Research Article |
Corresponding author: Ryan A. St Laurent ( rstlaurent@flmnh.ufl.edu ) Academic editor: Donald Lafontaine
© 2020 Ryan A. St Laurent, Lawrence E. Reeves, Akito Y. Kawahara.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
St Laurent RA, Reeves LE, Kawahara AY (2020) Cicinnus chambersi: a new species of sack-bearer moth (Lepidoptera, Mimallonidae, Cicinninae) from southeastern Arizona, USA. ZooKeys 931: 49-71. https://doi.org/10.3897/zookeys.931.50203
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A new species of cicinnine Mimallonidae, Cicinnus chambersi sp. nov., is described from the Sky Islands Region of southern Arizona, USA. The new species is closely related to C. mexicana (Druce), type locality Veracruz, Mexico, based on morphology and genetics. The other Cicinnus species known from the United States, the common C. melsheimeri (type locality Pennsylvania, USA) is morphologically and genetically distinct from both C. chambersi and C. mexicana. The new species is compared to C. mexicana and C. melsheimeri, as well as other Mexican Cicinnus. The life history of C. chambersi is unknown, but its description should facilitate future studies on this rarely reported North American mimallonid, a species which may have only recently become established in the United States. Cicinnus chambersi is the fifth known Mimallonidae species from the United States, and the first described from the country in nearly half a century.
Cicinnus chambersi sp. nov., Mimallonoidea, Sky Islands, taxonomy
Mimallonidae are a family of approximately 300 species of moths endemic to the New World, with the vast majority of species found in Central and South America (
Two mimallonid species inhabit the eastern United States and extreme southern Ontario east of the Great Plains: L. chiridota and C. melsheimeri. Although the majority of C. melsheimeri records are from the eastern United States, this species is quite widespread in the Rocky Mountains, although it is rarely collected in the region. A single, predominantly Mexican species reaches its northernmost extent in southern Arizona: L. arizonicum (
In southern Arizona, L. arizonicum is a somewhat regularly collected, late night-flying denizen of mid-elevation oak forests, and its life history was recently published (
The Sonoran Desert’s Sky Islands Region is located at a biogeographic crossroads at the convergence of several biotic zones. Patterns of biodiversity are influenced by the ecological communities of the Sonoran and Chihuahuan Deserts, the Rocky Mountains, the Great Plains, the Sierra Madre and the Neotropics (
The combination of location, at the interface of biotic zones, the complex topography of the Sky Islands, and relative isolation of mountain range islands promotes unique ecological communities with high species diversity and endemism (
All dissections performed for this study followed
Specimens examined are deposited in the collections listed below. Figures in this paper were created with Adobe Photoshop as part of the Creative Cloud (Adobe 2019), and maps were built using SimpleMappr (
BME Bohart Museum of Entomology, University of California, Davis, California, USA;
BWC B. Walsh Private Collection, Tucson, Arizona, USA;
CJM Collection of José Monzón, Guatemala;
CRAS Research collection of R. St Laurent, Gainesville, Florida, USA;
MCZ Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts, USA;
MGCL McGuire Center for Lepidoptera & Biodiversity, Gainesville, Florida, USA;
PJD Collection of Paul J. Dennehy, Pennsylvania, USA;
VOB Becker Collection, Camacã, Bahia, Brazil;
We refer to the anchored hybrid enrichment (AHE) Mimallonidae phylogeny of
Maximum Likelihood (ML) phylogenetic analyses of unpartitioned COI data was performed using IQ-TREE v. 1.6.10, with branch supports reported as 1,000 Ultrafast Bootstraps (UFBoot) and SH-aLRT as a secondary measure of support (
All sequence data provided by this study will be made available on GenBank, with applicable accession numbers provided in Suppl. material
Maximum likelihood phylogenetic tree inferred with IQ-TREE based on the COI marker, rooted to Gonogramma hanseni. Black circles indicate SH-aLRT/UFBoot of 80/95 or greater for both values, and gray circles indicate SH-aLRT/UFBoot of 80–95 for both values. The scale bar represents expected number of nucleotide substitutions per site. See Suppl. material
Holotype. United States of America – Arizona • Arizona: Santa Cruz. Co., Peña Blanca Lake, Pajarito Mtns., Coronado NF; 750 W MV, 1000W MH, 31.402057, -111.084236, 21.VII.2015; leg. L.E. Reeves/ St Laurent dissection 2-20-17:1 Cicinnus sp./ St Laurent barcode 2-20-17:1 [barcode unsuccessful]/ St Laurent BC 5-6-19:1 [second barcode attempt]/ Holotype ♂ Cicinnus chambersi St Laurent, Reeves, Kawahara, 2020 [red label]/ (MGCL).
