1 April, 2020

Volume 8 Number 5

The Taxonomic Report

OF THE INTERNA TIONAL LEPIDOPTERA SURVEY

ISSN 2643-4776 (print) / ISSN 2643-4806 (online)

A phenotypic comparison of regional populations of

Hemileuca maia (Drury, 1773) with designations of new subspecies

(Bombycoidea, Saturniidae, Hemileucinae).

Harry Pavulaan

606 Hunton Place NE

Leesburg, VA. 20176

intlepsurvey@ gmail.com

ABSTRACT. Following refinement of the type locality of Hemileuca maia to the Long Island

Pine Barrens of New York State by the author (Pavulaan, 2020), an evaluation of phenotypic characters of

regional populations of H. maia is presented. The Long Island population is the nominotypical subspecies.

Populations in southeastern coastal New England and offshore islands are presently considered

nominotypical maia . However, several continental inland populations show evidence of subspecific

variation. Four new subspecies are designated. Detailed phenotypic information of other interior regions

is lacking.

Additional key words: Pitch Pine Barrens, Scrub Oak Plains, isolate, Menyanthes trifoliata.

ZooBank registration: urn:lsid:zoobank.org:pub:3595D21C-4FDE-4336-A588-4E68195E1118

INTRODUCTION

The Buckmoths of North America are a bewildering blend of intergrading phenotypes that have

been the subject of numerous studies (Ferguson, 1971; Tuskeset al., 1996; Rubinoffet al., 2017; Dupuiset

al., 2018). Results of these studies are inconclusive over where to draw taxonomc limits. Michener

(1952) proposed a subdivision of genus Hemileuca into four subgenera: Hemileuca (Walker, 1855),

Pseudohazis (Grote & Robinson, 1866), Euleucophaeus (Packard, 1872) and Argyrauges (Grote, 1882).

Nestled within subgenus Hemileuca is the Hemileuca maia complex, presently considered to be a closely-

related group of species and unnamed populations of species H maia. This group is characterized by

variation in ground color (gray to black), bold median bands (white to yellow), and scale translucence.

Described members include: H. maia (Drury, 1773), H. grotei (Grote & Robinson, 1868), H. nevadensis

(Stretch, 1872), H. juno (Packard, 1872), H. diana (Packard, 1874), H lucina (Henry Edwards, 1887), H.

artemis (Packard, 1893), H. peigleri (Lemaire, 1981), and H slosseri (Peigler & Stone, 1989). The

present paper focuses on subspecific divisions within the species H. maia.

Most members of the H. maia complex feed on various species of Quercus sp. (Oaks), but several

peripheral populations in the northern states specialize on non -Quercus hosts. Eggs are laid in lines

encircling thin branches of the host or alternate substrates. Eggs then overwinter. Larvae emerge in

spring and feed gregariously on the primary host until the last two instars, when they disperse seeking

alternate, unrelated hosts to complete development. Pupation occurs in soil, leaf litter, or gravel

throughout the summer. Eclosion generally occurs immediately after the first frost in late September in

northern latitudes, then progressively later with the advance of fall in the southern states. Emergence as

late as December in Louisiana and February in Florida is not unusual. Some pupae do not eclose during

the first year, delaying eclosion for two, three or more years. Delayed eclosion is likely a response to

unpredictable weather trends (e.g. rainfall amount) and other environmental conditions.

Tuskes et al. (1996) comprehensively reviewed adaptations to local environments within the H.

maia complex. These authors described varying degrees of ecologic and genetic differentiation. They

pointed out "some populations within the maia complex defy neat categorization into species and

subspecies". Rubinoff et al. (2017) suggested "most of the populations in the H. maia complex have been

viewed as isolated relics surviving on the remaining 'islands' of what were formerly more widespread

suitable habitat" which now tend to be localized and fragmented. They found that isolated populations

frequently gave rise to phenotypic variation in adults and larvae with some switching to non -Quercus

hosts. They concluded these differences were still within conspecific level and there was little to no

genetic difference among members of the complex. Branching topology of their genomic trees showed

some members of the complex clustered in neat groups. For example, Florida, Lousiana and Bog

Buckmoth populations each formed tight groupings in COI trees and concatenated mt-DNA and nuclear

DNA maximum likelihood trees. Their elongation factor 1-alpha (EFla) and carbamoyl phosphate

synthetase (cpsi) maximum likelihood trees showed New York samples of the Bog Buckmoth clustering

together, separate from the rest of the H. maia complex.

A subsequent analysis by Dupuis et al. (2018), to a large degree, corroborated the findings of the

Rubinoff et al. (2017) study. These authors expressed: "Local adaptation can be a fundamental

component of speciation, but its dynamics in relation to gene flow are not necessarily straightforward ...

The charismatic, day-flying moths of the Hemileuca maia species complex ... are such taxa, as they are

geographically widespread, exhibit considerable ecological and morphological variability and host and

habitat specificity, but lack genetic differentiation across their range." They noted that previous studies

based on sequences of one mitochondrial and three nuclear genes failed to identify genetic divergence

even between taxa treated at species rank within the complex. To assess population structure and

genomic relationships of the H. maia complex, their study employed genome-wide single nucleotide

polymorphisms in a combination of population genomic and phylogenomic approaches. They uncovered

fine-scale population divisions among members of the complex. A maximum-likelihood consensus tree

and a distance-based phylogenetic network showed distinct clustering of various groups, including some

named species within the complex and other groups not currently recognized at any level below species

H. maia. Of note, their Floridian and Louisiana populations clustered distinctly and separately from other

members of the complex. They also found the Bog Buckmoth samples from New York clustered

separately from the rest of the complex, primarily due to geographic isolation, and that this "definitely

satisfy[ies] the criteria of genetic differentiation." The authors suggested that interpretation of their

results would support treating any cluster within the maia complex as a distinct species or subspecies.

