Discovery of a new gall-inducing species, Aciurina luminaria (Insecta, Diptera, Tephritidae) via multi-trait integrative taxonomy

Abstract

Integrative taxonomic practices that combine multiple lines of evidence for species delimitation greatly improve our understanding of intra- and inter-species variation and biodiversity. However, extended phenotypes remain underutilized despite their potential as a species-specific set of extracorporeal morphological and life history traits. Primarily relying on variations in wing patterns has caused taxonomic confusion in the genus Aciurina, which are gall-inducing flies on Asteraceae plants in western North America. However, species display distinct gall morphologies that can be crucial for species identification. Here we investigate a unique gall morphotype in New Mexico and Colorado that was previously described as a variant of that induced by Aciurina bigeloviae (Cockerell, 1890). Our analysis has discovered several consistent features that distinguish it from galls of A. bigeloviae. A comprehensive description of Aciurina luminaria Baine, sp. nov. and its gall is provided through integrative taxonomic study of gall morphology, host plant ecology, wing morphometrics, and reduced-representation genome sequencing.

Full text

217
Discovery of a new gall-inducing species, Aciurina luminaria
(Insecta, Diptera, Tephritidae) via multi-trait integrative taxonomy
Quinlyn Baine1, Branden White1, Vincent G. Martinson1* , Ellen O. Martinson1*
1 Department of Biology, University of New Mexico, 219 Yale Blvd, Albuquerque, NM 87131, USA
Corresponding author: Quinlyn Baine (quinbaine@gmail.com)
Copyright: © Quinlyn Baine et al.
This is an open access article distributed under
terms of the Creative Commons Attribution
License (Attribution 4.0 International – CC BY 4.0).
Research Article
Abstract
Integrative taxonomic practices that combine multiple lines of evidence for species de-
limitation greatly improve our understanding of intra- and inter-species variation and bio-
diversity. However, extended phenotypes remain underutilized despite their potential as a
species-specic set of extracorporeal morphological and life history traits. Primarily rely-
ing on variations in wing patterns has caused taxonomic confusion in the genus Aciurina,
which are gall-inducing ies on Asteraceae plants in western North America. However,
species display distinct gall morphologies that can be crucial for species identication.
Here we investigate a unique gall morphotype in New Mexico and Colorado that was
previously described as a variant of that induced by Aciurina bigeloviae (Cockerell, 1890).
Our analysis has discovered several consistent features that distinguish it from galls of
A. bigeloviae. A comprehensive description of Aciurina luminaria Baine, sp. nov. and its
gall is provided through integrative taxonomic study of gall morphology, host plant ecol-
ogy, wing morphometrics, and reduced-representation genome sequencing.
Key words: Bigeloviae, candle, ddRAD, Ericameria, ame, marshmallow, nauseosa,
rabbitbrush, tephritid, trixa, wing
Introduction
Species delimitation is an essential step in our collective goal as biologists to cal-
culate the total diversity of life on the planet (Dayrat 2005), and is particularly vital
in the midst of ongoing decline of insects – the world’s most species-rich group
of animals (Dirzo et al. 2014; Stork et al. 2015; Sánchez-Bayo and Wyckhuys
2019). However, species divisions are frequently unclear, particularly where pure-
ly morphological descriptions include high levels of intra-species variation, which
poses a signicant challenge in taxonomy, as it can result in unreliable diagnosis
of species boundaries (Gentile et al. 2021). Most descriptions of insect species
are made from adult morphological characters alone and are presented as a list
of traits that have some author-determined signicance in recognition (e.g., col-
oration, wing patterns, integumental texture). This issue can be addressed by
combining multiple lines of evidence, such as morphological, ecological, and
geographical data, to make more accurate and robust species delimitations;
a long-standing practice coined in recent decades as “integrative taxonomy”
Academic editor: Marc De Meyer
Received:
2 July 2024
Accepted:
27 August 2024
Published:
7 October 2024
ZooBank: https://zoobank.
org/317A7E22-B53E-4A35-9065-
F5CB66180350
Citation: Baine Q, White B,
Martinson VG, Martinson EO (2024)
Discovery of a new gall-inducing
species, Aciurina luminaria (Insecta,
Diptera, Tephritidae) via multi-trait
integrative taxonomy. ZooKeys 1214:
217–236. https://doi.org/10.3897/
zookeys.1214.130171
ZooKeys 1214: 217–236 (2024)
DOI: 10.3897/zookeys.1214.130171
* Contributed equally to this work.