Paratypes. (3 ♂, 2 ♀ total) United States of America – Arizona – Cochise County • 1 ♀; Copper Canyon, Huachuca Mts; 31.363, -110.300; 6,000 ft [1,828 m]; 4.VII.2018; C.W. Melton [leg.]; photo ID no. 18070692, St Laurent dissection: 5-9-19:1; (MGCL). – Santa Cruz County • 1 ♂, 1 ♀; California Gulch, Pajarito Mountains; 31.422N, 111.245W; 3800 ft [1,158 m]; 27.VII.2017; J.B. Walsh leg.; MV/UV; (BWC) • 1 ♂; Peña Blanca Lake/ Ruby Rd area; 31°23'16"-24'N, 111°05'25"-07'W; 2–4.VIII.2017; James Adams & Lance Durden; light traps, LEP-58833 [MGCL AHE voucher number and St Laurent dissection number], St Laurent BC 5-6-19:2 [barcode]; (MGCL) • 1 ♂; Peña Blanca Canyon; 31.3844N, 111.0935W; 3.VIII.2017; P. Dennehy leg.; (PJD). Paratypes with the following yellow label: Paratype ♂/ ♀ Cicinnus chambersi St Laurent & Reeves, 2020.
Adult ♂ Cicinnus a dorsal b ventral 6 C. chambersi holotype, USA, Arizona, Santa Cruz Co., Peña Blanca Lake, Pajarito Mtns., Coronado National Forest (MGCL) 7 C. chambersi paratype, USA, Arizona, Santa Cruz Co., Peña Blanca Lake/ Ruby Rd area (MGCL) 8 C. melsheimeri, USA, Florida, Alachua Co., Micanopy (MGCL) 9 C. chabaudi, Mexico, Oaxaca, ca. 15 km SE San Martín Huamalulpan, Cabañas Yucunuvichi (MGCL). Scale bar: 1 cm.
[not included in type series]. United States of America – Arizona – Santa Cruz County • 2 ♂, 1 ♀; California Gulch; 31°25'18.33"N, 111°14'40.02"W; 3,790 ft [1,155 m]; 23.VII.2015 [2 ♂], 21.VII.2017 [1 ♀]; E. Rand leg. (Coll. E. Rand, Arizona) • 1 ♂; Peña Blanca Canyon; 31°23'18.38"N, 111°5'33.00"W; 3895 ft [1,187 m]; 17.VII.2009; E. Rand leg. (Coll. E. Rand, Arizona) • 1 ♂; Jct. FR 49 & FR 812; 31°27'54.88"N, 110°43'9.94"W; 4960’ [1,512 m]; 8.VII.2010; E. Rand leg. (Coll. E. Rand, Arizona).
[not collected and not included in type series]. United States of America – Arizona – Pima County • 1 ♀; Box Canyon; 31.799198, -110.798744; photographed by Salvador Vitanza, Entomologist (Identifier) at APHIS-PPQ, Arizona (Fig.
In southern Arizona, there are no other moths with which this species could be confused, the only other congener found in the United States, C. melsheimeri, has not been found to be sympatric with C. chambersi, but occurs farther north in mountainous northern Arizona, north of the Mogollon Rim, and northeast into central and northern New Mexico. Because C. chambersi and C. melsheimeri are both found in Arizona, we compare them here, although they are not each other’s closest relatives within Cicinnus (see remarks later).