The status of proserpina (Fabricius, 1775) is uncertain. The holotype and co-type in the Hunterian

Museum (University of Glasgow) do not reliably fit the nominotypical phenotype or any of the inland

phenotypes (Pavulaan, 2020). The original description merely gives "America". It is not possible to

reliably determine where the specimens originated (Jeanne Robinson, pers. corr.) or whether they

represent any population of maia or of a different species. Currently, there are two possible treatments of

proserpina. One is to treat proserpina as as a junior synonym of H. maia maia , and the other is to treat it

as nomen incognitum (my preference). Lastly, the one specimen residing in the Macleay Museum

(University of Sydney), often thought to be the model for Drury's maia , does not match the drawing in

the original description (Pavulaan, 2020).

PHENOTYPIC FINDINGS & TAXONOMIC DECISIONS

Recent studies have demonstrated that several "ecotypes" in the H. maia complex display low

levels of genetic divergence (Rubinoff et al ., 2017; Dupuis et al., 2018), while maintaining their

ecological differences in sympatry. My analysis of phenotypic traits of members of the complex reveals a

number of stable phenotypes, which I propose to be subspecific. Some of these subspecies may

eventually warrant elevation to species. Hopefully this paper will help to further conservation efforts at

the federal level.

The Bog Buckbean population of northern New York State, utilizing Menyanthes trifoliata as

larval host, has unique status and is recognized as endangered in that State. However, it is unnamed and

lacks formal description. To complicate matters, a large, complex zone of integration occurs in the Great

Lakes region from Ohio to Minnesota that is still unresolved and under study as of this writing.

Compheating any resolution are a broad variety of full and intermediate phenotypes encompassing maia,

nevadensis , lucina , and the undescribed "Bogbean Buckmoths", ah of which reside in a complex

patchwork of habitats, mainly wetland/bog types. Scholtens & Wagner (1994) studied and compared

"transitional phenotypes" in Michigan to maia , nevadensis and lucina and concluded that the Michigan

populations are a single variable species, spanning the entire range of phenotypes, hosts and habitats

(mainly wetlands) of the three named species, thus calling to question the taxonomic distinctness of the

three Hemileuca species. Recorded hosts (both primary and secondary) in the Great Lakes complex

include: Betula nana (Dwarf Birch), B. pumila (Bog Birch), Dasiphora ( =Potentilla ) fruiticosa (Shrubby

Cinquifoil), Menyanthes trifoliata (Buckbean), Populus deltoides (Eastern Cottonwood), P. tremuloides

(Quaking Aspen), Quercus velutina (Black Oak), Salix behbiana (Bebb's Willow), S. Candida (Sageleaf

Willow), S. discolor (Pussy Willow), S. exigua (Narrowleaf Willow), S. humilis (Prairie Willow), S.

pedicellaris (Bog Willow), S. petiolaris (Meadow Willow), S. serissima (Autumn Willow), Spiraea alba

(White Meadowsweet) and Vaccinium macrocarpon (Cranberry). Metzler & Lucas (1990) successfully

reared larvae from northern Ohio on S. nigra (Black Willow), but larvae reared on "Pin Oak" did not

complete development and died. Gratton (2006) documents use of Lythrum salicaria (Purple Loosestrife)

in Wisconsin.

Tuskes et al. (1996) consider Great Lakes transitional populations reflect incomplete reproductive

isolation between maia and nevadensis with introgression occurring over a broad area. This view suggests

nevadensis may be better treated as a subspecies of maia. It also suggests that the Great Lakes complex

may be viewed as three species if only adult morphology is considered: dark southernmost populations

(southern Ohio) being maia; central and western Great Lakes populations being nevadensis ; and the

northern translucent populations being lucina. An outlying wetland-dwelling group of populations in

northwestern New Jersey feeds on B. pumila (Bog Birch) and Dasiphora ( [=Potentilla ) fruticosa (Shrubby

Cinquefoil); this group may represent a nevadensis isolate.

Scholtens & Wagner (1997) studied possible north-south clinal variation in a line running through

Michigan and Ohio. They concluded the Great Lakes group of populations comprised a single species in

which small, washed-out H. lucina-Yke forms occupied wetlands in the upper Michigan peninsula and

larger, heavily-marked maia- like forms occurred the upland wooded area in southern Ohio. Two

intermediate study sites in the lower peninsula of Michigan were attributed to nevadensis-Vke phenotypes.

In an earlier significant study , Metzler & Lucas (1990) noted ecological differences between southern and

northern Ohio populations of H maia. The northern Ohio population resided in oak barrens and utilized

Salix sp. (Willow) as larval hosts, while the southern Ohio population inhabited mature oak forests,

utilizing on Quercus sp. (oaks). They associated the northern Ohio population with the upper Midwest

zone of intergradation between H. maia and H. nevadensis. They suggested the northern and southern

Ohio populations may not be conspecific. Given the considerable distance between these populations, one

might consider them as separate taxa with the southern ones being more aligned with true H. maia.

