTCACCAAATATACTCTGAAGGTTAGGATTTGTATTTTTATTTACAATAGGAGGATTAACAGGAATTATTCTTTCAAATTCTTCTATTGATATTATTCTTCATGATACCTACTATGGATTCTATTAAGGAGAAACTGATTCTTCCGTTTCTGGACATAGACTTGCATATTTACGATCTTGGGATGGAAAACAGGGACGCGACCAACGATCAAGTAACCGT
GGACTGCGCGGAGGCTATAAAAAAATACAACGTTGGCATCAAGTGCGCTACGATCACGCCCGACGAGAAACGCGTGGAAGAGTTCAAGTTGAAACAAATGTGGAAGAGCCCGAACGGAACCATCCGAAACATCCTTGGCGGAACGGTGTTTCGGGAGGCGATTATTTGCAAGAATATACCAAGGCTCGTGACCGGATGGAATCAACCCATCATAATCGGACGTC
ACGCGCACGCCGATCAGTACAAAGCCACTGATTTCCTTGTTCCCGGCGCTGGAAAGCTGGAAATTAGCTGGACAGGGACAGACGGCAAGAAGATCCAACACACTGTATACGAGTTCAAGGGTGCGGGCGTCGCTCAGGCACAATTCAACACCGACGAGAGTATCGAGGCTTTCGCTCACAGTTCCTTCCAGTACGCGCTGTCCCGCTCGTACCCGTTGTACTTG
TCAACCAAAAACACCATCTTGAAGAAATACGACGGCAGATTCAAGGATATATTCCAAGCGATCTACGAGAAGGATTACAAGTCCAAATTCGAAGCCAAGAACATCTGGTACGAACATCGTCTCATCGACGACATGGTTGCCTATGCTATGAAATCCGAGGACATGCAACTGATTTGCGAGGCCTACCACGTCATGCGAGACGGATTGGGTCTCAGTCCACAGGA
GATGAGTGACGTGTTCGACGAGTGGAACAAAGGCGTGCTCGACTCGTTTCTGATCGAGATCACGCGAGACATTTTGAAGTACAAGGACGAGAAGGGCTACTTATTGGAGCGTATCAGGGACACGGCGGGTCAAAAGGGCACTGGGAAATGGACTGCGATCGCTGCTTTGGATTATGGCATGCCAGTTACCCTCATCGGAGAGTCAGTCTTCTCYCGATGTCTGT
CTGCACTACAGAGCGAAAGAATAGAAGCGAGTGCCGTTTTGAC------------------
AGGACCCAATGCTCGGTATCAGGGGGATAAAAAACAGTTCCTCGAACATCTCAGAAAAGCTCTCTATATGTCGAAAATCATCTCTTATGCTCAAGGATTCATGCTTCTAAGAGAAGCTGCAAAGATTCACAACTGGAATTTGAACTACGGTGGCATTGCGCTT??????????????????????????????
TTCCTGAAAGCCGGCCCGCTTGACCCCAACGTCGGTGAGTTTACAGAGGTCGTCGTCGGAGTGCCGGCTATATACTTGAACTACGCCAAGAGCATTTTACCGCCTAACGTTCAAGTTTCCGGCCAAAATACCTACAAGGTGGCCAAGGGAGCGTTCACCGGAGAGCTCAGTCCCGCTATGCTTCTGGACAACGACGTTCCCTGGGTGATTTTGGGACACTCGGA
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 Joseph spoke truly. Peter had at last fallen into the hands of
guardians of the law, and this time it was the King’s own men that
held him. It was evident, too, from the way they held him that they
thought they had a prize. They did not stop at the Town House where
offenders against municipal law were kept but marched straight
along Castle Street in the direction of the royal castle on Wawel Hill.
