22 March 2021

The Taxonomic Report

OF THE INTERNATIONAL LEPIDOPTERA SURVEY

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

Evaluation of the taxonomic status of Eurytides marcellus

form “‘floridensis” (W. Holland, 1898)

(Papilionoidea, Papilioninae, Leptocircini)

Harry Pavulaan

606 Hunton Place NE

Leesburg, Virginia, USA, 20176

ABSTRACT. The purpose of this paper is to firmly identify subspecific authorship of the name floridensis for the

Floridian population of Eurytides marcellus (Cramer, 1779), which I recognize as a distinct, though slightly differentiated,

subspecies ranging north and west along the Atlantic and Gulf of Mexico coastal regions. Though the name has been in historical

use by multiple authors since description by William J. Holland (1898), it has not been readily evident which, if any, published

work to date clearly and validly elevated the name to subspecific rank. The name “floridensis” is not preoccupied by any other

members of the butterfly family Papilionidae. | determine the authorship to be floridensis Klots (1951).

Additional key words: Infrasubspecific, International Code of Zoological Nomenclature.

ZooBank registration: urn:lsid:zoobank.org:pub:43AC7666-360C-48 1 F-81C4-1A94F94FDA0C

INTRODUCTION

Pieter Cramer (1779) was the first to illustrate the familiar Zebra Swallowtail (as ‘Papilio

marcellus’) of the eastern United States (Fig. 1). Other than a brief etymology, there was essentially no

descriptive text, nor reference to a type locality other than an elusive reference under ‘Maja’ [Hemileuca

maia| that all the specimens illustrated in plate XC VIII were taken in Virginia. The painted illustration is

clearly of a specimen originating north of Florida by the primary distinguishing character of the forewing

(see below) and appears to match Virginia spring form specimens very closely.

Fig. 1. Papilio marcellus (Cramer, [1779]), in “De Uitlandsche Fig. 2. Papilio ajax, Linnaeus, var. floridensis, Holland

Kapellen...” Plate XCVIII, Fig. F. (1898), in “The Butterfly Book”, Plate XLIV, Fig. 2.

Chainey (2005, page 305) provides an account of the possible disposition of Cramer’s original

specimen: “There is one specimen ex Felder collection but no other data (BMNH(E)#665081) ...As this

species was described from van Lennep’s collection it is possible that this specimen is a syntype.” Note in

Fig. 4 that the label is correctly numbered: 665381. The purported syntype is the only specimen in the

Natural History Museum, London, bearing any data label in relation to any “types” of marcellus. There is

no collection data label or information associated with the specimen. Also, while the specimen represents

the northern population of marcellus, it is a SUMMER phenotype by its elongated tails (Fig. 3). Cramer’s

image (Fig. 1) depicts a SPRING phenotype by its short tails, only tipped with white, and an enlarged red

hindwing anal mark. Thus, the purported syntype is clearly not the specimen illustrated in Cramer (1779),

though representative of the summer form of nominotypical marcellus. [Photos courtesy of Blanca Huertas,

Natural History Museum, London].

Fig. 3. Purported “syntype” of Papilio marcellus in the Fig. 4. View of same specimen showing data labels. Photo

Natural History Museum, London. Photo © by permission © by permission of the Natural History Museum, London.

of the Natural History Museum, London.

“FLORIDENSIS” HOLLAND (1898)

William J. Holland (1848-1932), born on the island of Jamaica, received a theological education

during his childhood in Salem, N.C. through several prestigious colleges, then continuing his education at

Princeton Theological Seminary, graduating in 1874. Holland then became a Presbyterian pastor in

Pittsburgh, PA., later assuming chancellorship of Western University, also in Pittsburgh, and

simultaneously served as chair of Zoology and Comparative Anatomy. MHolland’s studies found him

travelling extensively throughout the U.S. and worldwide. He is known to have traveled to Florida several

times and certainly collected butterflies there. In 1885, he was a founder and first president of the Academy

of Science and Art of Pittsburgh, then in 1901 became director of the Carnegie Museum, until 1922, while

maintaining membership in several entomological societies (Johnson & Brown, 1904). Holland was known

to be an ambitious, even obsessive collector, and considered butterfly collecting as a means to high social

rank among peers (Leach, 2013), thus amassing a sizeable personal collection. Holland is best known for

his landmark “Butterfly Book’, in which he described winter form “floridensis’, first published in 1898,

then reprinted several times until 1951.