3

The present paper focuses on regional phenotypic variation, describing phenotypic characters and

providing taxonomic names. I have not relied heavily on life history studies. The hope is this paper will

lay the groundwork for continued studies of subspecific taxa and promote conservation efforts such as in

the case of Bogbean Buckmoth. Specimens used in this study are mainly in the author's personal

collection and in the McGuire Center for Lepidoptera and Biodiversity. Access to specimens in other

institutional collections was generally unproductive; specimen numbers were often insufficiently small or

too disorganized.

SUBSPECIES

Hemileuca maia maia Drury (1773)

Coastal Barrens Buckmoth

The nominotypical taxon (Figs. 1, 2 & 3) was described from "New York" and determined to most

likely originate in the Pitch Pine Barrens region of eastern Long Island. Common name Coastal Barrens

Buckmoth (for Hemileuca maia ssp. 5) per New York Natural Heritage Program (2020) and NatureServe

Explorer (2020) which reflects its limited distribution in coastal Pitch Pine Barrens habitat in southeastern

New England and on Long Island, N.Y.

Type locality : Neotype male: Oct. 21, 2017, Long Island Avenue, north of Deer Park train station,

Edgewood, Suffolk County, New York, leg. H. Pavulaan (Pavulaan, 2020). Neotype is deposited in the

McGuire Center for Lepidoptera & Biodiversity, Gainesville, EL. [5 additional male and 5 female

specimens (Westhampton, Suffolk Co., N. Y.) analyzed in this study are deposited in the McGuire Center

for Lepidoptera & Biodiversity, Gainesville, FL.; additional specimens remain in the author's collection.]

Range : Confined to eastern Long Island, New York (Fig. 16). Most frequently observed in the

Westhampton Dwarf Pine Plains Preserve and in the Edgewood Oak Brush Plains Preserve. Additional

populations are in southeastern New England, Cape Cod, Martha's Vineyard and Nantucket, which are of

the same phenotype, are often referred to as the same taxon.

Habitat : Primarily pitch pine barrens, scrub oak plains and maritime shrublands or sandplains where the

hostplant Quercus ilicifolia abounds. Selfridge et al. (2007) state that H. maia is restricted to remnant

Pinus rigida (Pitch Pin e)/Quercus ilicifolia (Scrub Oak) habitat in the northeastern United States. This

habitat is becoming more fragmented and threatened by multiple factors, thus bringing conservation

attention to the nominotypical subspecies. Fire suppression in the fire-dependent habitat is a threat and

requires considerable management considerations.

Flight period : NY (Long Island): Oct. 4 - 28. Southeastern New England: Sept. 27 - Oct. 25.

Hosts : Quercus ilicifolia (Scrub Oak) on Long Island, N.Y., Plymouth County, MA. and Martha's

Vineyard, MA. Also on Quercus prinoides (Dwarf Chinquapin Oak). Haggerty (2006) reared larvae on

Q. alba (White Oak) and Q. stellata (Post Oak). Larvae are known to wander to additional, non -Quercus,

hosts in the later instars, such as Salix (Willow sp.) and Populus (Aspen sp.) (NatureServe Explorer,

2020). In Edgewood, N.Y., I have found them frequently on Prunus serotina (Black Cherry) in July. In

West Greenwich, R.I., I once found a clutch of first instar larvae on P. serotina but could not locate an

egg ring to confirm oviposition on that host.

Description : Male FW length 21-28 mm., female FW length 28-30 mm. The smallest of all maia

populations. Forewings slightly more rounded than other subspecies as described. Wings generally with

a variable dark gray appearance, darker toward the outer margin, and highly translucent (thinly-scaled)

[text can be read through the wings when placed against the wings]. Median bands primarily white with a

bare hint of yellow tint in some specimens. The forewing band directly intersects the discal streak which

usually breaks the band into two segments in most examined specimens. The posterior portion of the

median band may be partially faded in many individuals. Females are more uniformly gray than males,

and have a distinct black wing margin. The Long Island, N.Y. population has larvae characterized by

having a distinct yellow lateral line.

Hemileuca maia sandra Pavulaan, 2020 - New Subspecies

Eastern Buckmoth

ZooBank registration: urn:lsid:zoobank.org:act:9EDBE62F-2415-4EC0-BE24-6E2C4BB20712

This taxon (Figs. 3, 4 & 12) represents interior continental (inland) populations. Insufficient

published or available collected material makes it difficult to ascertain the full extent of its distribution.

Common name Eastern Buckmoth per NatureServe Explorer (2020) reflects distribution throughout the

interior eastern United States. The New York Natural Heritage Program (2020) refers to this as Inland

Barrens Buckmoth and considers this the "nominal" subspecies because adults are determined to fall

within the range of variation of southern populations. The subspecies is named in honor of my older

daughter, who provides invaluable field assistance in all my lepidoptera studies.

Type locality : Holotype (male): Oct. 15, 2019, ex-larva, Chatsworth, Woodland Township, Burlington

Co., New Jersey, leg. I. Osipov. 24 paratypes (20 males, 3 females) from same location. Holotype,

allotype (female) and 8 paratype specimens (6 males, 2 females, from TL) are deposited in the McGuire

Center for Lepidoptera & Biodiversity, Gainesville, FL.; the remainder in the author's collection. A

series of 32 southern Indiana specimens (males) were analyzed for morphological comparison and

matched the range of variation in the New Jersey series [10 specimens from Mishawaka and West

Lafayette, IN. are deposited in the McGuire Center for Lepidoptera & Biodiversity, Gainesville, FL.; the

remainder in the author's collection.]

