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ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN
1175-5334
(online edition)
Copyright © 2016 Magnolia Press
Zootaxa 4109 (3): 372
–
380
http://www.mapress.com/j/zt/
Article
http://doi.org/10.11646/zootaxa.4109.3.7
http://zoobank.org/urn:lsid:zoobank.org:pub:23EFD7CD-1C77-4789-BB9E-C497D19F6D2E
The Lepidoptera of White Sands National Monument, Otero County, New Mex-
ico, USA 10. A remarkable new white species of Chionodes Hübner (Gelechiidae)
ERIC H. METZLER
1
& JEAN-FRANÇOIS LANDRY
2
1
Adjunct Curator of Lepidoptera Michigan State University; Research Collaborator U.S.N.M.; Research Associate Museum of South-
western Biology the University of New Mexico. P.O. Box 45, Alamogordo, NM 88311-0045 USA. E-mail: metzlere@msu.edu
2
Agriculture and Agri-Food Canada, Ottawa Research and Development Centre, Central Experimental Farm, Ottawa, Ontario
K1A 0C6, Canada. E-mail: jean-francois.landry@agr.gc.ca
Abstract
The U.S. National Park Service initiated a 10-year study, in late 2006, of the Lepidoptera at White Sands National Mon-
ument, Otero County, New Mexico. Chionodes bustosorum sp. n., described here, was discovered in 2010, during the third
year of the study. The male imago and male genitalia are illustrated, and its DNA barcode is compared to that of seven
other species of Chionodes from western North America.
Key words: Endemism, evolution, U.S. National Park Service, U.S. Army, White Sands Missile Range, Tularosa Basin,
biological diversity, white gypsum dunes, DNA barcodes
Introduction
The White Sands National Monument preserves 284.9 km
2
(110 square miles), about 40%, of the world’s largest
snow-white gypsum dune field. The remainder of the 275 square miles dune field is under the jurisdiction of the
U.S. Army’s White Sands Missile Range. The dune field is located in the northern Chihuahuan Desert in southern
New Mexico’s Tularosa Basin (Schneider-Hector 1993).
In 2006 White Sands National Monument invited Metzler to conduct a 10-year study of moths at the
Monument. The primary purposes of the 10-year study were to compile an inventory of moths and describe new
species in habitats within and immediately adjacent to the white gypsum dunes in the Monument. In 1950 Stroud
reported 20 species of Lepidoptera from White Sands National Monument, none of which is unusual for the region.
In the period 9 February 2007 through 30 January 2016, Metzler recorded more than 650 (unpublished data)
species of described Lepidoptera from White Sands and approximately 40 undescribed species of moths. This is the
twelfth description of a new species of moth emanating from the study (see Metzler 2014b, Metzler et al. 2009,
Metzler & Forbes 2011b, 2011c, Metzler & Lightfoot 2014, Wright 2012, 2014, Wright & Gilligan 2015).
Hodges (1999) revised the species of Chionodes Hübner, [1825], found in North American north of Mexico.
As evidence of the poorly known status of the genus, Hodges described 115 (62%) of the 187 known species as
new to science. The distributions of many species appear to be highly disjunct. Additional collecting of small
moths should fill in some gaps as well as disclose undescribed species. When Hodges (1999) specifically
mentioned the need for more collecting in southern New Mexico, he did not know about the high rate of endemism
of moths in White Sands National Monument (Metzler 2014a).
Methods and materials
Moths and other night flying insects were collected in U.S.D.A. type black-light traps, as described in Smith et al.
(1974), in diverse habitats within the white gypsum dunes. A detailed description of the study methods is given by
Metzler et al. (2009).
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All but the most easily identified species of moths (e.g., Hyles lineata) were retained for further study. The
specimens were sorted, and selected specimens were spread and labeled. All nonlepidopterous insects from the
traps were placed in 95% ethanol and deposited in the Museum of Southwestern Biology at the University of New
Mexico, Albuquerque, New Mexico.
Genitalia were examined following procedures generally outlined in Clarke (1941), Hardwick (1950),
Lafontaine (2004), and Pogue (2002). Abdomens were removed from the moths, dipped in 95% ethanol, and
soaked in 10% KOH for 30 minutes at 50°C. Genitalia were dissected in 5% ethanol, dehydrated in 99.9% ethanol,
stained with Chlorazol Black in water and Safranin O in 99.9% ethanol. The genitalia were cleared in oil of clove,
rinsed in ethyl acetate, and slide mounted in Canada Balsam.