W.J. Holland (1898) described the taxon “floridensis” as an infrasubspecific form of Papilio ajax

Linnaeus, as follows:

“Another winter form, for which I propose the name floridensis, is represented in Plate XLIV, Fig. 2, by a male

specimen. It is characterized by the great breath and intensity of the black bands on the upper side of the wings,

which are quite as broad as in the summer form marcellus. I find this form prevalent in the spring of the year on the

St. Johns River, in Florida. Expanse, 2.50-2.75 inches.”

The caption for Plate XLIV, Fig. 2 in Holland describes the illustrated specimen (Fig. 2) as

follows:

“Papilio ajax, Linnaeus, var. floridensis, Holland, 3. (This is the dark form found in Florida in the early spring.)”

Literature treatment of “floridensis” since its original description as either a seasonal

(infrasubspecific) form or as a subspecies, remained inconsistent by various authors for over a century.

Evaluating literature and internet-sourced imagery, specimens in institutional collections, and from personal

field experience, it is evident that the Floridian population of Eurytides marcellus is represented by a

weakly-differentiated subspecies, with either an inland contact zone or cline with nominotypical marcellus

(Fig. 12). [The dynamics of clines are a result of a long period of evolution with many factors at work, and

are frequently more complex than simply zones where one phenotype gradually grades into another. This

is not the focus of the present paper and will not be addressed here.] The question remains over which

author rightfully first assumed authorship of the name in a subspecific capacity. In an attempt to apply

the principles of priority as established by the International Commission on Zoological Nomenclature it is

necessary to review the historic literature. The name floridensis was first assigned subspecific rank by

Klots (1951), as discussed below, though more recent works incorrectly attribute subspecific rank to

Holland (1898).

HISTORICAL SYSTEMATIC TREATMENT FOLLOWING ORIGINAL DESCIPTION

(including major synonymic treatments)

Rothschild & Jordan (1906), in their authoritative ‘A Revision of the American Papilios’,

described, under a summary of “spring forms” (page 689): “P. marcellus f. loc. hib. floridensis Holl.

(1899)” indicating that they also considered “‘floridensis” to be an infrasubspecific local spring form (“‘hib.”

= from hibernation [of pupae]). They describe “the black bands broader than in f. hib. marcellus.”

Grossbeck (1917), lists “Form floridensis” as an infrasubspecific form of Papilio marcellus with

only flight dates and locations.

Barnes & McDunnough (1917), in their synonymic ‘Check List of the Lepidoptera of Boreal

America’, the authors list “floridensis” as an infrasubspecific form of Papilio marcellus.

Seitz (1924) lists, under Papilio marcellus: “forma hib. loc. floridensis Holl. is the spring form from

Florida’, indicating that he considered “floridensis” to be an infrasubspecific local spring form (“hib.” =

from hibernation [of pupae]).

McDunnough (1938), in “Check List of the Lepidoptera of Canada and the United States of

America’, the author again lists “floridensis” as an infrasubspecific form of Papilio marcellus.

Clark & Clark (1951), under Graphium marcellus, give a detailed description indicative of an

intergrade zone into eastern North Carolina: “On the outer Coastal Plain of Virginia the size of the early-

spring form increases considerably, the fore wings being up to 40 mm. in length, though the color remains

the same. Farther southward the dark bands become broader, and finally as broad as in the summer form,

but the tails have only the tips white, and the red anal spot on the hind wings is large and undivided. This

early-spring form with the broad dark bands (floridensis Holland) ranges northward...to eastern North

Carolina, where it intergrades with the normally colored but large early-spring form found along the coast

farther north.” The authors imply infrasubspecific treatment by use of the term “early-spring form’. They

continue: “Early in September 1940 we were surprised to find occasional individuals in the Dismal Swamp

and elsewhere on the Coastal Plain... They resembled closely the southeastern spring form “floridensis”’

and, were the origin and date of capture unknown, would certainly be referred to it.”