At the church they found Pan Andrew in a very sweat of anxiety
and fear lest something of harm had befallen them. He caressed
them all one after another and then said to Joseph earnestly:
“I want you to remain here and sound the Heynals for the rest of
the night. There is much work to be done in the quarter where the
fire is, and every man’s hand is needed to stay the flames. . . . I see
that Pan Kreutz is not with you. He stayed, too, I presume, to work
with the rest of the men?”
“Indeed, father, I know not. We called many times, but his loft
was a mass of flames like to a roaring furnace, when we were driven
down the stairs.”
“I must see, then, if I can find aught of him. He has been on a
previous occasion our very great benefactor and it would but suit us
ill not to seek at least his body in the ruins. Should he not have
perished, as I pray God he has not, then we can offer him shelter
here until such time as he can find a roof again.”
But when Joseph told him of the capture of Peter he looked very
serious and said that if such people were in the city then he had
better not leave his wife and the young people. On second thought,
however, it seemed right for him to go, for the city was now lighted
by flames and it would be easy to summon aid if they were attacked.
And so he, with thousands of other valiant men, fought the fire in
Krakow that night. They formed in a ring about the conflagration and
tore down all the buildings across which it might run. The Collegium
Minus was the last building to catch on the side toward the city wall,
and then the fighters tore down the houses near the old Jew Gate
and stopped the fire there. The flames swept around the other
buildings of the university, destroyed one or two, though not all, and
were finally halted on the second street above St. Ann’s. Sweeping
in the other direction, the fire had early in its progress destroyed the
monastery and adjoining houses of the Church of the Franciscans,
and had crossed over to Castle Street where it burned flat a whole
line of buildings.
On these and the other edges of the district a wide belt of
destruction was created by the fighters. This belt, the tradesmen,
running to and fro with water wagons filled constantly at the
aqueduct, wet down and soaked until it was almost a water wall. So
furiously had they worked that the main progress of the fire was
checked in seven or eight hours, and although certain buildings and
ruins smoked and even blazed for several days afterward, yet the
great danger passed when this well-soaked belt of destruction was
completed.
When Pan Andrew returned to the tower in the full blaze of the
morning sun, nearly one third of the city of Krakow lay in ruins.
Fortunately it was not the better portion of the city, and many of the
old wooden-built hovels had been there since before the days of
Kazimir the Great; that monarch had successfully converted about
one half the city of Krakow from wood to stone more than one
hundred years before; had he not done so it is probable that the fire
of 1462 might have utterly consumed it.
Elzbietka and Joseph’s mother were asleep on the trumpeter’s
bed clasped in each other’s arms. Joseph sat outside the
compartment with the hourglass before him on a beam, gazing out
over the smoking ruins of the university quarter.
“Is the city saved?” was the first question he asked his father.
“It is not now in danger,” answered Pan Andrew. “But there are
many homeless souls in the city this day.”
“Did you see the alchemist?”
“I did not. He has disappeared as if he had flown away on the
clouds of smoke that covered the city.”
“Poor Elzbietka,” exclaimed Joseph.
 The girl inside the compartment moaned slightly as her name
was spoken, although she was deep in a heavy slumber.
“I wonder if he was caught in the loft?” mused Pan Andrew. “It
was in the very center of the burned district.”
The answer to his question came with sudden unexpectedness.
There was a sound of footsteps on the stairs and Jan Kanty’s head
appeared from below. The scholar was leading another man by the
arm, a man who had been in the fire—his charred clothes and
blackened face showed it; around his shoulders and falling to his
waist was all that remained of what had once been a black robe. He
kept his hands beneath this robe.
“Pan Andrew,” whispered Jan Kanty softly, “I have found in the
street—Pan Kreutz.” And, checking the other’s startled exclamation,
he explained, “He is not in his right senses. Something has affected
his brain. But he has here something of interest to us all.”
Pan Andrew turned toward Kreutz—he never would have
recognized him had not Jan Kanty identified him; Joseph felt his
eyes glued with strange eagerness upon the eerie, blackened figure
and the mysterious folded hands beneath the robe; it had been a
scholar’s robe once.