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ZooKeys 1214: 217–236 (2024), DOI: 10.3897/zookeys.1214.130171
Quinlyn Baine et al.: Aciurina luminaria sp. nov.
(Schlick-Steiner et al. 2010). Taxonomists are also now equipped with molecular
tools that can provide deep genome-wide datasets to investigate intra-taxon dis-
tinctions, even in cryptic or rare groups of arthropods (Hebert et al. 2003; Sheikh
et al. 2022). An integrative taxonomy approach to species description improves
our overall species estimates and identication of signicant radiations in evolu-
tionary history. By embracing a holistic approach, integrative taxonomy allows for
a more nuanced understanding of biodiversity, leading to more precise species
identication (Schlick-Steiner et al. 2010). This not only enhances our knowledge
of the natural world but also is crucial for conservation efforts, as accurately iden-
tifying species is foundational to protecting them and their habitats.
Though many have adopted integrative taxonomic description, a potentially
powerful tool for species delimitation remains under-utilized: the extended phe-
notype. This refers to an organism’s genetic expression that can be observed
beyond their own bodies, particularly in the case of animals that construct or
modify unique structures such as bird nests and spider webs (Blamires 2013;
Mainwaring et al. 2014). These extensions of the phenotype represent spe-
cies-specic behaviors and adaptations, establishing them as a key piece of the
species’ ecology, and a set of additional morphological traits that can be used in
species delimitation (Bailey et al. 2009; Freudenstein et al. 2016). For example,
gall-inducing insects create structures that are so frequently species-specic
that they can be used as a diagnostic character for identication (Raman et al.
2005; Bailey et al. 2009; Redfern 2011; Russo 2021). Integrating extended phe-
notypes in species description will likely enhance the resolution of taxonomic
classications, especially with ecosystem engineers like gall-inducing insects.
The genus Aciurina (Diptera: Tephritidae) are gall-inducing flies on Asterace-
ae shrubs in western North America (Foote et al. 1993). Many species in this
genus are informally recognized by gall morphological characters, and, simi-
larly to many tephritid “picture-wing” ies, formally identied with diagnostic
black and transparent markings of the wings. The common and widespread
species Trypeta bigeloviae was rst described only as a “white, woolly [sic], and
conspicuous” gall on the plant Bigelovia (Cockerell 1890a). The y was then
described in the same year as both T. bigeloviae and T. bigeloviae var. disrupta
based on a single distinction in the postero-distal hyaline region of the wing: in
disrupta this area is divided (disrupted) into two by a complete black marking
(Cockerell 1890b). Bates (1935) later re-assigned T. bigeloviae to the genus
Aciurina, including the variety disrupta which he did not consider distinct, citing
variation of this character in specimens from the same locality. Wing marking
variation led Steyskal (1984) in his revision of Aciurina to then synonymize the
type species Aciurina trixa Curran, 1932 and Aciurina semilucida Bates, 1935
with Aciurina bigeloviae (Cockerell, 1890), which he characterized as being the
most variable species of the genus. However, Dodson and George (1986) soon
after described the likely recently diverged relationship of A. bigeloviae and A.
trixa based on thorough examination of gall morphology, host plant ecology,
hybrid breeding success, and genetic allelic frequencies. First Goeden and
Teerink (1996) reinstated A. semilucida, then Headrick et al. (1997) ocially
reinstated A. trixa as a species distinct from A. bigeloviae and provided a larval
description. The most reliable diagnostic character established between the
sister species was gall morphology: A. bigeloviae has a white, wooly “cotton”
gall and A. trixa has a resinous, waxy “smooth” gall. Both species form galls on

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Quinlyn Baine et al.: Aciurina luminaria sp. nov.
Ericameria nauseosa (Pall. ex Pursh) G.L.Nesom & G.I.Baird, however on differ-
ent varieties of the species.
Dodson and George (1986) also described in detail the confusing variation
in wing patterns noted by the other authors above by establishing three pattern
categories among the two species: 1) the Type I pattern that included the orig-
inally described wing pattern for A. bigeloviae plus that of var. disrupta, 2) the
Type II pattern which matched the description of A. trixa, and 3) the Type II’ pat-
tern (hereafter referred to as Type III) as a modied version of the A. trixa wing
pattern, but from ies reared from A. bigeloviae-type cotton galls (Fig. 1). The
authors admit that this third pattern, with its unexpected gall-wing morpholog-
ical pairing, left them somewhat stumped: “Whether they belong to bigeloviae,
trixa, or a third species probably will not be resolved until further studies parallel
to those reported here are carried out” (Dodson and George 1986).
Here we follow this thread and using an integrative taxonomic approach that
employs gall morphology, previously unexplored host plant ecology, extensive
wing morphometric and character analysis, and multi-locus reduced represen-
tation genome sequencing, provide evidence that the Type III ies are a third
species. We provide a name for this species, Aciurina luminaria, and a complete
morphological description of the adult y and its gall.
Materials and methods
We observed in previous collections of A. bigeloviae that “cotton” galls in New
Mexico could be categorized into two groups by general gall shape: spherical and
teardrop-shaped (Fig. 2). We were able to conrm from rearing haphazardly col-
lected galls that the spherical cotton galls were induced by ies with Type I wing
morphology (A. bigeloviae), and the teardrop-shaped galls were induced by ies
with Type III wing morphology. The ability to recognize the different gall morphs
in the eld allowed us to perform targeted collections for each morphotype.
We systematically collected and reared A. bigeloviae and A. trixa galls in
New Mexico between 2021–2022 following methods outlined in Baine et al.
(2023a). Type III galls were collected haphazardly from identied populations
throughout New Mexico and Colorado in April and May of 2021–2023 by clip-
ping sections of stem with gall attached and transporting them to the University
of New Mexico. In the lab, galls were placed in insect rearing cages (BugDorm)
and kept at 45% relative humidity. Adult ies were removed from cages as they
emerged and preserved in 100% EtOH at -20 °C for the following analyses. A
subset of pinned ies and galls was examined for morphological characters
and photographed using an EOS 40D camera tted with a 65 mm MP-E macro
photo lens (Canon) mounted on Stackshot macro rail with controller (Cogni-
sys), and then focus stacked with Zerene Stacker software. Representative ies
and galls of each sampled population, plus other material examined, including
the holotype of A. luminaria, were deposited in the following collections: The
Museum of Southwestern Biology, Arthropods Division, New Mexico (MSBA);
Smithsonian Institution, National Museum of Natural History, Washington DC
(USNM); and William F. Barr Entomological Museum, Idaho (WFBM).
Aciurina species are frequently documented as specialists on particular
varieties of E. nauseosa. For example, A. bigeloviae is associated with E. n.
subsp. nauseosa var. graveolens, and A. trixa in New Mexico is associated