Range : Known to be common in parts of the southern New Jersey Pine Barrens region. N.J. phenotype

matches specimens from Virginia, Georgia, southeastern Ohio, southern Indiana, northern Alabama and

northern Mississippi. This subspecies likely ranges west to Oklahoma (Fig. 16). It is likely that "inland"

populations in New York, from the Shawangunk Mountains to Albany are represented by this taxon.

Habitat : In New Jersey and inland New York, primarily Pitch Pine/Scrub Oak barrens but also in dry,

mixed forest dominated by Quercus. In northern Virginia, mixed deciduous forest with presence of

different species of Quercus , but apparently Q. ilicifolia is absent. Reportedly associated with pine

barrens and sand barrens in other northern states: NY (inland sites), GA, OH, PA, VA, WV. In the

Appalachian Mountains, the habitat is mixed, dry Quercus-dormnaX^d woodland and granite balds. In the

southeastern coastal plain, the habitat is described as sandhills. Fragmentation and fire suppression of

Pitch Pine/Scrub Oak habitat in northern portions of the species' range are a threat to the habitat.

Flight period : Southern IN: Sep. 27 - Nov. 2. NY (populations around Albany, believed to be this

subspecies): Sept. 25 - Oct. 5. NJ: Sep. 27 - Nov. 1. PA: Sept. 25 - Nov. 1. MD: Oct. 25 - Nov. 12.

VA: Oct. 17 - Nov. 12. WV: Oct. 8 - Oct. 19. NC: Nov. 4 - Dec. 26. SC: Jan. 1. Southeastern OH: Oct.

18 - Nov. 4. KY: Oct. 3 - Nov. 14. TN: Nov. 14 - Nov. 30. GA: Nov. 4 - Dec. 2. Northern AL: Nov. 7

- Dec. 3. Northern MS: Dec. 7.

Hosts : Quercus alba (White Oak), Q. bicolor (Swamp White Oak), Q. falcata (Southern Red Oak), Q.

hemisphaerica (Darlington Oak), Q. ilicifolia (Scrub Oak), Q. laevis (Turkey Oak), Q. marilandica

(Blackjack Oak), Q. montana (Chestnut Oak), Q. muehlenbergii (Chinquapin Oak), Q. prinoides (Dwarf

Chinquapin Oak), Q. rubra (Red Oak), Q. stellata (Post Oak) and Q. velutina (Black Oak). Larvae are

known to wander to additional, non -Quercus, hosts in the later instars, such as Prunus cerasus (Sour

Cherry). In Clifton, VA found on Rosa multiflora (Multiflora Rose) in July; in KY and MD, found on

cultivated Fragaria (Strawberry sp.). In laboratory experiments, Smith (1974) successfully reared larvae

from eggs collected on Q. ilicifolia in Colonie, N.Y. on alternate hosts Q. chrysolepis (Canyon Live Oak)

and Salix hindsiana (Sandbar Willow). Leeuw (1974) reported rearing them on Salix babylonica

(Weeping Willow). Many other hosts are listed in Tietz (1952, 1972) and Heppner (2003), but they do

not specify what states or regions these records are from.

Description : Male FW length 24-31 mm., female FW length 29-36 mm. Larger than ssp. maia. Wings

black, opaque [text cannot be read through the wings when placed against the wings]. Median bands

white but with slightly more yellow tint than in ssp. maia. Fore wing median band intersects the discal

streak which breaks the band into two segments in less than half of individuals, but in most specimens the

median band is located exterior to the discal streak, thus maintaining a continuous band. Indiana and

Virginia specimens tend to have the continuous median band (inner edge of the median band intersects the

discal streak) and the bands are clearly cream-colored. Specimens from northern Alabama and northern

Mississippi are deep black, matching northeastern populations, and do not possess the brownish-black

color of the nearby Gulf Coast population.

Hemileuca maia warreni Pavulaan, 2020 - New Subspecies

Florida Buckmoth

ZooBank registration: urn:lsid:zoobank.org:act:281 A5C71-A821-47B4-BDA6-CFDBF4A5E251

This taxon (Figs. 5, 6 & 13) represents peninsular Florida populations and was only recently

thought to be a distinct subspecies. The subspecies is named in honor of Dr. Andrew D. Warren, Senior

Collections Manager, McGuire Center for Lepidoptera & Biodiversity, who provided considerable

information regarding Florida populations.

Type locality : Holotype (male): Jan. 24, 1984. Deltona, Volusia Co., Florida, leg. L. C. Dow. 50

paratypes from Alachua (Archer, Gainesville), Clay (Camp Blanding), Duval (Jacksonville), Putnam

(Katherine B. Ordway Preserve), Seminole, Taylor (Steinhatchee) and Volusia (Deltona, Cassadaga)

Counties. [Holotype, allotype (female) and all paratype specimens deposited in the McGuire Center for

Lepidoptera & Biodiversity, Gainesville, FL.]

Range : Primarily peninsular Florida (Fig. 16): Duval to Palm Beach counties along the east coast, west to

Taylor County.

Habitat : Pine-Oak scrub.

Flight period : Dec. 25 - Feb. 4 (peak flight in Jan.).

Hosts : Quercus laevis (Turkey Oak) reported from northern parts of Florida (e.g. western Marion,

Alachua, Clay counties), Quercus myrtifolia (Myrtle Oak) reported from southeastern portions (e.g.