Terms for elements of wing pattern, color, morphology, and genital structures follow Hodges (1999). Terms for
regions of the wing follow Mikkola et al. (2009). Forewing length was measured to the nearest 0.1 mm, from the
base to the apex excluding fringe, using a Leica MZ 12 stereo-microscope with a Wild 15x ocular micrometer.
The photograph of the adult was taken with a Nikon D7100 equipped with an AF-S Micro Nikkor 105mm 1.28
GED lens and a small homemade lightbox of 4” diameter x 4” long white PCV pipe, illuminated with a 60 LED
ring light. The photographs of the genitalia and slide-mounted structures were taken with a Nikon D7100 digital
camera mounted on a Leitz Aristophot, a Summar 42 mm objective, and a 24 mm condensor. The images were
processed with Zerene Systems software and Photoshop CS6 software.
The coordinates for latitude and longitude on the labels of the specimens from the study are in degrees and
decimal minutes. The coordinates were obtained with the aid of a Garmin II Plus and confirmed by reference to
Google™ Earth Pro 7.1.4.1529. The specimens cited in this paper are deposited in the United States National
Museum of Natural History (USNM), Smithsonian Institution, Washington, DC.
Systematics
Chionodes bustosorum Metzler, sp. nov.
BOLD:ACS7412
Figs 1–3
Holotype ♂, pinned, double-mounted, with labels as follows: “New Mexico: Otero Co.,White Sands Nat[ional]
Mon[ument]; Interdune vegetation; 106˚11.38'W; 32˚46.60'N 4,000'; 11 June 2010 WSNM8; Eric H. Metzler uv
tr[a]p; Accss # White Sands National Monument 00131.” [blue label] “Barcode of Life Project Leg(s), DNA
extracted”; “USNMENT01142737”; [green label] “Genitalia slide by EHMetzler, ♂ USNM 146317” [red label]
“HOLOTYPE USNM; Chionodes bustosorum Metzler 2016”. Deposited in USNM.
Paratype ♂, pinned, double-mounted, same locality/date as holotype, [blue label] “Barcode of Life Project
Leg(s) removed, DNA extracted”; “USNMENT01142738” [green label] Genitalia slide by EHMetzler, ♂ USNM
146318.” Deposited in USNM.
Description. Adult male (Fig. 1). Head: Front and vertex scales broadly spatulate, erect, cream-white; front
smooth, scales spatulate, directed forward and ventrally, cream-white; palpi upturned, basal segment scales
appressed, mid-segment = ½ length of palpi, slightly shaggy, scales spatulate, apical segment = 1/3 length of palpi,
divergent apically, scales appressed, cream-white; haustellum base densely scaled, cream-white. Antenna, each
segment basally ringed with semi-erect cream-white scales. Thorax: dorsal and ventral surfaces with appressed,
cream-white scales. Legs with appressed, cream-white scales. Forewing: Length 5.4 mm, mean 5.4 mm, n = 2;
Uniformly cream-white, including fringe, apex rounded; underside pale yellow with concolorous fringe. Hindwing
mirror-like reflective-white with concolorous, long fringe, apex slightly produced; underside white, male with
patch of pale yellow sex scales arising from wing base and extending along inner margin. Abdomen: Scales
appressed, pale yellow. T8 (Fig. 2d) with lateral sides parallel, posteriorly broadly convex, anteriorly broadly
concave. Male genitalia (Fig. 2) (2 preparations examined) with uncus broad, spoon shaped, setose laterally;
culcitula absent; gnathos with base sclerotized, lobed laterally, lobes extending 1/3 length of tegumen, each lobe
with a posteriorly directed finger-like projection enclosing a diamond-shaped lateral process; gnathos sharply
curved at 0.2x length, gently curved most of length, apex sharply recurved; tegumen broadly A-shaped, excavated
to 1/3, robust suture separating pedunculi, each pedunculus narrowing to junction with vinculum, base of each
pedunculus twisted 180˚ at junction with vinculum; vinculum = 0.8x length of tegumen, abruptly narrowed
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immediately anterior of juncture with tegumen, distal 2/3 trough-shaped, sides parallel extended to blunt rounded
apex; posterolateral lobe from vinculum an extension of twisted pedunculus, sclerotized, trough-shaped, apex
bluntly rounded, length = width of base of pedunculus of tegumen; saccus not differentiated from vinculum; valvae
asymmetrical, unequal in length and dissimilarly shaped: right valva, maximally extended to 1/2 length of
tegumen, then strongly recurved mesially at 2/3 length, robust where recurved, apex weakly bifurcate, posterior
projection thorn-like, robust anterior projection pointed, beak-like; left valva narrow, curved, bow-like mesially or
laterally, extended to middle of uncus, terminal 1/10 bent approximately 90˚, apex doubly bifurcate, posterior
projection robust pointed, beak-like, anterior projection bifurcate, not robust. Phallus with distal part sculpting
complex, caecum approximately 5x length of distal part, longitudinal sclerotized bar at anterior end.