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ZooKeys 1214: 217–236 (2024), DOI: 10.3897/zookeys.1214.130171
Quinlyn Baine et al.: Aciurina luminaria sp. nov.
with E. n. subsp. nauseosa var. latisquamea (Dodson and George 1986). To
determine the host plant identity of Type III galls, we additionally returned to a
subset of sites during the owering season in the fall of 2022 and 2023 to ob-
tain plant voucher specimens. Plant samples were identied using Anderson
(2006) and Allred and Jercinovic (2020), and deposited in the MSB Herbarium.
Wing morphology
A total of 62 female Aciurina specimens of the three morphotypes from 19 pop-
ulations were selected for morphological assessment. Both wings of each spec-
imen were carefully removed by pulling at the connection point where the tegula
meets the thorax. Wings were mounted on glass slides with a Euparal mounting
medium (Hempstead Halide). The edges of the cover slide were then sealed with
clear nail polish and slides were left to dry at room temperature for 24 hours. Im-
ages of each wing were taken using a Axiocam 208 color camera mounted on a
Stemi 508 microscope (Zeiss). Measurements were taken in ZEN 3.6 blue edition
(Zeiss). The methodology used to take standardized proxy measurements of the
wing width (represented by distance from apex of vein R1 to junction of vein M4
and crossvein dm-m) and length (represented by distance from junction of vein
M4 and crossvein bm-m to apex of vein M4) follows Baleba et al. (2019). A total
of seven different measurements and three different categorical observations
were made for each wing (Fig. 1A). To facilitate comparison without inuence of
difference in overall body size, each measurement was divided by our measure-
ment for wing length. Finally, measurements from each wing pair were averaged.
We then compared morphotype III to both morphotype I and II using each set
of measurements by analysis of variance (ANOVA, aov) or Kruskal-Wallis where
assumptions for ANOVA were not met (kruskal.test), and each set of categorical
variables by Pearson’s 𝛘2 test (chi.test) in R version 4.2.2 (R Core Team 2021).
The terminology we used for venation and cells follows Cumming and Wood
(2017), and our selected measurements and categorical variables are dened
as follows (Fig. 1A):
• brL: The maximum diameter of the subapical hyaline spot of cell br. This
spot has a circular-elliptical shape, so the measurement typically follows
a line from one elliptical-vertex to the other.
• rrL: The maximum width of the hyaline region located within cells r1 and
r2+3, crossing vein R2+3: measurement taken from the midpoint of the vein
C to the parabolic vertex of the shape. This pattern occasionally reaches
vein R4+5; in this case the midpoint that borders this vein is used instead of
a parabolic vertex.
• rrH: The maximum length of the same region hyaline in cells r1 and r2+3,
measured along vein R2+3.
• mrL: The maximum length of the subapical hyaline region in cells r4+5 and
m: length taken from the postero-distal corner to the parabolic vertex of
the region.
• C1: The presence or absence of a complete medial stripe within the anal
lobe from veins CuA+CuP to the posterior wing margin.
• C2: The presence or absence of a complete medial stripe within cell m4
from vein M4 to the posterior wing margin.

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Quinlyn Baine et al.: Aciurina luminaria sp. nov.
Figure 1. Wing morphotypes and measurements taken A diagram of characters and measurements dened and used in
analysis B A. bigeloviae C A. bigeloviae D A. trixa E A. luminaria sp. nov. Not to scale.
• C3: The extent of black medial stripe in the large subbasal hyaline region
on the posterior margin of the wing within cell m. Three conditions exist:
stripe absent, stripe incomplete, and stripe completely bisecting the re-
gion (var. disrupta morphology).
Genomics
From three populations each of A. bigeloviae, A. trixa and Type III, we extracted
whole-body DNA from three replicates (total n = 27) using the DNeasy Blood &
Tissue kit and protocol (Qiagen), and quantied nucleic acid with a Qubit 3.0
uorometer (Invitrogen). We generated genotypes for each sample from sin-
gle-nucleotide polymorphisms (SNPs) derived from double digest restriction site
associated DNA sequencing (ddRADseq) (Peterson et al. 2012). DNA was di-
gested with enzymes EcoRI and MseI, and fragments were coupled with Illumina
adaptors through T4 ligation. Pooled fragments were used for PCR with a proof-
reading enzyme (Iproof; BioRad), and fragment size selection for 300–450 bp

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ZooKeys 1214: 217–236 (2024), DOI: 10.3897/zookeys.1214.130171
Quinlyn Baine et al.: Aciurina luminaria sp. nov.
Figure 2. Galls of A. luminaria and A. bigeloviae A immature A. luminaria galls B internal view of immature A. luminaria gall
with early instar larva burrowing into the stem while gall develops C mature A. luminaria galls D internal view of mature A.
luminaria gall with full-sized larval chamber E mature A. bigeloviae galls F internal view of A. bigeloviae gall with a mature
larva in the larval chamber. Side by side comparison of A. bigeloviae (left) and A. luminaria (right) tomentum texture and
uniformity G external view H internal view.