Marion Co. (Ocala National Forest), Palm Beach Co.).

Description : Male FW length 23-28 mm., female FW length 27-35 mm. [Holotype (male) FW length is

23.7 mm.] Smaller than the other subspecies except ssp. maia. Forewings have a slightly more elongated

appearance than other subspecies. In many individuals, the forewings are decidedly more "pointed" than

other subspecies. Wings black to dark brownish-black, solid opaque. Median bands cream-white with a

tendency for the forewing bands to be quite narrow in most specimens. Some individuals have the

posterior portion of the fore wing median band faded or reduced to small white patches. In some, the

posterior portion of the band is absent and the anterior portion reduced to a small white triangle adjacent

6

to the discal streak, leaving the fore wing almost all black. The fore wing median band is positioned

exterior to the discal streak in most specimens, thus maintaining a continuous band (inner edge of the

median band intersects the discal streak). The discal streak is enlarged in most specimens, more so than

in other subspecies.

Hemileuca maia Orleans Pavulaan, 2020 - New Subspecies

Gulf Coast Buckmoth

ZooBank registration: urn:lsid:zoobank.org:act:EB65773F-B937-45F5-A58D-9C30478A8281

This taxon (Figs. 7, 8 & 13) represents Louisiana populations, mainly around New Orleans and the

Mississippi River delta. This is a unique phenotype with a variable brownish-black ground color.

Type locality : Holotype (male): Dec. 11, 2003. 4.2 mi. NE of Abita Springs, sec. 24.T6.SR12E, St.

Tammany Parish, Louisiana, leg. V. Brou. 45 paratypes (males) from St. Tammany Parish, 1 paratype

(female) from Ascension Parish. [Holotype, allotype (female) and 16 male paratype specimens deposited

in the McGuire Center for Lepidoptera & Biodiversity, Gainesville, FL., the remainder in the author's

collection.]

Range : Primarily southern portions of Louisiana around New Orleans and adjacent portions of

Mississippi (Fig. 16). Range into eastern Texas requires morphological examination of specimens, but all

viewed images of eastern Texas maia are black. Some Texas records may be confused with H. peigleri

but adult morphology is very different.

Habitat : Known to be an urban pest on host trees in New Orleans and Baton Rouge, experiencing heavy

infestations, leading to mass defoliation of city trees (Martinat et al., 1997).

Flight period : LA (New Orleans region): Nov. 15 - Jan. 9. Eastern TX records are: Dec. 4-17 but may

not be ssp. Orleans.

Hosts : Quercus virginiana (Live Oak). Martinat et al. (1997) conducted host suitability studies to

determine alternate host suitability. They report that late-instar larvae have been found on almost any

type of foliage but it was unknown whether they can complete development on non -Quercus species. In

their study, 1 st through 4 th instar larvae were raised on Quercus nigra (Water Oak), then switched to

various alternate hosts in the 5 th instar, while others were successfully reared to pupation on Q. nigra.

Suitable alternate hosts included: Q. velutina (Black Oak), and Prunus serotina (Black Cherry). Lower

survival rates occurred on Lagerstroemia indica (Crape Myrtle), Liquidambar styraciflua (Sweetgum),

and Salix nigra (Black Willow).

Description : Male FW length 25-32 mm., female FW length 35 mm. The largest of all maia populations.

Wings decidedly brownish-black, solid opaque. The brownish tint is variable. While some individuals

are black, a small percentage are clearly brown. Median bands are cream-colored. The forewing median

band is located exterior to the discal streak in most specimens, thus maintaining a continuous band (inner

edge of the median band intersects the discal streak). Some individuals have the posterior portion of the

forewing median band faded or reduced to small white patches

Hemileuca maia menyanthevora Pavulaan, 2020 - New Subspecies

Bogbean Buckmoth

ZooBank registration: urn:lsid:zoobank.org:act:B7D0030D-lC45-4733-AlCE-79B6515AC4El

This taxon (Figs. 9, 10 & 15) represents a small grouping of wetland-associated populations in

northern New York State, around Ottawa, Ontario and in southeastern Wisconsin. Rubinoff & Sperling

(2004) found no marked genetic divergence from H. maia populations nearby in New York State, lending

to its status as a recent postglacial taxon. Long considered to be a unique ecological "form" of H maia ,

adapted to use of a particular hostplant ( Menyanthes trifoliata ), but the lack of an adequate description

and naming hinders conservation efforts. In the absence of formal description, and to further conservation

efforts in New York and Ontario, the Evolutionary Significant Unit (ESU) concept was employed. Fraser

& Bematchez (2001) define the ESU concept as: "Segments of species whose divergence can be

measured or evaluated by putting differential emphasis on the role of evolutionary forces at varied

temporal scales [within a species]".

The present paper resolves at least one issue, that of formal description and naming. One of the

most difficult hindrances to the study of this taxon is locating any specimens able to serve as types. Only

the McGuire Center for Lepidoptera and Biodiversity had institutional specimens, which serve as the

holotype and paratypes. Ross Layberry provided information on specimens collected in Ontario in 1984

(pre-act). Additional analysis of New York, Wisconsin, and Ontario phenotypes came from many internet

resources and published works (Handfield, 2011). It is not clear whether the New York, Wisconsin and

Ontario populations all represent the same exact taxon. COSEWIC (2009) states that genetic exchange

between U.S. and Canadian populations does not occur due to distance between population groupings.