Adult female. Unknown.
FIGURE 1. Chionodes bustosorum adult male paratype (scale bar = 1 mm).
Diagnosis. Ron Hodges (personal communication to EHM) considered C. bustosorum to be undescribed based
on the diagnosis in combination with the details of the description. Chionodes bustosorum (Fig. 1) is a small
(forewing length 5.4 mm) creamy white moth with no discernable markings on the surface. Two nearly identical
specimens were captured in black light traps with Scythrididae of similar size and appearance. The apex of the
forewing of C. bustosorum is rounded, whereas the forewing of Scythrididae terminate in an acute apex. The apex
of the hindwing of C. bustosorum is rounded and produced, a wing shape that is typical for most Gelechiidae. The
apex of the hindwing of Scythrididae is acute. The male genitalia of C. bustosorum are typical for the genus
Chionodes (Fig. 2) with a broad prominent spoon-shaped uncus. Chionodes bustosorum keys out to couplet 6,
obscurusella or formosella groups, in the Hodges (1999: 25) key. The male sex scales of C. bustosorum are on the
ventral surface of the hindwing, similar to species in the obscurusella group. Based on characters of the male
genitalia as figured by Hodges, C. bustosorum appears to be in the abella complex of the formosella group,
however, C. bustosorum fails to match any features in the early couplets in the key to species of the formosella
group (see Hodges 1999: 33). The diagnostic features of the male genitalia of C. bustosorum are the markedly
asymmetrical valvae. The left valva is long and narrow, whereas the right valva is shorter, more robust, and
recurved mesially like the cutting blade of a hand-held scythe. In a comparison of male genitalia, based on the
photographs in Hodges (1999), the genital capsule of C. bustosorum is most similar to a lateral mirror image of C.
abella as illustrated on p. 261, fig. D-1 of that work. The angle of the recurved right valva of C. bustosorum is
broad, like a hand-held scythe, whereas the angle of the recurved left valva of C. abella is acute. The area between
the pedunculi of the tegumen of C. bustosorum is not sclerotized, whereas in C. abella it is lightly sclerotized. The
lateral sides of the uncus of C. abella appear to be nearly parallel; the uncus of C. bustororum is oval. The genital
capsule also resembles a mirror image of that of C. abdominella as illustrated on p. 263, fig. F-20 of Hodges
(1999).
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FIGURE 2. Chionodes bustosorum male genitalia and male eighth abdominal segment. 2a, male genitalia paratype USNM
slide # 146318; 2b, male aedeagus paratype USNM slide # 146318; 2c, male eighth abdominal segment (tergum on left)
holotype USNM slide # 146317 (scale bars = 1 mm).
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FIGURE 3. Chionodes bustosorum type locality and distribution map. 3a, type locality; 3b, Chionodes bustosorum is known
from White Sands National Monument, Otero Co., New Mexico.
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Remarks. This new species is placed in the genus Chionodes based on the presence of the caecum on the
aedeagus as defined by Hodges (1999). The holotype and paratype are identical in habitus and genital structure.
The paratype imago is illustrated because it possesses complete antennae. The holotype was selected because it is
the specimen that yielded a nearly complete barcode sequence.
Etymology. The specific name of this species, bustosorum, a noun in the genitive case, recognizes David
Bustos, Chief of Resources at White Sands National Monument since 2007, and his wife, Andrea. David
aggressively pursues research; his efforts contributed greatly to the immense accumulation of scientific data during
his years at White Sands National Monument. Metzler knows from personal experience that David works long and
crazy hours in support of the Monument. He is enthusiastic about the research on moths, to wit, he often sends
emails to Metzler from home, long after Metzler is in bed. David is the recipient of the U.S. National Park Service’s
2014 Director's Trish Patterson Student Conservation Association Award for Natural Resource Management in a
Small Park. His efforts would not be possible without the support and encouragement of his lovely wife Andrea.