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Quinlyn Baine et al.: Aciurina luminaria sp. nov.
was performed using a Pippin Prep quantitative electrophoresis unit. An Illumina
NovaSeq S2 housed at the University of Texas at Austin Genomic Sequencing
and Analysis Facility (Austin, TX) generated sequences of ~100 bp from input
fragments. Trimmomatic 0.39 was used to truncate and lter demultiplexed sin-
gle-end reads to a threshold of 85 bp (Bolger et al. 2014). Sequence les are
deposited in NCBI GenBank under BioProject ID PRJNA1075688.
We mapped trimmed reads de novo using the Stacks 2.61 (Catchen et al.
2011) wrapper denovo_map.pl with the selected parameters: 5 minimum reads
per stack, 3 maximum mismatches per locus, 3 maximum mismatches per
stack, and minimum 80% individuals required per locus (parameter optimization
was performed generally following Paris et al. 2017). We then ltered stacks in R
with SNPltR (DeRaad 2022) and the settings: maximum depth of 50, maximum
missing per sample 80%, minimum SNP completeness 80%, and minimum mi-
nor allele count of 1. Filtered SNP loci were used for principal component analy-
sis (PCA) using packages vcfR (Knaus and Grünwald 2017), dartR (Gruber et al.
2018) and adegenet (Jombart 2008). We also calculated π for each population
as a measure of nucleotide diversity. Structure was estimated using sparse non-
negative matrix factorization (sNMF) with package LEA (Frichot and Francois
2015). For each K value, where K represents the number of clusters ranging from
1 to 9, we performed 50 iterations. We then selected and graphed the best result
from the most optimal K value, as identied through the cross-entropy criterion.
SNPs per sample were concatenated to generate sequences for each indi-
vidual, and sequences were aligned in Stacks. Phylogeny was inferred by max-
imum likelihood (ML) tree with IQ-TREE 2.2.0 (Minh et al. 2020). ModelFinder
(Kalyaanamoorthy et al. 2017) was used to select the best model as deter-
mined by Akaike Information Criterion (AIC). We calculated branch support
from 1000 ultrafast bootstrap (UFBoot) (Hoang et al. 2018) and 1000 Shimo-
daira–Hasegawa approximate likelihood ratio test (SH-aLRT) (Guindon et al.
2010) replications. We report the selected model and basal branch support
values. Sequence reads used in all above genomic analyses are deposited in
NCBI GenBank within BioProject PRJNA1075688.
Results
Wing morphology
Morphometric comparisons of A. bigeloviae and A. trixa with the Type III mor-
photype highlighted regions of the wing that differ signicantly in relative size
and can therefore be used as diagnostic characters. The greater length of the
hyaline spot within cell br (measurement brL) in Type III is the most notably
distinct wing measurement from the other two morphotypes (A. bigeloviae 𝛘2=
20.57, p < 0.0001; A. trixa 𝛘2 = 20.83, p < 0.0001). Type III also differs in the
dimensions of the hyaline spot in r1 and r2+3, being longer than in A. bigeloviae
(measurement rrH, F = 30.61, p < 0.0001), and wider than in A. trixa (measure-
ment rrL, F = 7.99, p < 0.01). Finally, the measurement mrL is slightly greater in
A. trixa than in Type III (F = 4.44, p < 0.05).
Despite variation, A. bigeloviae wings had the darkened stripes represented
by the selected characters consistently present, while in Type III they were ab-
sent (Table 1). The absence of the dark stripe within the hyaline region in cell

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ZooKeys 1214: 217–236 (2024), DOI: 10.3897/zookeys.1214.130171
Quinlyn Baine et al.: Aciurina luminaria sp. nov.
m (C3) that frequently separates A. trixa from A. bigeloviae, also separates the
Type III morphotype from A. bigeloviae, but not from A. trixa. See Suppl. material
1: g. S1 for wing character variation in each of the three morphotypes sampled.
Table 1. Signicance values from 𝛘2 tests to compare the presence/absence of wing
characters in the three morphotypes.
Wing pattern character A. bigeloviae ~ Type III A. trixa ~ Type III
C1 𝛘2 = 50.22, df = 1, p < 0.0001 𝛘2 = 36.29, df = 1, p < 0.0001
C2 𝛘2 = 68, df = 1, p <0.0001 𝛘2 = 65.33, df = 1, p < 0.0001
C3 𝛘2 = 47.28, df = 2, p < 0.0001 NS
Genomics
From the ddRAD sequencing of the three morphotypes, 42,838 SNPs were re-
tained after ltering. A single sample was ltered out for high relative missing-
ness, so we used 26 samples total for the following analyses. The rst principal
component (PC1) in the PCA performed accounted for 42.3% of the variance,
splitting the morphotypes into discrete clusters that match to gall morpholo-
gy (Fig. 3A). The second principal component (PC2) explained an additional
21.2% of the variance, splitting Type III from A. bigeloviae, and also with mini-
mal loadings on geographic variation within A. bigeloviae populations (Fig. 3A).
The PCA represents the greatest amount of variance exists between the three
morphotypes. Average genetic diversity across all sites (π) was lower in Type III
(0.0008) than in A. bigeloviae (0.0014) or A. trixa (0.0011) across all populations
(0.0775, 0.1324, 0.1083, respectively across variant sites). Genetic diversity
was also lower by a comparable margin in Type III across the populations at the
sympatric site for all three morphotypes, “SNM” (0.0767 versus 0.12 A. bigelo-
viae and 0.1006 A. trixa). The most optimal K for structure-like sNMF analysis
was 3, and plots reveal that there is virtually no admixture present between the
morphotypes, even in the populations at the sympatric sites (Fig. 3C).
Finally, to construct the phylogeny, we selected with ModelFinder a trans-
version substitution model of AG=CT and empirical unequal base frequencies,
plus a FreeRate model (Yang 1995; Soubrier et al. 2012) for rate heterogeneity
across sites with 2 categories, and an ascertainment bias correction appropri-
ate for SNP alignments (TVM+F+ASC+R2). The output tree supports the pat-
tern observed in the results above, that the morphotypes are consistently dis-
tinct, and that variation is lower in A. bigeloviae (Fig. 3B). The tree also suggests
that Type III is more closely related to A. bigeloviae than A. trixa is, which may
mean that their shared gall morphological traits are ancestral.
Taxonomy
Aciurina luminaria Baine, sp. nov.
https://zoobank.org/BC9BB3A4-E1CC-4793-AC8D-B8A4E779FFBF
The Candle Flame Gall Tephritid
Type material examined. Holotype (Fig. 4A, C) USA • NM; Santa Fe Co.; ♀; Tesuque
E arroyo crossing Road 74; 35.77686°N, 105.92904°W; 5 May 2021; Q. Baine leg.;
MSBA81957. Allotype (Fig. 4B, D) USA • NM; Santa Fe Co.; ♂; 1 mi SE Chupadero