Further research is warranted. This may actually represent a full species. The Bogbean Buckmoth was

granted state protection as threatened in 1999 in New York State, though it has not been formally

described or named. It is also protected as endangered in Canada as of 2009, and also in Ontario under

the Endangered Species Act, 2007.

Common name Bogbean Buckmoth (. Hemileuca sp. 1) per New York Natural Heritage Program

(2020) and NatureServe Explorer (2020).

Type locality : Holotype (male): Sep. 22, 1985. Brennan Beach Camp, Pulaski, Oswego Co., N.Y. Leg.

P. Savage. Allotype (female), 9 male paratypes and 4 female paratypes, same data as holotype. All

specimens deposited in the McGuire Center for Lepidoptera & Biodiversity, Gainesville, EL.

Range: Confined to six known fens in Oswego County, N.Y. and in four fens near Ottawa, Ontario (Fig.

16). Kruse (1998) provides detailed information on two sites in Marquette and Ozaukee Counties in

Wisconsin.

Habitat : Described as minerotropic [stream or spring-fed] fens at the east edge of Lake Ontario and open

graminoid fens (New York Natural Heritage Program, 2020); open, low-shrub fens (Government of

Canada Species Profile, 2020), and open, calcareous, graminoid and low shrub fens in Ontario

(COSEWIC, 2009). COSEWIC (2009) gives the habitat description of the two groupings of New York

as: (1) a complex of several discrete, rich, shrub-dominated fen openings behind barrier dunes on Lake

Ontario at one site, and (2) an inland complex of edge-dominated floating peat mats on lake edges. This

subspecies (or species) is extremely vulnerable to extirpation in New York, because its habitat is rapidly

degrading and is significantly threatened (New York Natural Heritage Program, 2020). Habitat threats

include succession by invasive Typha angustifolia (Narrowleaf Cattail), Phragmites australis (Common

Reed), Lythrum salicaria (Purple Loosestrife), and Frangula alnus (Glossy Buckthorn); hydrologic

alteration resulting in lowering of the Lake Ontario water level; nutrient pollution run-off from

surrounding areas; surface flooding at two fen sites; and parasitoid wasps and other predators. Pesticide

application in the habitat could be disastrous. In Ontario, there are nearly identical threats to the habitat

(COSEWIC, 2009).

Flight period : Sept. 9 to Oct. 12.

Hosts : A rather complicated series of choices (COSEWIC, 2009; Government of Canada Species Profile,

2020) across a broad range of suitable hostplants. In Ontario, females were observed depositing egg rings

mostly on Myrica gale (Sweet Gale), Betula pumila (Bog Birch), but also other low shrubs such as

Spiraea alba (Narrow-leaved Meadowsweet), Salix petiolaris (Slender Willow), Salix bebbiana (Bebb's

Willow) and odd choices such as Phragmites australis (Common Reed), Muhlenbergia glomerata (Marsh

Muhly), Solidago (Goldenrod sp.), Aster (Aster sp.), and Scirpus (Rush sp.) [The primary host Menyanthes

trifoliata has already dried by the time females are looking to oviposit]. Many of these plants are

unsuitable for larval development. Newly-hatched larvae in the following year seek out Vaccinium

macrocarpon (Bog Cranberry), later switching to M. trifoliata as it leafs out. Late instars disperse widely,

being found on B. pumila, S. petiolaris , S. bebbiana , Salix pedicellaris (Bog Willow), and Spiraea alba

(Narrow-leaved Meadowsweet).

In a New York study (Pryor, 1998), similar observations were obtained of ovipositions on M. gale ,

Acer rubrum (Red Maple), Woodwardia ( =Anchistea) virginica (Virginia Chain Fern), Alnus incana

rugosa (Speckled Alder), Carex (Sedge sp.), Chamaedaphne calyculata (Leatherleaf), Salix pedicellaris

(Bog Willow), S. alba , and Cornus sericea (-stolonifera ) (Red Osier Dogwood). Likewise, many of

these plants are unsuitable for larval development. Larvae have been observed leaving unsuitable plants

and seeking out V. macrocarpon and later switching to M. trifoliata as it leafs out. Larvae were further

found throughout summer feeding on: A. incana rugosa, Aronia melanocarpa (Black Chokeberry), Carex

(Sedge sp.), C. calyculata, Ilex verticillata (Common Winterberry), M. trifoliata, Quercus (Oak sp.), S.

pedicellaris , S. alba, and V. macrocarpon. [The name Buckbean Buckmoth might thus be a misnomer.]

Description : FW length: males 22-32 mm; females 26-36 mm. [Holotype (male) FW length is 27.6 mm.]

Despite published claims, this subspecies is very distinct and nearly identical to H lucina. Larger than

ssp. maia. Wings light gray and highly translucent, with the cream-tinted median bands nearly

transparent [text can easily be read through the wings when placed against the wings]. The median bands

on both sets of wings are much wider than in all other maia subspecies, generally completely enclosing

the forewing median discal streak and covering about 1/3 of the interior of the wings. The basal third of

the forewings are darker than the outer third. Female wings are not as broad as the other maia subspecies,

the wings are more rounded and posess a dark gray wing margin on both sets of wings. The median band

pattern in females is remarkably similar to H. nevadensis females.