“Ha, ha, ha!” laughed the alchemist suddenly, “up to heaven goes
everything in fire and yet no gold is found anywhere. Johann Tring!”
he looked about anxiously. “Where is Johann Tring? He answers me
not. He is lost in the flames, the flames that came so red and purple
when niter mixed with charcoal. Oho, Johann Tring! Come, Johann
Tring, and see what I have carried this whole long night for you.”
Throwing back the black robe, he held up the object that he had
been concealing there, and at the same moment the sun, streaming
in through the little window on the east side, fell full upon that object;
fell upon it and made it sparkle like the myriad of dew diamonds
shining upon a morning lawn new-mowed; sparkled like the
thousand chandeliers in the King’s great hall in the palace on the
Wawel Hill; sparkled like the rubies and emeralds that gleam in the
Queen’s crown; sparkled like the wondrous thing that it was, all
touched by the red rays of the morning sun—the Great Tarnov
Crystal!
“Now whence has that come?” shouted Pan Andrew so loudly
that the sleepers in the next room awoke. “Where by all that is good
and holy in the world have you found the gem which has been in my
family for years and years, which all my ancestors and I have sworn
to guard forever and to surrender to no person except to the King of
Poland? How has it come into your hands after it was stolen from
me, and my heart was nearly broken? Did you get it perhaps from
that ruffian who has been captured by the King’s guards? Did you
find it perhaps in the ruins of the town? Did you perchance——” The
truth suddenly flashed upon him and he was speechless.
 “It is an accursed thing,” cried out the alchemist suddenly, reeling
in Jan Kanty’s arms as if he were gone faint. “There is blood upon it,
and fire! It has lured princes and kings to their destruction! It has
made men’s brains mad with lust for want of it! It has caused good
men to steal, and evil men to kill. I will have none of it. I will have
none of it, I say.” He was growing almost boisterous, yet there was
something in this attack of madness that had much of reason and
determination in it. “I will have no more of it,” he repeated, “and no
more of Johann Tring.”
At that he fell fainting to the floor.
Jan Kanty raised him, and Elzbietka who had run out from the
trumpeter’s room rushed to him and kissed and fondled his
blackened hands.
 Pan Andrew picked up the Great Tarnov Crystal and held it at
arm’s length with a smile.
“Now may peace come upon us all,” he said, “for I may fulfill the
oath that my family has taken and deliver this to the King. While the
secret of its hiding place remained with me I might keep the crystal
as long as I chose, but now that the secret is out, there is but one
place where it may be guarded safely and that is in the palace of the
King. Pan Kreutz is right. This jewel has already done too much
harm in the world.”
“Then you may rid yourself of it at once,” broke in Jan Kanty. “The
King returned to Krakow two days ago, and we may find him at the
castle this very morning.”
 CHAPTER XV
KING KAZIMIR JAGIELLO

O f all the wonders that the capital city of Poland possessed

Joseph knew of none that stirred his imagination more than
did this royal castle of the kings upon the Wawel.
Impregnable through many sieges, its great rock base had stood,
brick and stone heaped high above it in a great mass of towers,
turrets, and walls. At its very heart, high above the winding Vistula
and the town, stood a strangely built round tower enclosed and
protected by the palace wings, where men in prehistoric times
worshiped the old nature gods of the Slavs; a place of rest and
seclusion where on rare occasions, when townsfolk might visit the
castle, Joseph had stood thinking of things that had been in the old
days.
He knew well the legend of Krakus, the hero of the dark ages,
who slew a dragon that had once made this hill his habitation. There
was a cave, so Joseph heard, that ran from the fortress underground
beneath the river, a secret exit in time of siege; here had been the
dragon’s lair, until the hero overcame him, and from that day men
made the Wawel a home, from which might be seen climbing into the
air, spires and belfries. All this Joseph had seen; he had fed his
fancies upon every object that graced the bleak, majestic rock, and
yet there remained one glory that had never yet met his eyes. That
glory was Poland’s King.