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ZooKeys 1214: 217–236 (2024), DOI: 10.3897/zookeys.1214.130171
Quinlyn Baine et al.: Aciurina luminaria sp. nov.
RGG
MAL
SNM
RGG
RGG
SNM
MAL
MAL
MAL
SNM
MAL
MAL
SNM
SNM
TES
0.02
A. bigeloviae
A. luminaria
TES
TES
SNM
SNM
SNM
TES
TES
TES
RGG
RGG
RGG
A. trixa
100/100
100/100
100/100
100/100
−40
−20
0
20
−50 −25 025
PC1 (42.3%)
PC2 (21.2%)
RGG
MAL
MAL
SNM
SNM
TES
RGG
SNM
TES
0.00
0.25
0.50
0.75
1.00
SNM MALTES
SNM
MAL RGG
SNM TESRGG
AB
C
Figure 3. Results from SNP comparisons across populations (labeled as three-letter abbreviations) of the three morpho-
types (labeled by color) A PCA plot B Maximum likelihood tree with basal support values UFBoot + SH-aLRT C sNMF
structure-like admixture plot.
off St Rd 592; 35.814°N, 105.907426°W; 7 May 2023; Q. Baine leg.; MSBA81946.
Paratypes USA • CO; Alamosa Co.; 2♀ 1♂; S of Mosca side of Hwy 17; 37.62534°N,
105.86636°W; 20 May 2021; V. Martinson leg.; MSBA81880–81882 • 3♀ 3♂ 1
gall; San Luis State Wildlife Area, Lane 6 N; 37.66256°N, 105.72293°W; 20 May
2021; E. Martinson & V. Martinson leg.; MSBA81892–81893, MSBA81896–
81897, USNMENT02011093–USNMENT02011095 • 4♀ 4♂; San Luis State
Wildlife Area, Lane 6 N; 37.66256°N, 105.72293°W; 14 May 2023; Q. Baine leg.
MSBA81926–81933 • 7♀ 6♂; W of San Luis Valley Regional Airport entrance, Ala-
mosa; 37.4442°N, 105.86753°W; 14 May 2023; Q. Baine leg.; MSBA81898–81901,
USNMENT02011096–USNMENT02011097 • 2♀ 3♂; 2 mi S Zapata Falls turnoff
Hwy 150; 37.59092°N, 105.6015°W; 15 May 2023; Q. Baine leg.; MSBA81909–
81913. 2♀ 1♂; Road N 110 and Lane 1 N off Hwy 17; 37.59092°N, 105.6015°W;
15 May 2023; Q. Baine leg.; MSBA81914–81916 • 2♀ 1♂; Corner of Cortez Rd

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Quinlyn Baine et al.: Aciurina luminaria sp. nov.
Figure 4. Aciurina luminaria sp. nov. A holotype lateral habitus B allotype lateral habitus. Difference in eye color is a result
of the age of mounted specimen C holotype dorsal habitus D allotype dorsal habitus E holotype head, anterior F holotype
abdomen, dorsal G–H variation in dorsal abdomen color G mostly orange morph H dark morph.

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Quinlyn Baine et al.: Aciurina luminaria sp. nov.
and Van Iwarden Dr, Alamosa; 37.43771°N, 105.88511°W; 14 May 2023; Q. Baine
leg.; MSBA81922–81925. Chaffee Co. • 2♀ 2♂; Hwy 285 W of Johnson Village;
38.80956°N, 106.11603°W; 20 May 2021; E. Martinson leg.; MSBA81883–81886
• NM; Cibola Co.; 2♀ 2♂; El Malpais National Conservation Area Narrows;
34.96499°N, 107.81464°W; 15 May 2022; V. Martinson leg.; MSBA81876–81879.
San Juan Co. • 1♂; S Bloomeld Hwy 550 Kutz Wash; 36.64524°N, 108.00264°W;
18 March 2022; E. Martinson leg.; MSBA81891. Sandoval Co. • 4♀; N side Hwy
550, La Jara; 36.0581873°N, 106.9749619°W; 31 May 2020; D. Hughes leg.;
MSBA81887–81890. Santa Fe Co. • 5♀ 4♂, 1 gall; Tesuque E arroyo crossing
Road 74; 35.77686°N, 105.92904°W; 7 May 2023; Q. Baine leg.; MSBA81934–
81938, MSBA81941–81942, USNMENT02011090–USNMENT02011092 • 2♀;
Tesuque E arroyo crossing Road 74; 35.77686°N, 105.92904°W; 5 May 2021;
Q. Baine leg.; MSBA81956–81957 • 4♀ 2♂; 1 mi SE Chupadero off St Rd 592;
35.814006°N, 105.907426°W; 7 May 2023; Q. Baine leg.; MSBA81943–81948 • 3♀
4♂; Tesuque Arroyo Ancho and Meredith Dr on Tesuque Village Rd; 35.752991°N,
105.934346°W; 7 May 2023; Q. Baine leg.; MSBA81949–81955. Taos Co. • 2♀
3♂; 3 mi N Ojo Caliente off Hwy 285; 36.33046°N, 106.00581°W; 14 May 2023; Q.
Baine leg.; MSBA81917–81921 • UT; Kane Co.; 1♀ 1♂; Coral Pink Sand Dunes; 3
July 1966; E.J. Allen leg.; WFBM0050980–0050981.
Other material examined. USA • NM; Alamosa Co.; 4 galls, 1 pupa; S Nageezi
side of Pueblo Pintado Rd; 36.21480°N, 107.69633°W; 23 May 2024; S. Rollins
leg • 4♀ 3♂; Corner of Cortez Rd and Van Iwarden Dr, Alamosa; 37.43771°N,
105.88511°W; 14 May 2023; Q. Baine leg. • 3 larvae; San Luis State Wildlife
Area, Lane 6 N; 37.66256°N, 105.72293°W; 20 May 2021; E. Martinson leg.
Diagnosis. This wing pattern of the adult A. luminaria can be distinguished most
easily from both A. bigeloviae and A. trixa by the elongate hyaline spot in cell br,
consistent dark brown region surrounding crossvein r-m, and lack of dark stripe
in anal cell; it further from A. bigeloviae by lack of dark stripe in the postero-distal
region of cell m and lack of medial dark stripe in cell cua1 (frequently present in A.
trixa also). It differs from the similar-looking A. maculata (Cole, 1919) and A. lutea
(Coquillett, 1899) by the hyaline cell bc and hyaline basal region of cell br. The ex-
tent of bright orange on the abdomen of many A. luminaria specimens also distin-
guishes it from A. maculata which has a more red abdomen, and from A. bigeloviae
and A. trixa which frequently have a dark orange, brown, or black abdomen. Gen-
italia structures are highly similar to that of A. bigeloviae, except perhaps for the
rounded tips of the prensisetae which differ from illustrations in Steyskal (1984).
However, Steyskal describes A. bigeloviae (at the time synonymized with A. trixa
and A. semilucida) as being highly variable in male terminalia characters, so this
may or may not be reliably diagnostic. The gall can be distinguished from A. bige-
loviae and A. maculata by the pointed, teardrop shape, and from all remaining galls
in the genus by the thick layer of dense tomentum covering the surface (Fig. 2).
Description. Female body length (minus terminalia) 6 mm.
Head (Fig. 4E) uniformly pale yellow except for occiput and narrow intero-
cular margin grey and moderately pilose. Compound eye bright green, drying
to dull red. Three pairs of frontal setae, two pairs of orbital setae, and one pair
of ocellar setae present. All setae pale yellow in color matching frons in color.
Antenna yellow with black arista.
Thorax. Scutum and dorsal portions of pleura dark gray in background color with
pale gray pollinosity and dense pale yellow setulae making the scutum appear pale