This subspecies is the most unique of all the maia- complex populations and likely represents a

different species-level taxon. It appears to be nearly identical in morphology to H. lucina. Handfield

(2011) describes it as Hemileuca sp. near lucina due to great similarity to H lucina. Legge et al. (1996)

suggest this taxon is an evolutionary significant unit, possibly a separate species. Crandall et al. (2000)

consider this an evolutionary significant unit (ESU), based on ecological and genetic data, and further

suggest that these are more relevant for conservation efforts. They consider the evolutionary heritage of

ESU's based on ecology will better foster conservation management. Buckner et al. (2014) state that Bog

Buckmoth populations are not consistently distinguishable from other H. maia populations on the basis of

any diagnostic set of color or other morphological characters. This paper presents a different conclusion.

9

of*?

Fig. 1. Hemileuca maia maia Drury (1773). Neotype. Oct. Fig. 2. Hemileuca maia maia Drury (1773). Neotype (same

21, 2017, Edgewood, Long Island, N. Y. Leg. H. Pavulaan specimen), ventral view.

Male. Dorsal view.

1

of

Fig. 3. Hemileuca maia sandra Pavulaan (2020). Holotype. Fig. 4. Hemileuca maia sandra Pavulaan (2020). Holotype

Chatsworth, Woodland Township, Burlington Co., N.J., leg. (same specimen), ventral view.

S&J

Fig. 5. Hemileuca maia warreni Pavulaan, 2020. Holotype. Fig. 6. Hemileuca maia warreni Pavulaan, 2020. Holotype

Jan. 24, 1984. Deltona, Volusia Co., FL. Leg. L. C. Dow. (same specimen), ventral view.

Male. Dorsal view.

10

12

13

SUBSPECIES RANGE MAP

ACKNOWLEDGMENTS

Many thanks to: Dr. David M. Wright for very helpful comments and editorial review; Dr.

Andrew D. Warren (Senior Collections Manager, McGuire Center for Lepidoptera and Biodiversity) for

early comments on this project; Crispin Guppy for comments; Vernon A. Brou, Jr. for Abita Springs,

Louisiana specimens; Dr. Ilya Osipov for New Jersey specimens; Jeanne Robinson (Curator of

Entomology, Hunterian Museum) for discussions pertaining to the historical status of proserpina\ Ross

Layberry for information on Ontario bog populations.

REFERENCES

Buckner, J., A. B. Welsh & K. R. Sime. 2014. Evidence for population differentiation in the Bog

Buckmoth of New York State. Northeastern Naturalist 21(4): 506-514.

Crandall, K. A., O. R. P. Bininda-Edmonds, G. M. Mace & R. K. Wayne. 2000. Considering

evolutionary processes in conservation biology. Trends in Ecological Evolution 15:290-295.

Dupuis, J. R., R. S. Peigler, S. M. Geib & D. Rubinoff. 2018. Phylogenomics supports incongruence

between ecological specialization and taxonomy in a charismatic clade of buck moths. Molecular

Ecology 27(22): 4417-4429. https://onlinelibrarv.wilev.com/doi/abs/10. Ill 1/mec. 14883

Ferguson, D. C. 1971. The Moths of America North of Mexico. Fascicle 20.2A, Bombycoidea,

Saturniidae (Part). E. W. Classey Limited and R. B. D. Publications, Inc. London, U. K. 153 pp.

+ n pi.

Fraser, D. J. & L. Bernatchez. 2001. Adaptive evolutionary conservation: towards a unified concept for

defining conservation units. Molecular Ecology 10: 2741-2752.

Gratton, C. 2006. Interactions between a native Silkmoth Hemileuca sp. and an invasive wetland plant,

Lythrum salicaria. Annals of the Entomological Society of America 99(6): 1182-1190.

14

Haggerty, S. A. 2006. Land management implications for Hemileuca maia (Lepidoptera: Saturniidae)

habitat at Manuel F. Correllus State Forest, Martha's Vineyard, Massachusetts. Masters thesis,

University of Massachusetts Amherst. 72 pp.

Handfield, L. 2011. Les Papillons du Quebec. Broquet Inc., Saint-Constant, Quebec, Canada. 672 pp. +

166 pi.

Heppner, J. B. 2003. Lepidoptera of Florida. Arthropods of Florida and Neighboring Land Areas, Vol.

17, Part 1, Introduction and Catalog. Florida Department of Agriculture & Consumer Services,

Gainesville, Florida: x + 670 pp.

Iftner, D. C., J. A. Shuey & J. V. Calhoun. 1992. Butterflies and Skippers of Ohio. Bulletin of the Ohio

Biological Survey 9(1): x + 212 pp.

Kruse, 1998. New Wisconsin records for a Hemileuca (Lepidoptera: Saturniidae) using Menyanthes

trifoliata (Solanales: Menyanthaceae) and Betula pumila (Betulaceae). Great Lakes Entomologist

31(2): 109-112.

Leeuw, I. 1974. A further note on the acceptability of an alternate foodplant for Hemileuca maia (Drury)

(Saturniidae). Journal of the Lepidopterists' Society 28(4): 301.

Legge, J. T., R. Roush, R. Desalle, A. P. Vogler & B. May. 1996. Genetic criteria for establishing

evolutionarily significant units in Cryan's Buckmoth. Conservation Biology 10(1): 85-98.

Martinat, P. J., J. D. Solomon & T. D. Leininger. 1997. Survivorship, development, and fecundity of

Buck Moth (Lepidoptera: Saturniidae) on common tree species in the Gulf Coast urban forest.

Journal of Entomological Science 32(2): 192-203.