Metzler takes great pleasure in naming this moth in honor of David and Andrea Bustos.
Distribution and biology. Chionodes bustosorum occurs in White Sands National Monument, Otero County,
New Mexico (Fig. 3). The immature stages and host plant are unknown.
DNA barcode analysis
Tissue samples (dry legs) were shipped to the Canadian Centre for DNA Barcoding in Guelph for DNA extraction,
amplification, and sequence analysis. The protocol followed is as outlined in Landry & Hebert (2013). Barcoding
efforts included the holotype and paratype of C. bustosorum. Records of other Chionodes species already in BOLD
were selected for barcode comparison and analysis. The Barcode Identification Numbers (BINs) (Ratnasingham &
Hebert 2013) in BOLD are used as registry designations for barcode clusters. Neighbor-joining trees and genetic
distances were calculated with the Taxon ID Tree and Distance Analysis tools available in BOLD using the Kimura
two-parameter (K2P) model of base substitution with pairwise deletion for ambiguous positions, and Kalign
sequence alignment. Details of the barcoded specimens (collecting data, photos, sequence data, GenBank
accessions) are available through the online dataset http://dx.doi.org/10.5883/DS-CHIOWHIT.
TABLE 1. Percentage of divergence in DNA barcode (cytochrome c oxidase I gene) sequences among eight species of
Chionodes. “BOLD:ABC1234” = Barcode Index Number. Diagonal cells = intra-specific distances; cells below diagonal
= inter-specific distances; means with standard errors in parentheses.
abdominella
abella
bustosorum
fructuaria
landryi
pinguicula
sistrella
sp near
fructuaria
abdominella
BOLD:ACG5519 (n=4)
0.2
(±0.10)
abella BOLD:AAB6065
(n=6)
8.0
(±0.08)
0.0
(±0.00)
bustosorum BOLD:ACS7412
(n=1)
5.9
(±0.14)
7.4
(±0.00)
-
fructuaria BOLD:AAN5596
(n=4)
6.3
(±0.19)
6.2
(±0.14)
5.0
(±0.22)
0.4
(±0.21)
landryi [no BIN] (n=1) 4.0
(±0.13)
5.9
(±0.00)
4.8
(±0.00)
4.1
(±0.25)
-
pinguicula BOLD:AAH5117
(n=2)
6.6
(±0.12)
8.4
(±0.00)
6.9
(±0.25)
6.8
(±0.49)
5.4
(±0.00)
2.0
sistrella BOLD:ACI6713
(n=7)
8.3
(±0.36)
8.4
(±0.28)
5.5
(±0.53)
6.1
(±0.39)
6.8
(±0.63)
6.8
(±0.31)
0.4
(±0.37)
sp near fructuaria
BOLD:AAN5597 (n=2)
7.2
(±0.13)
7.6
(±0.09)
5.5
(±0.12)
5.8
(±0.20)
4.2
(±0.19)
4.2
(±0.21)
7.0
(±0.33)
0.8
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The holotype of C. bustosorum yielded a slightly abbreviated sequence of 636 bp which was of sufficient
quality to be assigned to a BIN (BOLD:ACS7412). Barcoding of the paratype failed. Twenty-six specimens
representing seven other Chionodes species were selected from BOLD for comparative barcode analysis. To make
that selection, an initial neighbor-joining (NJ) tree was generated by searching BOLD for all available Chionodes
records from North America: this resulted in a large tree comprising over 1500 sequences representing more than
100 species (not shown). A smaller subset of species was then selected based on their genetic proximity to the type
of C. bustosorum in the larger tree. These included C. abdominella (Busck), C. fructuaria (Braun), C. landryi
Hodges, C. pinguicula (Meyrick), C. sistrella (Busck), and one unidentified BIN provisionally named C. sp. near
fructuaria. Additionally, Chionodes abella (Busck), although appearing genetically quite distant, was also selected
because it is the species whose male genitalia are compared to those of C. bustosorum in the diagnosis. A tree was
then generated and distances calculated for the smaller subset (Fig. 4; Table 1).
Chionodes bustosorum is the sole representative of its BIN, congruent with the distinctiveness in morphology
and confirming its status as a uniquely distinct species. It is nested within a cluster with C. sistrella as nearest
neighbor, and with C. pinguicula and C. sp. near fructuaria forming a neighboring subcluster. It is interesting to
note that the similarity in male genitalia with C. abella is not reflected in barcode proximity, as the latter is the most
distant of all clusters included in the subset. Chionodes abella is also the most different in coloration, being a dark-
colored species. It is noteworthy that the genetically closest species all have a substantial amount of pale creamy
white on the head, thorax, legs, and forewings, suggesting the possibility that C. bustosorum may be derived from
an ancestor with a predominantly pale coloration.