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Quinlyn Baine et al.: Aciurina luminaria sp. nov.
yellow-gray in color at a distance. Scutellum pale orange-brown at apex, narrowly
gray at base. Subscutellum with anterior half pale yellow, posterior half and all of
mediotergite black with pale gray pollinosity. Ventral part of pleura yellow-orange.
The following setae are present, and pale yellow: basal scutellar, postalar, intra-alar,
acrostichal, postsutural dorsocentral, presutral supra-alar, postsutural supra-alar,
two notopleural, postpronotal, anepisternal, and katepisternal. Anepimeral seta in-
distinguishable from surrounding setulae. Legs wholly orange in color except for
black tarsal claw and apical tarsal setae. Forefemur with elongate comb-like setae.
Wing 4.2 mm in length. Costa pale orange. Setae narrowly present dorsally at junc-
tion of R2+3 and vein R4+5. Wing coloring is dark brown to black with the following
hyaline regions: cell bc, base of cell br, two vertical bands in cell c, the proximal one
extending posteriorly halfway into cell bm, two marginal spots in r1 with apical spot
extending into r2+3, large (2× wide as high) subapical spot in cell br, medial spot in
cell bm, large basal and small apical spot in cell cua1, subapical spot (1.5× high as
wide) in cell dm, entire cell cup except for narrowly at apex, alula, anal lobe, large
basal marginal spot in cell m, and subapical band extending from posterior margin
in cell m into cell r4+5 reaching vein R4+5. Halteres bright yellow.
Abdomen bright red-orange and shiny. Oviscape wholly black and shining.
Eversible membrane brown, with shallowly semicircular cuticle denticles. Ac-
uleus short (0.8 mm), notched at basal edge. Apical one third of aculeus with
minute denticles covering medial edge (Fig. 5A).
Male body length (minus terminalia) 4 mm. Matching female in all respects
except for terminalia. Epandrium black and shining, and proctiger pale yel-
low-orange. Surstylus pale brown, and prensisetae paired, bluntly rounded at
the tips, and black. Phallus (1.25 mm long) and glans dark brown (Fig. 5B).
Variation. Ventral thoracic pleura (including episternum, meron, anatergite
and katatergite) in darker morphs are black with gray pollinosity, as on the scu-
tum. Abdomen color ranges from wholly orange, orange with black tergite 6 (5
in male), orange with lateral black spots on tergites 5 and 6 (4 and 5 in male), to
mostly black with orange background in dark morphs of both sexes (Fig. 4F–H).
Immature. Second instar larva: Body white, elliptical-oblong and rounded on
both anterior and posterior ends. Body segmented by rows of acanthae. Gna-
thocephalon conical and generally smooth. Mouth hook black and bidentate.
Posterior spiracular plate with three pale brown rimae. Puparium: length 4.00
mm, width 1.62 mm. Dark brown, shining, elliptical-oblong, and rounded on
both anterior and posterior ends. Anterior end with invagination scar and ante-
rior thoracic spiracle. Posterior spiracular plate with spiracle darkened and at.
Gall relatively large at maturity (7.24 mm mean latitudinal diameter), has a
mostly rounded oblong to tapered teardrop shape and is covered uniformly in
dense off-white cottony tomentum (Fig. 2C).
Biology. Aciurina luminaria is univoltine and has a life cycle and phenology
similar to A. bigeloviae and A. trixa (Baine et al. 2023a; 2023b). Eggs are laid
singly into the leaf bud of a distal plant stem. The gall forms at the oviposition
site and the developing larva feeds on the tissue surrounding the central cham-
ber of the gall. By fall, the gall reaches full size, and the larva reaches its nal
instar and chews to the outer layer of the gall to create a circular trap door. The
larva ceases feeding and overwinters inside the gall, then pupates in the spring.
Adults eclose in summer and push their way through the door to emerge from
the gall and nd a mate. The period of emergence from galls reared by these

