Research Article |
Corresponding author: B. Christian Schmidt ( christian.schmidt@canada.ca ) Academic editor: Donald Lafontaine
© 2018 B. Christian Schmidt, J. Bolling Sullivan.
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:
Schmidt CB, Sullivan BJ (2018) Three species in one: a revision of Clemensia albata Packard (Erebidae, Arctiinae, Lithosiini). In: Schmidt BC, Lafontaine JD (Eds) Contributions to the systematics of New World macro-moths VII. ZooKeys 788: 39-55. https://doi.org/10.3897/zookeys.788.26048
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Clemensia albata Packard, previously thought to be a single, widely distributed North American species, is here shown to consist of three distinct, primarily parapatric species: Clemensia albata Packard, C. umbrata Packard, stat. rev., and Clemensia ochreata Schmidt & Sullivan, sp. n. Adults and genitalic structures of the three species are illustrated.
algivory, cryptic species, lichen moth, lichenivory
The genus Clemensia Packard encompasses about 55 described species (
The variability of Clemensia albata was recognized as early as
Repository abbreviations. Voucher specimens (Suppl. material
JBS J. Bolling Sullivan Collection, Beaufort, North Carolina, USA
JTT Jim T. Troubridge Collection, Hagersville, ON
MEM Mississippi Entomological Museum, Starkville, MS
Due to the difficulty in species identification based solely on photographs, records from citizen-science groups such as BugGuide.net and iNaturalist.org were not included here, with the exception of a few records that could be identified with reasonable confidence, and that represent significant range-gap or range-edge records (Suppl. material
The three Clemensia phenotypes that occur in North America are shown in Figs
Examples of all three phenotypes resolved into three distinct DNA barcode sequence clusters, or BINs (Barcode Index Numbers; see
Clemensia
umbrata
Packard, 1872: 85. Type locality. Congress Springs, Santa Clara Co, California [lost] male holotype. Note. The type locality was given as “California” in the original description, and
Clemensia irrorata H. Edwards, 1874, p.185. Type locality. “Victoria, V.I. [Vancouver Island, British Columbia]”
Clemensia umbrata is most similar to C. albata; flight time and locality aid in separating the two. Both species occur together only from southern Quebéc and eastern Ontario southward; C. umbrata is the only Clemensia species across the boreal forest region and the Pacific Northwest (Figure
Externally, C. umbrata differs from C. albata in its larger size in regions of sympatry (northern boreal C. umbrata are smaller and not noticeably significantly larger than C. albata), with male forewing length of 12.3 mm (n = 9) versus 10.8 mm (n = 6) in C. albata. Wing pattern differences are difficult to discern, especially flight-worn individuals, but C. umbrata has a more contrasting forewing pattern that is more suffused with grey and black, and often with a diffuse dark grey postmedial patch near the anal margin; this patch is absent or much more restricted in C. albata.
Internally, the male genitalic structure of C. umbrata and C. albata differs in the shape of the basal ventral diverticulum of the vesica, which is bilobed in C. umbrata versus heart shaped in C. albata (Figs
Clemensia umbrata occurs from Nova Scotia across the boreal region to the Pacific coast, southward into central California and northern Idaho (Figure
As defined here, C. umbrata represents the taxon that occurs across most of Canada and western USA that was previously called C. albata. In most of its range (except eastern North America), it is the only Clemensia species. Although no California specimens of C. umbrata were available for DNA analysis, examined California material was not distinguishable morphologically from that of the Pacific Northwest, with the latter genetically very similar to the transboreal/Appalachian taxon. The distribution of C. umbrata is continuous along the Pacific coast from southern British Columbia to central California (Figure
Clemensia adults. 1–4 C. albata, 1 ♂, Manitoulin Island, Ontario, Canada 2 ♂, Grandfather Mountain, 4850’, Avery Co., North Carolina, USA 3 ♂, Langley, British Columbia, Canada 4 ♀, Tweed, Ontario, Canada 5–8 C. umbrata, 5 ♂, Backus Woods, Ontario, Canada 6 ♀, Backus Woods, Ontario, Canada 7 ♂, Starkville, Oktibbeha Co., Mississippi, USA 8 ♀, Barksdale A.F.B., Bossier Parish, LA, USA; 9–12 C. ochreata, 9 ♂, Gainesville, Paynes Prarie State Park, Alachua Co., Florida, USA 10 ♀ holotype, Anthony, Marion Co., Florida, USA 11 ♂, Middleton Creek, Franklin Co., Mississippi, USA 12 ♀, Sweetbay Bogs, Stone Co., Mississippi, USA.
Clemensia albata Packard, 1864, p. 101. Type locality. “Norway, Me. (Mus. Comp. Zool., Smith), Brunswick, Me., August.” [unknown]. Note. The holotype of albata should be with other Packard types at MCZ, but the online inventory of MCZ types (mczbase.mcz.harvard.edu) indicates it is not, and as such iot may no longer be extant.
Uxia
albida
Walker, 1866, p. 1897. Type locality. “North America;” female type [
Repa cana Walker, 1866, p. 1898. Type locality. “United States;” male type [BMNH]. Note. The holotype is a male without an abdomen.