FIGURE 4. Neighbor-joining tree of K2P distances for the barcode region of the cytochrome c oxidase I gene among 27
specimens representing eight species of Chionodes. End-branch labels are Specimen IDs. Barcode Index Numbers (BIN) where
available are indicated below each species name.
Discussion
Species of the family Gelechiidae are diverse, and a satisfactory way to identify genera from external characters is
lacking. Fortunately, Hodges (1999) provided a definition of Chionodes in his revision of species in America North
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WHITE SANDS CHIONODES (GELECHIIDAE)
of Mexico. His work greatly aids identification of species of this large genus. The surprisingly rich endemic fauna
of moths from White Sands National Monument includes several very pale or nearly white species, including
Chionodes bustosorum described here. At first the plethora of white and pale species in the snow white dunes
seems an obvious outcome of crypsis, yet the explanations are probably more complex. Because most
lepidopterans are herbivores, an analysis of the plants growing in the gypsum soils is needed. David Bustos’
preliminary findings show that some of the species of plants in the dunes have genetic markers and associated
microbes that differ from the same species of plants growing outside, and immediately adjacent to the gypsum
soils. These data suggest that larvae eating plants inside the dunes are consuming diets unlike larvae eating the
same species of plants outside the dunes, but the relationship between diets of plants in dunes that are genetically
different from the same species outside the dunes is unknown relative to the evolution of white colors of
individuals. More research must be done before drawing any conclusions, yet the natural, nearly unspoiled by
humans, laboratory of the Monument offers an unusual opportunity to explore the evolution of so many species of
moths. The lack of lepidopteran specimens from the Monument seen until now can probably be attributed to the
dearth of insect collecting in the gypsum dunes ecosystem in New Mexico because the dunes were formerly private
property, and are now under the control of the U.S. National Park Service and the U.S. Army.
Acknowledgements
Three non-profit organizations contributed funding for travel and logistics in support of this study of Lepidoptera at
White Sands National Monument. Metzler is especially grateful for the substantial financial support of The
Western National Parks Association, Tucson, Arizona. The El Paso Zoo Conservation Committee, El Paso, Texas,
and the Association of Zoos and Aquariums’ Terrestrial Invertebrate Taxon Advisory Group (TITAG), Seattle,
Washington also contributed small grants. Their commitment to this research is rewarding. Several executives from
White Sands National Monument, including David Bustos, Marie Frias-Sauter, Becky Burghart, Joseph Roberts,
and Cliff Spencer, were instrumental in arranging and promoting this study of the moths. We single out David
Bustos for his enthusiastic support and for getting things done.
The National Park Service provided permits to take samples of moths and provided access to areas normally
closed to the public. Representatives from research collections and other institutions provided insect pins, alcohol,
identification services, research consultation, and storage space for specimens collected leading to discovery and
naming of this species. Metzler thanks the following persons for offering support from their respective institutions:
Kelly B. Miller, Sandra L. Brantley, and David C. Lightfoot (University of New Mexico), Frederick W. Stehr,
Anthony I. Cognato, and Gary L. Parsons (Michigan State University), J. Donald Lafontaine, Vazrick Nazari, and
B. Christian Schmidt (Canadian National Collection of Insects, Arachnids, and Nematodes), Larry Berger (Ohio
Department of Agriculture), and David Adamski, John W. Brown, Mark E. Metz, David G. Furth, Patricia Gentili-
Poole, and M. Alma Solis (United States National Museum of Natural History). Patricia A. Metzler faithfully
assisted the first author on many aspects of this study, and she provided funding. We thank Marie Frias-Sauter,
Peter Huemer, and one anonymous reviewer for reading the paper and offering valuable suggestions.
We are grateful to the team at the Biodiversity Institute of Ontario, University of Guelph, Ontario, Canada for
their great assistance in the production of DNA barcodes. Funding for DNA barcoding and sequence analysis was
partly provided by the Government of Canada through Genome Canada and the Ontario Genomics Institute in
support of the International Barcode of Life project, and by NSERC. Our work was also aided by the BOLD
informatics platform whose development is funded by the Ontario Ministry of Economic Development and
Innovation.
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