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Quinlyn Baine et al.: Aciurina luminaria sp. nov.
authors is May 29th – June 28th. The latest date of emergence in examined ma-
terial from Utah is July 18th (gall collected 3 July 1966).
Associated arthropods. The most common parasitoid by far is Eurytoma
chrysothamni (Hymenoptera: Eurytomidae). We reared very few unidentied Hal-
ticoptera, Pteromalus (Pteromalidae), and Torymus (Torymidae) wasps, which
may be the same species as those associated with A. bigeloviae (Baine et al.
2023a). We also observed and reared a small number of Rhopalomyia (Diptera:
Cecidomyiidae) hypergalls on the surface of galls collected in northwestern
New Mexico. The hypergall system has been previously documented on both A.
bigeloviae (Baine et al. 2023b) and A. trixa (Russo 2021), but whether the midge
species is the same is unknown. Unexpectedly, and unknown from other Aciuri-
na systems, a single Synergus (Hymenoptera: Cynipidae) wasp was also reared.
Host plant. The known host plant is strictly Ericameria nauseosa subsp. am-
mophila L.C. Anderson, which was described from the San Luis Valley in Colora-
do (Anderson 2006). This plant is restricted to sandsheet and sand dune habitat
and is known from southern Colorado (Anderson 2006) and here we add to its
range northern New Mexico. Floral specimens from galled plants in New Mexico
are deposited in MSB Herbarium (UNM0143677–UNM0143682). The host plant
of material from Pink Sand Dunes, Utah is only identied as “Chrysothamnus nau-
seosus” [sic] and the host plants of gall observations in Arizona are unidentied.
Figure 5. Aciurina luminaria terminalia A female B male.

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Quinlyn Baine et al.: Aciurina luminaria sp. nov.
Geographical range. Beyond the localities of the examined material above, we
have conrmed the presence of this species in some locations reported by Dod-
son and George (1986): Great Sand Dunes National Monument, CO, and near the
cities Grants and Gallup, NM. We are aware of a specimen collected from Kanab,
UT (A. Norrbom, pers. comm. Aug 2024). We are also able to denitively identify
from photos the distinctive tomentum and shape of this gall on iNaturalist. Thus,
the following localities are added to our own observations to the range of A. lumi-
naria from public user observations: Petried Forest National Park, AZ (obs. no.
2848593 & 112933554); Porcupine Spring, AZ (170773584); Brown’s Canyon, CO
(151317053); Nageezi, NM (141568708); Aztec, NM (151553769); White Sands Na-
tional Park, NM (199614395); and Kodachrome Basin State Park, UT (57095918).
Etymology. The species epithet is a noun derived from the Spanish word for
“light” which is specically used in the southwest United States for small dec-
orative lanterns traditionally displayed during the winter leading up to Christ-
mas. We chose this epithet because the shape of this species’ gall is similar
to that of a small ame on a candle, like those inside luminarias. Furthermore,
this species’ galls are easiest to nd when they are mature, and after the host
leaves have dropped, so they are also associated with display in wintertime in
the Southwest. The tradition of luminarias is common and adored in New Mex-
ico, the type locality of this species. We elected to use the more widespread
term luminaria over northern New Mexico regionally specic “farolito” because
the species’ range extends into other regions in the West.
Discussion
We used multiple lines of evidence to illuminate the species boundaries in an
oft-confused complex of gall-inducing ies in the southwestern United States.
Aciurina luminaria induces galls of a distinct and diagnostic shape on a dif-
ferent E. nauseosa subspecies than its sympatric relatives, A. bigeloviae and
A. trixa. It can further be consistently separated from these species by con-
sistent differences in the adult wing pattern, and by genotyping via reduced
representation genome sequencing.
We provide the following supplement, modied from that of Headrick et al.
(1997) couplets to modify the key to species of Aciurina by Foote et al. (1993),
with gure citations referencing Foote et al. (1993) except as noted. We have
also removed a character from the key of Headrick et al. (1997: 419) that we
found to be inconsistently present in both A. bigeloviae and A. trixa in New
Mexico, and therefore not reliable as a diagnostic character: “pterostigma of at
least one wing with a proximal hyaline or subhyaline incision”.
Additions to the key to species of Aciurina by Foote et al. (1993)
10 Pterostigma along costa no more than 1.5× as long as its greatest width (g.
121c); vein dm-cu nearly straight (g. 121e), the lower apical angle of cell dm
~ 65° (g. 121f); wing predominantly hyaline ................. A. notata (Coquillett)
– Pterostigma along costa at least 2.0 × as long as its greatest width (g.
124a); vein dm-cu usually bowed apicad (g. 124b), the lower apical angle
of cell dm seldom less than 90° (g. 124c); wing with approximately equal
area hyaline and brown, or predominantly brown .....................................11