 But this morning after the fire, when the little company set out
from the Church of Our Lady Mary toward the Wawel Hill, Joseph felt
his heart leaping strangely in his breast at the thought of the
adventure that was to be theirs. To see the King, to have audience
before him—it made the blood sing in his ears and tingle in his finger
tips.
They took the alchemist with them, on Jan Kanty’s advice,
although he still seemed like a man in a dream.
“I found him wandering through the fire-swept streets early this
morning,” said the scholar. “He had been running hither and thither
all night long in the most dangerous parts of the city, and how he has
escaped death from falling timbers and burning coals is more than I
know. . . . The man has something on his mind, something that
troubles him hugely, and with it all he seems to be acting like one in
a spell.”
“Do you think it well to bring him with us?” asked Pan Andrew,
who had doubted from the beginning that there would be any benefit
from the man’s presence.
“Yes—I have a curious notion,” answered the scholar, “that he
may be able to help us. We have much to explain to the King, and
the man’s presence will make our story more credible. And who
knows, perhaps the alchemist himself may get help—he needs some
light thrown into that brain of his, and since he is harmless, it will do
no damage to take him.”
Pan Kreutz’s hands and face had been washed and dried, and
most of the fire grime had left him; the scholar’s robe was useless,
however, and Pan Andrew hung a kontusz or long coat about his
shoulders.
Joseph was there with the three men; Wolf had been left behind
sleeping upon the floor of the high tower room. Joseph’s mother and
Elzbietka were under the protection of the day watchman who
relieved Pan Andrew at dawn. It was necessary for Pan Andrew and
Jan Kanty to assist the alchemist in walking at times when his feet
would shuffle curiously like those of a man walking in his sleep, but
he plodded along bravely, not yet realizing quite clearly what was
happening about him yet confident that the two men near him were
his friends and were leading him to some good place.
 From Castle Street they turned at length up the long slope
leading to the castle on the Wawel. Behind them lay street after
street of desolation, of smoking ruins, of masses of wood still
flaming; amidst these ruins men were still working valorously tearing
down charred beams and hurling in tons and tons of water from the
water wagons, which were now all drawn by horses. One side of
Castle Street had suffered badly, the houses on the Street of the
Pigeons were entirely destroyed, St. Ann’s Street had but few
buildings left, while much devastation had been done along the
Street of the Bakers, the Street of the Goldsmiths, and the Street of
the Jews and Broad Street.
Jan Kanty’s company was challenged twice by guards on the way
to the palace, but when the soldiers recognized the good father, they
were at once passed along without question. It was another proof to
Joseph of the esteem in which the man was held; in himself,
however, there was not the least indication of pride and ostentation,
he was as simple as a child in most matters affecting worldly things,
and yet his name was as magic even in the court of the King. At
length they all stood in the little passageway on the Wawel through
which one passes to the palace, and here the guard, with spear
raised in salute to the scholar, bade the company wait until he went
to see if an audience might be had.
The soldier came back quickly. “The King,” he said gravely, “will
grant any request that may be made by Father Jan Kanty; he only
begs that the company wait a few minutes until a present audience is
finished.”
They waited perhaps fifteen minutes until an important-looking
functionary in a blue robe came to announce that King Kazimir
Jagiello would receive Jan Kanty and his friends.
Out into a wide court they went, following the courtier in blue, up
a marble staircase to the left and along a balcony. Then suddenly a
door was flung back and they were in the presence of the King.
To Joseph, remembering it afterward, it all seemed like a dream,
it was all so quiet and without ceremony. King Kazimir had chosen to
receive them in a small antechamber in which he often met certain
persons who were to be received without the usual ceremony of
presentation, and Jan Kanty was one of the privileged ones that he
met in such fashion.