W. J. Holland (1951): In the revised edition of ‘The Butterfly Book’, the author briefly describes

form “floridensis” in slightly different wording than the earlier editions but still implies it is

infrasubspecific: “The spring form in Florida is very dark...”

Klots (1951), under Papilio marcellus, treats floridensis as follows, though he does not make a clear

differentiation from nominotypical marcellus: “In central Florida is the weakly distinguished P. m.

floridensis Holland (similar to /econtei) with heavy, dark markings in even the early brood; the

southernmost population of a cline.” This is the first reference I can find, that treats floridensis at

subspecific rank.

Forbes (1960), under Papilio marcellus, states: “Floridensis Holland represents the southern

(chiefly Florida) race.” thus indicating subspecific rank. However, no description of the phenotype is

provided.

dos Passos (1964) lists Graphium marcellus form “floridensis” (Holland), 1898, at infrasubspecific

rank.

Kimball (1965), under Graphium marcellus, merely cites notes by George D. Morgan

(unpublished?): “Of the three subspecies described as differing slightly in size, hairiness, color, pattern, and

length of tails, and supposed to be restricted to certain seasons or regions, all may be matched by

Hillsborough County specimens throughout the year. While it is convenient to follow the line of least

resistance and call all our Florida specimens floridensis (Holland), it is perhaps more accurate to separate

them by color pattern into three series corresponding to marcellus, telamonides (Felder), and floridensis.

The rest will be found to vary in all sorts of ways between these.” Kimball stated also: “...the whole subject

of subspeciation in marcellus needs to be worked out.” Thus, Kimball (1965), following the reasoning of

Morgan, recognizes “floridensis” at infrasubspecific rank.

Mather (1970) gives extensive discussion of seasonal variation in marcellus in Mississippi. Citing

Holland (1931), Mather states: “the form represented as # 2 “floridensis” by Holland...is matched by a few

Mississippi specimens but is not differentiable as a seasonal form in the present sample.” Mather goes on

to conclude that two primary seasonal forms occur in Mississippi: spring (gen. vern. “marcellus’’) and

summer (gen. aest. “lecontei”). This implies infrasubspecific treatment.

Maudsley (1973), in his thesis, studied the influence of temperature, photoperiod and diapause on

producing seasonal polymorphism in marcellus. Maudsley recognized “three distinct forms”: marcellus

(Cramer, 1777), telamonides (C. Felder & R. Felder, 1864) and lecontei (Rothschild & Jordan, 1906),

stating: “All three forms occur throughout the butterfly’s range...Populations in peninsular Florida and

adjacent areas of southern Georgia represent the subspecies G. m. floridensis Holland. G. m. floridensis

differs from the nominate subspecies, which would now be G. m. marcellus, in the extent of black banding,

G. m. floridensis having comparatively wider bands in each form than G. m. marcellus. The subspeciation

is clinal with odd occurrences of floridensis-like specimens in northern G. m. marcellus populations and

vice versa.” Maudsley compares respective “early spring”, “late spring” and “summer” forms in both

subspecies and gives detailed character analyses. He also goes on to note that only two forms occur in

southern Florida: the “late spring” and “summer” forms. Thus, he considers floridensis to be subspecific.

Emmel (in Howe, 1975) states under Graphium marcellus: “The subspecies G. marcellus

floridensis (Holland) is similar to “lecontei” and weakly distinguished; it is found in central Florida.” The

statement clearly applies subspecific rank.

Tyler (1975) lists, under Eurytides marcellus: “A fourth form, “floridensis”...was described as a

subsp. by Holland; its status and relationship to the other forms should be studied.” Tyler thus appears to

imply that he considers “floridensis” to be infrasubspecific.

Miller & Brown (1981), under Eurytides marcellus, list = f. “floridensis’”’, implying infrasubspecific

rank.