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Quinlyn Baine et al.: Aciurina luminaria sp. nov.
11 Proximal marginal hyaline incision in cell m lacking median, dark mark ..... 12
– Proximal marginal hyaline incision in cell m with a median, dark, often
elongate mark (g. 122), which sometimes divides the incision (Steyskal
1984: g. 13) (Dodson and George 1986: g. la, b); galls spheroid with
cottony tomentum ..................................................A. bigeloviae (Cockerell)
12 Anal cell bisected at least partially by medial brown mark from veins
A1+CuA1 extending posteriorly, often reaching posterior wing margin; hy-
aline spot within cell br of the wing subcircular, 1–1.5 × as long as wide;
brown region surrounding vein r-m paler in color than remaining dark part
of wing, appearing like a diffuse orange spot; submedial dark mark usually
present crossing cell cua1 from vein CuA1 to posterior wing margin; galls
without tomentum .................................................................. A. trixa Curran
– Anal cell without medial brown mark; hyaline spot within cell br of the
wing elongated longitudinally, 1.5–2.5 × as long as wide; brown region sur-
rounding vein r-m consistent in color, no diffuse spot present; submedial
dark mark in cell cua1 absent; galls frequently ovoid or teardrop shaped
with dense cottony tomentum .......................... A. luminaria Baine, sp. nov.
The adaptive signicance of melanized wing patterns present on tephritid
y species is unclear as studies have found evidence that these patterns could
play a role in sexual communication (Benelli et al. 2014; Hippee et al. 2022),
thermoregulation (Sivinski and Pereira 2005), or predator deterrence by salti-
cid mimicry (Mather and Roitberg 1987; Whitman et al. 1988; Rao and Díaz‐
Fleischer 2012). From this study, it is clear that A. luminaria wings are not sexu-
ally dimorphic, similar to A. bigeloviae and A. trixa. However, it is also clear that
A. luminaria wings have less melanized area on them than these species, and
in combination with their usually paler abdomens, indicates that there may be
an advantage to paler colors in their habitats.
The lower nucleotide diversity of A. luminaria suggests it is a more recently
speciated group, and potentially the result of a “founder-effect” in which very
few individuals from a population establish a lineage after colonizing a novel
niche on a different host plant variety (Balakrishnan and Edwards 2009). This is
supported by our observations of wing pattern variation, which is higher in A. bi-
geloviae and lower in A. luminaria. However, in the eld, populations of the host
plant E. n. subsp. ammophila appear largely fragmented across the range, often
separated by dozens of miles, so the dispersal mechanisms of this species, if it
was evolved from a single founding population, is mysterious.
Because A. luminaria occurs in sympatry with A. bigeloviae but on a unique
host plant subspecies, speciation may be a result of host-race formation. Ev-
idence of speciation via host-race formation, from a host switch specically,
is well-supported in the tephritid genus Rhagoletis, who display extraordinary
host delity similar to that observed in Aciurina (Abrahamson and Blair 2008).
Host-race formation is also documented in the closely related galling tephritid
Eurosta solidaginis (Fitch, 1855), where two populations that are specic to
distinct species of host Solidago (goldenrod) are reproductively isolated via
assortative mating, and oviposition preference to the plant species of maternal
provenance (Abrahamson and Blair 2008). Though we do not have documen-
tation of A. luminaria mating or ovipositional behavior, the lack of intermedi-
ates in both our wing and genomic analyses suggest a similar level of isolation

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Quinlyn Baine et al.: Aciurina luminaria sp. nov.
that may be maintained by similar barriers. In both Eurosta and Rhagoletis,
there is substantial evidence that host switches were advantageous for enemy
escape (Brown et al. 1995; Feder et al. 1995); a new niche, like a host plant
and/or altered gall form may be absent of, or inaccessible to, predators that
previously had a negative impact on tness of the y. In rearing A. luminaria
for this description, we observed many fewer enemy parasitoids than have
been reared from A. bigeloviae in a similar and overlapping range (Baine et al.
2023a), suggesting a lower rate of attack and therefore the possibility of ene-
my escape driving differentiation.
Although evolutionary biologists frequently view species delimitation as an
impossible task due to disagreement on signicant characters that dene spe-
cies concepts (De Queiroz 2007), we can describe species with relatively high
condence if we use “integration by congruence” which delimits based on mul-
tiple, independent, taxonomic characters (e.g., ecological niche + DNA) (Padial
et al. 2010). In the case of A. luminaria, we employ integrative taxonomy to use
the combined evidence of distinction in ecological niche, diagnostic morpholo-
gy, and genomic structure to recognize a new species.
Acknowledgements
The authors would like to thank Gary Dodson and Sarah B. George for essential
background work in Aciurina that enabled this species description. We thank M.
Londoño-Gaviria for preparation of sequence data for repository submission;
L. Leblanc and the William F. Barr Entomological Museum for specimen loan;
E. Gyllenhaal for support with RAD analysis; and D.W.W. Hughes, E.E. Casa-
res, S. Rollins, and H. Sikora for collection of what were previously known as
“marshmallow” galls and rearing support. We are grateful to D. Lightfoot and K.
Miller for MSB resources including specimen photography equipment, and the
M. Syed lab at UNM for wing photography equipment. We also thank our two
expert reviewers, S. Korneyev and A. Norrbom, for suggestions that improved
the quality of this manuscript.
Additional information
Conict of interest
The authors have declared that no competing interests exist.
Ethical statement
No ethical statement was reported.
Funding
This material is based upon work supported by the National Science Foundation under
Grant No. 2021744 and the University of New Mexico.
Author contributions
Conceptualization: VGM, EOM. Data curation: BW, QB. Formal analysis: BW, QB. Funding
acquisition: EOM, VGM. Investigation: EOM, QB, BW, VGM. Methodology: QB. Resources:
EOM, VGM. Supervision: EOM. Visualization: QB. Writing – original draft: BW, QB. Writing
– review and editing: EOM, VGM.

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ZooKeys 1214: 217–236 (2024), DOI: 10.3897/zookeys.1214.130171
Quinlyn Baine et al.: Aciurina luminaria sp. nov.
Author ORCIDs
Quinlyn Baine https://orcid.org/0000-0001-5025-3741
Vincent G. Martinson https://orcid.org/0000-0001-5824-3548
Ellen O. Martinson https://orcid.org/0000-0001-9757-6679
Data availability
All of the data that support the ndings of this study are available in the main text or
Supplementary Information. Sequence data used in phylogenetic analysis is stored in
NCBI GenBank within BioProject ID PRJNA1075688.
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Supplementary material 1
Selection of mounted wing pairs from each of the three sampled
morphotypes to show variation in wing pattern
Authors: Quinlyn Baine, Branden White, Vincent G. Martinson, Ellen O. Martinson
Data type: pdf
Copyright notice: This dataset is made available under the Open Database License
(http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License
(ODbL) is a license agreement intended to allow users to freely share, modify, and
use this Dataset while maintaining this same freedom for others, provided that the
original source and author(s) are credited.
Link: https://doi.org/10.3897/zookeys.1214.130171.suppl1