Cicinnus melsheimeri is a somewhat variable species, usually with brown shaded regions along the wing margins, in comparison, C. chambersi is more consistently uniformly pink in coloration with a homogenous ground color. The apex of the forewing of C. chambersi is much sharper than in C. melsheimeri, and the postmedial line of the forewing more distinctly forms a right or acute angle near the apex of the forewing, whereas this same angle is more obtuse in C. melsheimeri (see Figs
Genitalia of the two species of Cicinnus in Arizona are structurally very distinct, although they both have the characteristics deemed apomorphic of Cicinnus sensu stricto as defined by
The differences between C. chambersi and the other Mexican Cicinnus species are less obvious. The only names currently applied to similar Mexican species are C. chabaudi Dyar, 1914 (Figs
Adult Cicinnus a dorsal b ventral 11 C. chambersi ♀ paratype, USA, Arizona, Cochise Co., Copper Canyon, Huachuca Mts, 1828 m (MGCL) 12 C. melsheimeri ♀, USA, Texas, Cameron Co., Brownsville (MGCL) 13 C. near mexicana ♂, Guatemala, Zacapa, Sierra de las Minas, N Rio Hondo, E San Lorenzo, Cerro Monos env., 2243 m (MGCL) 14 C. mexicana ♂, Guatemala, Baja Verapaz, SE Purulhá, Ranchitos de Quetzal, Parque Ecológico Gucumatz, 1660 m (MGCL). Scale bar: 1 cm.
Cicinnus chambersi ♂ genitalia a ventral, vinculum extension in natural position, deciduous setae intact b ventral, vinculum extension in natural position, deciduous setae removed c ventral, vinculum extension held open d lateral 15 holotype, USA, Arizona, Santa Cruz Co., Peña Blanca Lake, Pajarito Mtns., Coronado National Forest, St Laurent dissection: 2-20-17:1 (MGCL) 16 paratype, USA, Arizona, Santa Cruz Co., Peña Blanca Lake/ Ruby Rd area, St Laurent dissection: LEP58833 (MGCL). Scale bar: 1 mm.
Cicinnus ♂ genitalia a ventral, vinculum extension held open b lateral 17 C. melsheimeri, USA, Ohio, Geauga Co., Thompson Township, St Laurent dissection: 8-10-18:1 [arrow denotes vincular arms which are unique to C. melsheimeri among Cicinnus] (MGCL) 18 C. chabaudi, Mexico, Oaxaca, ca. 15 km SE San Martín Huamalulpan, Cabañas Yucunuvichi, St Laurent dissection: 8-10-18:3 (MGCL) 19 C. near mexicana Guatemala, Zacapa, Sierra de las Minas, N Rio Hondo, E San Lorenzo, Cerro Monos env., 2243 m, St Laurent dissection: 7-25-18:1 (MGCL) 20 C. mexicana, Guatemala, Baja Verapaz, SE Purulhá, Ranchitos de Quetzal, Parque Ecológico Gucumatz, 1660 m, St Laurent dissection: 7-25-18:4 (MGCL). Scale bar: 1 mm.
Male. Head: Coloration pinkish beige with an ample speckling of dark brown petiolate scales. Antennae pale yellow with a covering of beige scales, occasionally speckled with darker brown scales, bipectinate to tip, distal quarter of pectinations dramatically shorter than basal three quarters of pectinations. Eyes very large, comprising more than two thirds area of head. Labial palpus exceedingly short, not extending beyond frons, coloration as for head though with darker gray scales dorsally; labial palpus apparently three-segmented though distalmost segment miniscule. Thorax: Dorsally light beige with profuse speckling of dark brown petiolate scales, prothorax lighter in color, pinker, ventrally thorax as above. Legs: Coloration mostly as for thorax. Tibial spurs small, about as long as one quarter length of first tarsomere. Forewing dorsum: Forewing length: 20–22 mm, avg.: 21 mm; wingspan: 39–45 mm, N = 3. Triangular, apex sharply falcate, outer margin mostly convex except for concavity below apex and slight tornal concavity. Ground color same light beige as thorax, with underlying pink hue throughout, profusely speckled with dark brown petiolate scales which are