Hodges (1983) lists “floridensis (Holl., 1898)” in the synonymy under Eurytides marcellus,

implying infrasubspecific rank.

Gillmore (1988), editor of the Southern Lepidopterists News, for the Zone V report, states: “Ron

Gatrelle commented that...the SC coastal marcellus are identical phenotypically with peninsular Floridian

marcellus! Unknown to many researchers is the University of Georgia PhD dissertation of member Jim

Maudsley of Athens, GA completed in 1970...The most significant aspect of Jim's research was his

recognition of the validity of Eurytides (Graphium) marcellus floridensis (Holland) as our SE subspecies,

whereas most authorities have considered floridensis as a form name...the major difference between

nominate marcellus and floridensis is the consistent enlargement of the dark bands on the upper surface of

the wings as demonstrated by the SE population in all broods. The material from other areas to the north

and west appear almost white by comparison...Clinal gradients between subspecific populations are

commonplace among lepidoptera in the SE Coastal Plain region.” Gillmore’s assessment clearly implies

subspecific recognition.

Gerberg & Arnett (1989) list Eurytides marcellus floridensis at full subspecific rank: “The

only...subspecies in Florida.” The authors give a complete description but fail to differentiate it from

nominotypical marcellus.

Minno & Emmel (1993) list Eurytides marcellus floridensis (Holland) at full subspecific rank but

did not differentiate it from nominotypical marcellus.

Tyler, Brown & Wilson (1994) simply list “floridensis” as a “spring form” under Protographium

marcellus, implying infrasubspecific rank.

Smith, Miller & Miller (1994) provide the following brief assessment under Protesilaus marcellus:

“Populations in Florida were separated as subspecies floridensis Holland, heavily marked and resembling

form ‘lecontei’ even in the earliest adults to emerge...”, thus implying subspecific rank.

Mohn (2002) treats “Neographium (Neographium) marcellus floridensis (Holland, 1898)’ as

follows (specimens are illustrated from Ocala National Forest, Marion Co. and West Palm Beach, Dade

County): “Often viewed as a synonym of m. marcellus. Specimens of the spring generation however, are

markedly darker than those of m. marcellus. Specimens which fly later are darker too. Flight period

substantially longer, from March — December. Distribution: USA, Florida (central Florida, Dade County,

Marion County; southern Florida, West Palm Beach). The author gives clear, descriptive status of the

taxon as the Floridian subspecies and differentiates it from nominotypical marcellus.

Heppner (2003) lists, under Eurytides marcellus - “f. floridensis (Holland, 1898)”, clearly

indicating he considers this an infrasubspecific form.

Lamas (2004) lists “f. floridensis” in the synonymy of Protographium marcellus.

Pelham (2008) lists the name “= floridensis (W. Holland, 1898)” as a synonym of Eurytides

marcellus (Cramer, 1777), thus implying availability as a subspecific name.

ESTABLISHING SUBSPECIES AUTHORSHIP

Holland distinctly described “‘floridensis” as “another winter form’, implying infrasubspecific rank.

All subsequent authors until 1951, maintained “floridensis” as a seasonal form. Klots (1951) was the first

author to apply subspecific rank to floridensis. The pertinent ICZN articles to apply are:

Article 10.2. Availability of infrasubspecific names

An infrasubspecific name is not available [Art. 45.5] from its original publication, unless it was published

before 1961 for a "variety" or "form" and is deemed to be available under Art. 45.6.4.1. If an author uses a

name, previously published at infrasubspecific rank, in a way which makes it available for a species or

subspecies, that author thereby establishes it as a new name and it takes his or her authorship [Art. 45.5.1]

(see also Articles 23.3.4 and 50.3.1).