Clemensia albata is the most common Clemensia in most of eastern North America south of the boreal forest region. In the northeastern US and the Appalachians, C. albata can be confused with C. umbrata, and a differential diagnosis is presented in the C. umbrata account. Along the Atlantic coastal plain from North Carolina to Florida and across the southern states to at least Mississippi, C. albata can occur with C. ochreata, and the two can be difficult to separate. Clemensia albata usually differs from C. ochreata in its pure white rather than ochre-white ground colour, less prponounced dark spots along the forewing costa, a less distinct antemedial dark patch, lack of a well-defined ventral hindwing medial band, and narrower uncus (Figure
The larvae feed on blue-green algae and lichens growing on tree trunks (
Examined specimens identified with certainty as C. albata are mapped in Figure
There is some uncertainty in the identity of the holotype of Clemensia albata, given the type locality and the similarity to C. umbrata, but two facts help in ascertaining what taxon the name albata applies to: the types were collected in August in southern Maine, pointing to second-brood specimens of the bivoltine eastern species (versus univoltine C. umbrata flying mostly in July), and the wing expanse given as 0.83 inch, or 21.1 mm, which is smaller than most eastern C. umbrata. Examined specimens from regions adjacent to the C. albata type locality in southern Maine (Figure
Holotype ♀ (Figure
The name ochreata is a noun in opposition and refers to this species’ characteristic ochre forewing tint.
Very similar to Clemensia albata, but differing from that species by the smaller mean forewing length, more extensive, and brighter ochreous scales along the forewing antemedial and postmedial lines; overall more contrasting pattern, especially the heavier costal dark spots (most pronounced on the forewing underside), and the more prominent and better-defined medial dark patch basad of the antemedial line. Internally, the basal diverticulum of the male vesica has much smaller spicules than in C. albata. The valve shape is proportionally shorter and stouter than that of C. albata. In females, the corpus bursae is more elongate than that of C. albata and with shorter spinules lining the interior; the appendix bursae is also smaller and more narrowly joined to the corpus bursae in C. ochreata. Compared to C. albata, overall shape and structure of the bursa copulatrix is very similar, but the spinules lining the inside of the bursae are smaller in C. ochreata.
Head: frons and vertex with mix of dirty-white and dull grey-brown scales; palpi grey brown, terminal (3rd) segment 0.5 × length of second segment; male antenna filiform and finely ciliate, segments approximately width 0.9 × that of length; dorsally with grey-brown scales, finely ciliate ventrally, with two mediolateral setae, these equal in length to antennal segment; female antenna similar to that of male but narrower and less ciliate. Thorax: vestiture predominantly dirty white with scattered dull grey-brown scales; pro- and mesothoracic legs appearing striped, grey brown with a ring of dirty white scales at apices of segments, in addition to a mid-tibial pale ring; metathoracic leg entirely dirty white; metepisternal tymbal unscaled, with 9–11 fine grooves. Abdomen: vestiture light grey brown dorsally and ventrally; females with apical tuft of incurved setae; males with a series of paired setal tufts ventrally on segments 7, 8 and 9, in addition to two paired lobe-like setal tufts situated within a pouch between segments 7 and 8. Forewing: ground color slightly ochre, dirty white; basal line indistinct greyish ochre; antemedial line double, often only with distal line defined; grey ochre; medial line grey ochre; postmedial line an irregular row of disconnected grey-black splotches; terminal line grey black, thin or incomplete, interrupted with white at vein termini; fringe dirty white, often interrupted with grey patch at apex and medial area; reniform spot usually very pronounced, grey black; orbicular spot small and grey black, often absent or tiny; ventrally with large dusky-grey patch medially, pattern inconspicuous, wing edges dull tan white, interrupted at costa by four costal grey-black markings; males with a brown setal (androconial?) tuft in medio-anal area; wing pattern of females similar, but usually with the dark markings reduced considerably, giving the impression of a paler moth. Hindwing: dirty white with faint, indistinctly delineated fuscous area in distal third; ventrally with slightly darker fuscous patch in anal angle and discal spot, and faint medial line. Male genitalia (Figure
The immature stages and larval hosts are unknown, but larvae likely feed on algae or lichens growing on tree bark. There are multiple broods starting in March and continuing into September.
The Atlantic coastal plain from North Carolina southward into Florida and westward to eastern Texas (Figure
The late Douglas Ferguson deemed C. ochreata to be closely related to the Mexican C. patella (Druce), and the latter name was therefore applied by him to this taxon (
Neighbour-joining tree of representative mtDNA barcode haplotypes in North American Clemensia species. Sample size and locality are given in brackets, with number of specimens indicated after two-letter state/province abbreviation. Lacinipolia sareta variation is divided into five haplogroups, A–E. Voucher specimen data is given in Suppl. material
Although Clemensia albata has been treated as a single species, the concordant variation in phenotype, morphology, phenology distribution, and DNA barcode clearly supports a concept of three species. Further research is needed on Clemensia life histories, given the current taxonomic changes. Specifically, it is not known if the differences in voltinism and phenology correspond to differing winter diapause strategies, different food plant requirements or how the larvae differ morphologically. The distributional limits of the genus also requires refining, such as the northwestern range limits of C. albata and the full extent of the distribution of C. umbrata in the eastern US.
Richard Brown, Martin Honey, Greg Pohl, Danny Shpeley, and Thomas Simonsen kindly provided access to specimens in their care. James Adams, Jason Dombroskie, and Val Deziel provided specimens and/or additional data. Jocelyn Gill and Christi Jaeger provided technical support.
Table S1. DNA barcode voucher specimen data.