less densely distributed submarginally. Antemedial line very faint, usually nonexistent but if present diffuse and wavy. Postmedial line fine, almost always nearly straight, well-defined, dark brown, perpendicularly angled toward costa after passing Rs4, line thickness variable but comparatively thicker near costa. Entire wing nearly concolorous except for postmedial line and discal spot though coloration grayer rather than pink along costa, especially apically. Discal spot variably developed ranging from faint comma-like mark to well-developed gray-brown oval situated at distal margin of discal cell. Fringe darker brown than ground color of wing. Forewing ventrum: Ground color similar to dorsum but suffused with bright orange-red especially medially and along veins, basally wing much pinker than dorsally. Bright orange-red patch of scales present submarginally between Rs3–M3. Postmedial line weakly defined, consisting of dentate, convex line that is neither straight nor distinctly angled toward costa. Discal spot may be more well-defined than on dorsum. Hindwing dorsum: Rounded, coloration and patterning as for forewing dorsum, but antemedial line absent, postmedial line outwardly convex, discal spot less defined. Hindwing ventrum: Follows similar pattern as forewing ventrum, postmedial line convex and more interrupted by veins than on dorsum, discal spot weakly developed. Abdomen: Robust, extending beyond anal angle of hindwing, coloration mostly as for thorax. Sternite of VIII anteriorly and posteriorly concave, with pair of short protuberances, one on either side of posterior concavity. Genitalia: (Figs
Cicinnus ♀ genitalia a ventral b lateral c dorsal 21 C. chambersi paratype, USA, Arizona, Cochise Co., Copper Canyon, Huachuca Mts, 1828 m, St Laurent dissection: 5-9-19:1 (MGCL) 22 C. near mexicana, Mexico, Chiapas, San Cristobal de las Casas env., nr. Hotel Flores, 2415 m, St Laurent dissection: 5-6-19:4 [note: ductus and corpus bursae not shown, but highly elongate as in Fig.
The life history of C. chambersi is unknown, but we expect the larvae feed on oaks (Quercus spp. Linnaeus) as do all Mimallonidae in Canada and the United States for which larval hosts are known. Cicinnus chambersi appears to be a denizen of mid-elevation oak-dominated habitats of the Sky Island Region. The type series and other examined specimens were collected at elevations ranging from 1,155 m to 1,828 m, all within the oak belt of sky island mountain ranges (
Cicinnus chambersi is known only from sky island mountain ranges of southeastern Arizona (Figs
This new species is named for Aaron Chambers of Tucson, Arizona, a desert dweller and dear friend of the authors, in recognition of his support of native biodiversity and for imparting his expansive ecological knowledge of the Sonoran Desert to us every monsoon season.
It is surprising that C. chambersi has been overlooked in North America due to the distinct morphological differences between C. chambersi from Arizona and the common C. melsheimeri with which the new species has been confused. However, C. chambersi appears to be a rarely collected moth considering the few specimens known to us and the intensity at which insect collecting occurs in southern Arizona, and this could be the reason C. chambersi has not yet been described. To our knowledge, no specimens of C. chambersi were available to Franclemont at the time of his 1973 revision (St Laurent pers. obs. of the Cornell University Insect Collection). In fact, we are not aware of any specimens collected in the United States prior to 2009. Therefore, it is also possible that C. chambersi is a relatively recent establishment from Mexico in southern Arizona.