Joseph and his father dropped upon one knee in front of the
King. He was sitting in a high-backed chair without a canopy, which
bore at its highest peak a royal crown; this crown was just above the
monarch’s head so that at first it seemed as if it were actually upon
his head and he were crowned. He wore a huge purple robe which
fell clear to the tops of his soft leather sandals; it had a great collar
embroidered with silks of many colors and in many patterns; a heavy
gold chain held the folds of the collar together, and beneath the
collar folds could be seen a rich vest embroidered with gold. The
sleeves of the robe were immense and hung down far below his
knees as he sat there; the robe itself was fringed with heavy fur. His
head covering was a simple cap of the same color as the robe, flat,
soft, and turned up a trifle at each side.
The King himself seemed the picture of comfort and informality;
not so his guards. On each side of him stood a guard in plate armor,
with stiff metal pieces over the arms, stomach, thighs, and legs. At
the waist they wore short, straight swords ready for action at a
second’s notice. These two men were as motionless as statues.
About the room stood knights in different kinds of armor, some in
light chain with long skirtlike coats, some in mail jackets that
resembled checkerboards in pattern and extended only from
shoulder to thigh, some in heavy armor and metal shoes armed with
spurs.
In front of the King were two pages carrying scepters. They, too,
stood motionless as he spoke.
“What is this?” he asked, as Jan Kanty came forward to kiss his
hand, which ceremony the King did not allow. “Have we here some
poor city dwellers driven forth by last night’s fire?”
“Yea,” answered Jan Kanty, “that is true, though we come not on
business of that sort. We are here upon some matter that may be of
deeper interest than one would suppose. These are Pan Andrew and
his son Joseph, by family name Charnetski, dwellers of the Ukraine
driven forth by violence and come here to have audience with your
Majesty.”
“So,” said the King with quick interest. “Stand, if you please, and
tell me the circumstances of your trouble for it greatly interests me at
this present time. I have much news from the Ukraine, and not so
pleasant, either. How came you by your misfortunes?”
“If your Majesty please,” began Pan Andrew, rising and taking out
the crystal from beneath his coat, “I wish to deliver to your Highness
the Great Tarnov Crystal.”
The sunlight touched it as he held it up, and the room and its
splendid company were suddenly agleam with wavy flecks of light,
red and orange and blue and yellow; there was a dazzling brilliance
to it that struck each eye with almost the force of lightning. The King
literally sprang forward to take the wondrous thing from Pan
Andrew’s hands.
“What a marvel! What a thing of beauty!” he exclaimed, while a
very murmur of astonishment ran through the circle of his attendants.
“Where in the world is to be found any jewel one-half so miraculous
as this?”
“I know not,” answered Pan Andrew, “but it has been in the
keeping of my family for many years.”
“Then why do you deliver it up to me?” demanded the King. “It is
worth a quarter at least of all the treasures in this palace.”
“That I will explain. My family has held it in trust these two
hundred years and more, and we have sworn to guard it until the
secret of its hiding place became known, and then, since there would
be great danger following such a discovery, to deliver it into the
hands of the King.”
“Then its hiding place has been discovered? But tell me first the
reason for concealing such a wondrous stone.”
“That, your Majesty, is a long story, which if your Majesty so
desires I will deliver shortly in writing, but I may say briefly that when
Tarnov fell before the Tartars these many years ago the citizens
entrusted this crystal to a member of my family. He took oath to
guard it zealously, with his life if need be, lest it fall into the hands of
people who might abuse its powers, for its beauty hides strange
properties which are allied to magic and sorcery and the Black Arts,
and it has been at times a curse, a thing of mystery and a source of
evil. When Tarnov was rebuilt new dwellers came there and the
crystal remained in our family.”
“How did the secret become known?”