Metzler, E. H. & V. P. Lucas. 1990. An endangered moth in Ohio, with notes on other species of special

concern (Lepidoptera: Saturniidae, Sphingidae, Notodontidae and Arctiidae). Ohio Journal of

Science 90:33-40.

Michener, C. D. 1952. The Saturniidae of the Western Hemisphere. Bulletin of the American Museum

of Natural History 98:335-502.

Pavulaan, H. 2020. Designation of neotype of Hemileuca maia (Drury, 1773) and refinement of its type

locality (Bombycoidea, Saturniidae, Hemileucinae). The Taxonomic Report 8(4): 1-12.

Pryor, G. S. 1998. Life history of the Bog Buck Moth (Saturniidae: Hemileuca) in New York State.

Journal of the Lepidopterists' Society 52(2):125-138.

Rubinoff, D., M. San Jose & R. S. Peigler. 2017. Multi-gene phylogeny of the Hemileuca maia complex

(Saturniidae) across North America suggests complex phylogeography and rapid ecological

diversification. Systematic Entomology 42(4): 621-634.

Rubinoff, D. & F. A. H. Sperling. 2004. Mitochondrial DNA sequence, morphology and ecology yield

contrasting conservation implications for two threatened buckmoths ( Hemileuca : Saturniidae).

Biological Conservation 118: 341-351.

Scholtens, B. G. & W. H. Wagner, Jr. 1994. Biology of the genus Hemileuca (Lepidoptera: Saturniidae)

in Michigan. The Great Lakes Entomologist 27(4): 197-207.

Scholtens, B. G. & W. H. Wagner, Jr. 1997. An example of clinal variation in estern North American

Buckmoths (Saturniidae: Hemileuca). Journal of the Lepidopterists' Society 51(1): 47-56.

Selfridge, J. A., D. Parry & G. H. Boettner. 2007. Parasitism of Barrens Buck Moth Hemileuca maia

Drury in early and late successional pine barrens habitats. Journal of the Lepidopterists' Society

61(4): 213-221.

Smith, M. J. 1974. Life history notes on some Hemileuca species (Saturniidae). Journal of the

Lepidopterists' Society 28(2): 142-145.

Tietz, H. M. 1952. The Lepidoptera of Pennsylvania: a Manual. Pennsylvania State College School of

Agriculture, Agricultural Experiment Station, State College, Pennsylvania: xii + 194 pp.

Tietz, H. M. 1972. An Index to the Described Life Histories, Early Stages and Hosts of the

Macrolepidoptera of the Continental United States and Canada, Vol. 1. Allyn Museum of

Entomology, Sarasota, Florida: iv + 536 pp.

15

Tuskes, P. M., J. P. Tuttle & M. M. Collins. 1996. The Wild Silk Moths of North America, a Natural

History of the Saturniidae of the United States and Canada. Cornell University Press, Ithaca, New

York: 250 pp.

Tuttle, J. P., T. W. Carr & M. M. Collins. 2020. An interpretation of the impact of the Wisconsin

Glaciation on the Hemileuca maia/nevadensis complex (Saturniidae) of the Great Lakes Region.

Journal of the Lepidopterists' Society 74(1): 18-30.

ADDITIONAL ONLINE RESOURCES

COSEWIC, 2009. COSEWIC assessment and status report on the Bogbean Buckmoth Hemileuca sp. in

Canada. Committee on the Status of Endangered Wildlife in Canada. Ottawa, Ontario, vii + 29 pp.

https://www.canada.ca/en/environment-climate-change/services/species-risk-pubhc-registry/cosewic-

assessments-status-reports/bogbean-buckmoth-2009.html . Accessed March 14, 2020.

Government of Canada Species Profile. Bogbean Buckmoth. https://wildlife-species.canada.ca/species-

risk-registrv/species/speciesdetails e.cfm?sid= 1058 . Accessed March 14, 2020.

NatureServe Explorer, https://explorer.natureserve.org/ . Accessed March 14, 2020.

New York Natural Heritage Program, 2020. Online Conservation Guide for Inland Barrens Buckmoth

Hemileuca maia maia. https://guides.nvnhp.org/inland-barrens-buckmoth/ . Accessed March 14, 2020.

New York Natural Heritage Program, 2020. Online Conservation Guide for Bogbean Buckmoth

Hemileuca sp. 1. https://guides.nvnhp.org/bogbean-buckmoth/ . Accessed March 14, 2020.

New York Natural Heritage Program, 2020. Online Conservation Guide for Coastal Barrens Buckmoth

Hemileuca maia ssp. 5. https://guides.nvnhp.org/coastal-barrens-buckmoth/ . Accessed March 14, 2020.

Submitted for review March 11, 2020, accepted for publication March 30, 2020.

ADDENDUM

While this paper was in review, an additional study was published (Tuttle et al., 2020) that

discussed the intrusion of a Xerothermic Period "prairie peninsula" (Iftner et al., 1992) across the upper

Midwest and into Ohio. This broad prairie region presented a post-glacial barrier to the movement of

many organisms between the glaciated Great Lakes region to the north and unglaciated regions south of

the prairie. In Ohio, this is well-reflected by the presence of the Great Lakes Complex of Hemileuca

populations north of the prairie peninsula utilizing Salix larval hosts and Hemileuca maia populations

south of the prairie peninsula utilizing Quercus larval hosts. This post-glacial arrangement is maintained

to the present day.

16

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