Apart from the differences between C. chambersi and C. melsheimeri revealed by genitalia dissections, ongoing phylogenomic work using anchored hybrid enrichment (
In describing C. chambersi, it is necessary to go into some additional depth in discussing the Cicinnus of Mexico in order to couch the new species within the broader context of its conspecifics in the region, where several described and undescribed taxa are found. The common North American C. melsheimeri, discussed above in the diagnosis of C. chambersi and in the phylogenetic justification for the validity of the new species, also appears to be found in Mexico. Cicinnus melsheimeri is found throughout the eastern United States and southeastern Canada, with sparse records in the Rocky Mountains and the western United States (Colorado, Utah, northern Arizona, central and northern New Mexico, and the Big Bend Region of western Texas, and southern Texas) (St Laurent unpublished). Additional populations of a taxon near C. melsheimeri are known from throughout the mountainous regions of Mexico (in Chihuahua, Distrito Federal, Hidalgo and Nuevo Leon) as well, these specimens display the typical genitalia of C. melsheimeri complete with the vincular arms (NHMUK dissection NHMUK010402293 of a specimen from Hidalgo, examined). The degree of cryptic diversity included under the name C. melsheimeri is yet to be fully resolved, though this will be a worthy area of research. It is clear however, that true C. melsheimeri (type locality, Pennsylvania, USA), which we here consider to include all populations morphologically most similar to the eastern USA species, as well as those in Mexico, can be readily differentiated from the remainder of Cicinnus (including C. chambersi) by the vincular arms that are present in the male genitalia. Cicinnus melsheimeri has been shown to represent a distinct lineage of Cicinnus sister to the remainder of the genus within a phylogenomic (
Other Cicinnus populations in Mexico, including C. chambersi which ranges as far north as southern Arizona, belong to the more typical, primarily tropical American group of Cicinnus that always lack vincular arms in the male genitalia. This putative species-group contains all Cicinnus sensu stricto (sensu
In northwestern Mexico there exists no name to adequately refer to Cicinnus species there, except for specimens clearly more allied to C. melsheimeri as discussed previously. Therefore, C. chambersi is the first named species belonging to the typical Neotropical Cicinnus species-group described from the arid southwestern United States and (likely) northwestern Mexico. We are aware of five specimens of Cicinnus from northwestern Mexico that are morphologically similar to C. chambersi but are from scattered localities with inadequate numbers of specimens from each location to allow for a convincing determination as C. chambersi. These specimens were also not barcoded. Therefore, none of these specimens are included in the type series of C. chambersi in order to conservatively restrict the type series to specimens collected at and around the type locality in southern Arizona. Each of the five northwestern Mexican specimens will be discussed below in order to bring attention to them in hopes that additional material will be discovered or collected to better determine their identities. Complete collecting data, including institutional depositions, for these specimens can be found in the supplemental appendix (Suppl. material
One male specimen from the
In the BME there are two apparently conspecific male specimens from Chihuahua, one from Temoris and another from Cuiteco, and a putatively conspecific female from Choix, Sinaloa. While these Chihuahuan males are externally very similar to C. chambersi from Arizona, and inhabit comparable habitats, the female from nearby Choix displays a postmedial line angled differently from any examined female C. chambersi, and thus makes the determination of the Chihuahuan males as C. chambersi inconclusive (if the two males and the female are regarded as conspecific).
A single large male from the
Finally, a single male from Minatitlán, Colima (in VOB) and a single female from nearby Michoacán (Barcode of Life Datasystems, BC-Her2624) are known, which are morphologically more similar to C. chambersi than C. mexicana and may represent another undescribed species near C. chambersi or the southern extent of the distribution of this species. We were unable to examine the genitalia of this population, though a barcode of the Michoacán specimen places it sister to C. chambersi (see Fig.
Aaron Chambers (Arizona), namesake of C. chambersi, assisted the authors with collecting on many occasions in Arizona. Trace Hardin, Isaac Powell, and Doug Main assisted with specimen collection at Pena Blanca Lake, Arizona. Salvador Vitanza provided his excellent photos of a female C. chambersi for publication in this article. James Adams (Dalton State College, Georgia), Charles Melton (Arizona), Paul Dennehy (PJD), and Bruce Walsh (BWC), each supplied us with specimens or photos of specimens designated as paratypes of C. chambersi. Stefan Naumann (Germany) donated many Mexican and Guatemalan Cicinnus specimens for study, this material was essential for understanding the distribution of this genus in Central America. We would also like to offer our sincere thanks to the staff and managers of the various collections that provided images and materials to make this study possible: Courtney Richenbacher, David Grimaldi (
Figure S1
Data type: PDF figure
Explanation note: Maximum likelihood phylogenetic tree inferred with IQ-TREE based on the COI marker, rooted to Gonogramma hanseni.
Table S1
Data type: Species data
Explanation note: Data for Cicinnini specimens used in this study, either sequenced de novo or extracted from publicly available data on BOLD (http://www.boldsystems.org/).
File S1
Data type: Species data
Explanation note: Data and remarks for additional Cicinnus examined in this study.
File S2
Data type: Genetic data
Explanation note: FASTA file of COI alignment for all taxa used in Figure 1/ Supplementary material
File S3
Data type: Phylogenetic data
Explanation note: Tree file in NEWICK format corresponding to phylogeny in Figure 1/ Supplementary material