“I had a servant, a Tartar. He was with me for many years. It was
my custom to conceal the crystal in the rind of a pumpkin, and many
a time this man must have seen me scraping out the inside of a
pumpkin and rubbing the shell with oils and gum in order to preserve
it. Because he was a simple fellow, I took no pains at any time to
conceal my task. But though lacking in wit, it seems that the man
was not lacking in curiosity. And his curiosity, I now believe, led him
to spy upon me, and eventually he discovered the use to which I put
the preserved pumpkin rind. He left me about a year ago, and it was
only a few months later that my house was attacked. I am sure that
he sold his information to some Tartar chieftain.”
“Could he suspect the value of the crystal?”
“That I do not know. I do know, however, that legends concerning
this crystal are everywhere to be found among the Tartars and
Cossacks. When they are children they are told tales of it, and all of
them grow up in the hope that some day they may find it.”
“Thou thing of beauty,” said the King, gazing at the crystal, “could
thou but speak and tell all that men have done to possess thee.
Thou cruel, marvelous thing.”
Pan Andrew fell upon his knees before the King. “Take this
crystal and guard it, your Majesty,” he exclaimed with great feeling,
the tears already streaming down his face. “It has already done
enough harm in the world. In my own family it has been nothing but a
burden, a source of endless anxiety and suffering. My father’s
fathers, years and years ago, even dug a passageway in the earth,
through which one might escape with it secretly in case of an attack,
and so cleverly was this passage concealed that for years no one
but the master of the family has known of its existence.
“In spite of the beauty of this jewel I hate it from the very bottom
of my heart and I hope that I may never look upon it again. For every
ray of light that it reflects, thousands of men have fought and died for
its possession, for every color that lurks within its depths miseries
and sufferings have swept over whole nations. I have guarded it
faithfully but no more shall I guard it. I am fulfilled of my oath.”
The King looked into the crystal fixedly, and then suddenly
shuddered as if he saw something fearful there.
“I shall be before many years an old man,” went on Pan Andrew,
in a pleading tone. “My home in the Ukraine exists no more. My
house is burnt, my fields are laid waste, and all because I had this
jewel in my possession and men envied me.”
He then went on to tell the story of the escape from the Ukraine,
the pursuit, the attempted robbery of his house, the attack on the
tower, and the persistency and repeated appearances of Peter of the
Button Face whom he had heard of in the Ukraine as Bogdan the
Terrible.
“I do not know,” he said, “who it might be that sent this man to
dog my steps, but my son Joseph has told me that your guards have
taken this same Peter a prisoner in the streets, and that he is a
captive of your men. Let me confront him here and perhaps I may
learn who it was that drove me from the Ukraine.”
While he was speaking the King gradually took his thoughts from
off the crystal, and when he mentioned the name Peter, the King
grew restless with excitement.
“I have the man,” he exclaimed, “and he shall be brought here.
My spies in the Ukraine reported recently that a great treachery was
afoot and that this man Peter or Bogdan was in Poland for the
purpose of consummating it. His description was given to my guards
and a reward was offered. Last night he was seen in the district
where the fire was raging and my guards brought him in. I shall have
him here directly.”
 Two spearmen brought him in; as he walked, the chains which
hung from his arms and legs clanked on the floor. He did not deign at
first to glance at Pan Andrew or any of his party, but simply looked at
the King and folded his arms defiantly and with spirit. Whereat the
two guards forced him to his knees.
His air of indifference disappeared, however, when his eyes fell
upon the Tarnov Crystal which the King had set down upon the floor
before him. He glanced left and right and favored the trumpeter and
the alchemist with a look of bitter hatred.
“You are accused of treason against the Commonwealth of
Poland,” said the King immediately. “Have you any plea to make?”
“Who accuses me?”
“The governors of the Ukraine,” answered the King. “And
moreover you are charged with other crimes, among them that of
persecuting this citizen here before me—you have destroyed his
home and fields, and attacked him while he was on duty in the
church tower. The penalty for any one of these is death.”
Peter did not lose his self-assurance for a moment. He realized
more quickly than another might that his plea of innocence would
soon be broken down. He fell back then quickly upon another means
of obtaining his end.
“I would buy my freedom,” he asserted.
“What have you that is worth while to me?” asked the King.
“Much. You are threatened in the Ukraine.”
The King thought for some minutes. It rather irked him to give this
man his life since he had already done such damage, but on the
other hand he might be able to obtain some really valuable
information. The whole Ukraine was in some kind of uproar, and
even his most trusted spies had not been able to get to the bottom of
it. The usual method of obtaining information from prisoners in those
days was torture, and in the field of battle it was employed widely,
but often in cases of such desperate men as Peter torture led them
to confess wildly but seldom with truth. The Cossack was ordinarily a
man of his word, and Peter had enough Cossack blood in him to
make him pass for a Cossack in the Ukraine and in the East.
“It pleases me to be merciful to-day,” replied the King. “There has
been too much suffering at my very gates to make me wish for more.
Your death would in no way pay for your crimes, and it is possible
that your information might be of service to the commonwealth. I
could wrest this information from you by torture, but I prefer an
easier way. . . . Now, mark,” he cautioned the Cossack, “I know
certain facts concerning what you have to tell, I have information
from my own men in the Ukraine, and if you utter so much as one
word of a falsehood to me, I will have you taken out and hanged by
the neck from the tower gate. Do you understand?”
“I understand,” answered Peter, turning just a trifle pale at the
threat. He was a bold man, he was a desperate man—otherwise he
would not have ventured back into Krakow after having been
defeated there twice—and he had no fear of death in any form, so
long as he was free and able to fight. But now his knees smote
together at the thought of hanging and he resolved that he would
keep close to the truth. After all, the whole affair was finished for him.
The crystal was in the hands of the King and he was not likely to part
with it easily.
“One thing,” he said in a low tone, “one thing, your Majesty, I beg,
and that is that you will let none talk of what I say. For if it were
known that I had spoken the truth, my life would not be worth—that,”
he snapped his fingers. “I have your promise, your Majesty.”
“You have.”
“Then hear what I have to say. I am Bogdan, known in the
Ukraine as the Terrible. Two years ago in March I was summoned to
Moscow by one in authority who said that a powerful lord had
something to say to me. Now having an open mind always for new
activities, I went, although our people have but little love for
Muscovites. And there I was taken to one Ivan.”
The King interrupted. “You mean——”
 “I mean Ivan himself, chief power among the Muscovites, son of
that blind one. He has the ambition to unite all lands thereabouts
under himself—as Emperor, men say.”
The King bit his lips and his eyes flashed. “This they have told
me,” he exclaimed in an angry voice. “I only wanted the confirmation
of it that you have given me. Ivan—Ivan—that one who makes
friendly proffers to one’s face and strikes in treachery when the back
is turned.” He strode up and down the room for a moment and then
turned to the captive again. His tone was as calm as it had been in
the beginning. “Proceed,” he ordered.
“In this he has partially succeeded, but his ambitions run higher,
and he dreams of establishing his power over the people outside the
borders, the Ruthenians and Lithuanians. Knowing them to be
willingly under Polish domain, however, even the city of Kiev which
fell beneath Tartar rule, he wishes instead to strike a blow at the
Poles in the Ukraine. Some one advised that he loose the Tartars
against the Poles, and an ambassador was even sent to find out
what would induce the Khan to send his warriors to fight the Poles.
The answer that he made was a curious one.”
“And that was——” asked the King.
“This was his answer. He would lead his Tartars against the Poles
in the Ukraine on one condition, and that was that Ivan should
deliver into his hands the Great Tarnov Crystal.”
At this the whole company started, chief among them Pan
Andrew, for none of them had suspected that such great importance
was even now attached to the stone.
“How did he know of the crystal?” asked the King.
“Every one in the east knows of the Great Tarnov Crystal,”
answered Bogdan. “Every worker of magic, every astrologer, every
chief, every prince, is desirous of possessing this treasure. For it is
said that in addition to being a jewel of great value it has this quality
also, that one may look into it and there read of the future—one may
also find there secrets of great worth, one may see the faces of men
long since in their graves. There are many legends and stories of it,
too, and since the days when it disappeared from Tarnov, when the
Tartars conquered Western lands, there has been search after
search to find it.”
The King thought for a few seconds. “Then the Khan of the
Tartars knew that he was asking Ivan for an impossibility when he
demanded the crystal? Does that mean that he meant to refuse to go
against the Poles?”
“Please—your Majesty—it was no such thing,” Bogdan stated
emphatically. “A short time ago a servant who had left the services of
this man here,” he pointed to Pan Andrew, “went to the land of the
Tartars and there spread the report about, that the crystal was to be
had for the taking, that it was hidden in a country house in the
Ukraine. You may be sure that this reached the ears of the Khan,
whose passion for curious jewels is almost a madness, and when I,
going from Ivan to Tartary, learned this, then Ivan promised the Khan
that he would get him the crystal if it could be gotten.”
“You were the go-between?”
Bogdan bowed.
“And Ivan sent you to get it from Pan Andrew?”
Bogdan bowed, though not quite so low.
Fire leaped into the King’s eyes. “Dog that you are,” he said.
“Less than beast in all things that Christians believe; for this you
must destroy a man’s house and ruin his fields, yes, and threaten his
life, too, if it would serve your purpose. . . . God knows, my kingly
duties lie heavily upon me. . . . All that I seek in this, my
commonwealth, is peace, peace with my neighbors and happiness
for my people. And yet Poland is ever insulted to the point where
nothing but war is possible. It is not enough that enemies on the
north and west threaten, there must be plots against our happiness
on the south and east. Oh, Poland, Poland, when will the day come
that thy sons and daughters may enjoy the tranquillity that God has
designed for all people? . . . As to you,” he turned again to Bogdan,
“what further have you to say?”
 “Only that I have failed,” answered Bogdan miserably. “And only
that I know that I shall go free, for there was never yet Jagiello who
did not keep his word. Though had it not been for this creature
here”—he pointed to the alchemist, who from the rear of the room
had been watching the scene through half-shut eyes—“I should have
had the crystal long ago.”

The King did not reply. “Take him away,” he said to a guard.
A captain in armor came forward. “Deliver this Bogdan at sunrise
to the guards of the Florian Gate. Tell them to see that he has safe-
conduct through to the border, but that his chains are not to be struck
off until he reaches the frontier. After that let happen what will, but if
he so much as sets foot again upon Polish soil he shall be hanged to
the nearest tree.”
When they had departed, he said to Pan Andrew:
“In this my right and duty of kingship in the Commonwealth of
Poland I commend you most heartily as a man who has been of
great service to his country. It is a most extraordinary and gracious
thing that a family such as yours should be so faithful to its word
through so many years and be willing to suffer so much for an oath
once given. Therefore to you go my whole thanks.”
He took the gold chain from his throat. It was a thing of wondrous
beauty, of heavy solid links cut out of the purest metal.
“Wear this,” he said placing it with his own hands over Pan
Andrew’s shoulders. “This chain shall ever be to you the token of
your faithfulness. I shall see to it that the state makes return to you
for the property which you have lost, for in so losing it you have
conferred a favor upon us all. Had the crystal been taken by these
thieves and delivered to the Khan of the Tartars it is probably true
that by now the Ukraine had begun to be overrun by Tartars and the
armies of Ivan. In due time I shall see to it that a more formal and
proper reward is given you.”
Here Jan Kanty made a sign that the interview was finished and
the whole company fell upon their knees before the King.
He, too, stooped, but only to pick up the crystal which had lain
upon the floor before him during the entire interview. It seemed to
Joseph, glancing up at that moment, that the instant the King’s eyes
were fixed upon the stone he became suddenly oblivious to
everything else that was before him, and stood as one in a dream or
trance gazing into the depths of the fearsomely beautiful thing.