Article 45.6. Determination of subspecific or infrasubspecific rank of names following a binomen

The rank denoted by a species-group name following a binomen is subspecific, except that 45.6.1. it is

infrasubspecific if its author expressly gave it infrasubspecific rank, or if the content of the work

unambiguously reveals that the name was proposed for an infrasubspecific entity (see also Article 45.6.4);

Article 45.6.4. It is subspecific if first published before 1961 and its author expressly used one of the terms

"variety" or "form" (including use of the terms "var.", "forma", "v." and "f."), unless its author also expressly

gave it infrasubspecific rank, or the content of the work unambiguously reveals that the name was proposed

for an infrasubspecific entity, in which case it is infrasubspecific [see also Art. 45.6.1]; except that

Article 45.6.4.1. a name that is infrasubspecific under Article 45.6.4 is nevertheless deemed to be subspecific

from its original publication if, before 1985, it was either adopted as the valid name of a species or subspecies

or was treated as a senior homonym

Article 50.3. Authorship unaffected by changes in rank or combination

50.3.1. The authorship of the name of a nominal taxon within the family group, genus group or species group

is not affected by the rank at which it is used. But if an infrasubspecific name that otherwise satisfies the

criteria of availability is used in a manner that makes it available for a species or subspecies, its author is the

one who first so uses it [Arts. 10.2, 45.5.1].

The code clearly states that an infrasubspecific name described prior to 1961, which is elevated by

a subsequent author to subspecific rank, the subsequent author is responsible for that usage and takes

authorship of the name. Klots (1951) thus takes authorship of the name floridensis at subspecies rank.

DESCRIPTION OF THE PHENOTYPE

The original description of Papilio ajax f. floridensis was published in Holland (1898). In the

original description, Holland simply indicates: “It is characterized by the great breadth and intensity of the

black bands on the upper side of the wings, which are quite as broad as in the summer form marcellus”’.

Klots (1951) describes it as similar to [summer form] lecontei “with heavy, dark markings in even the early

brood.” Maudsley (1973) describes floridensis in similar terms: “G. m. floridensis differs from the

nominate subspecies...in the extent of black banding...having comparatively wider bands in each form than

G. m. marcellus.”

The original “type” (now lectotype) of Papilio ajax f. “floridensis” (Holland, 1898) is located in the

collection of the Carnegie Museum of Natural History [photo courtesy of Vanessa Verdecia, Carnegie

Museum of Natural History, Pittsburgh]. The specimen’s right forewing had been detached and pinned

beneath the specimen (Fig. 5).

Fig. 5. Holotype specimen, floridensis, dorsal view (left), ventral view (right). Photo © by permission of the Carnegie

Museum of Natural History. [Right forewing (insets) is detached from the pinned specimen.|

Three diagnostic characters of the forewing primarily differentiate subspecies floridensis from

nominotypical marcellus (Fig. 6) and are based on the earliest Floridian spring form in comparison to the

earliest Virginia spring form. These characters apply to all seasonal phenotypes. Differences in hindwing

morphology between the two subspecies are not evident between respective seasonal phenotypes.

OUTER (WHITE)

MEDIAN BAR

) ‘A» BLACK MEDIAN BAR

SUBMARGINAL . fe - \NNER (WHITE)

BAND ~* a “a MEDIAN BAR

MEDIAN BAND

Fig. 6. Diagnostic characters of the forewing used to compare subspecies (the illustrated wing shows

subspecies floridensis).

The width of the median band is narrower in floridensis. The outer edge of the band generally aligns

with the outer edge of the outer (white) median bar. In nominotypical marcellus the outer edge of the

median band extends outward beyond the outer (white) median bar, reaching to about a halfway point up

on the inner edge of the black postmedian bar. Similarly, the inner edge of the median band in floridensis

generally aligns with the inner edge of the inner (white) median bar, while in nominotypical marcellus, it

extends inward.

In floridensis, the black median bar is well-developed, sharply defined and often noticeably

rectangular. In nominotypical marcellus, the bar is variably less well-developed, often forming a slight “v”

shape in the inward side of the discal cell. In many northern individuals, the bar may be faded.

In nominotypical marcellus the submarginal band is essentially straight, with only a slight concavity

in the lowest two wing cells. In floridensis the cells that comprise the band each tend to show well-

developed concavity except in the apical area.

In general, the white areas of nominotypical marcellus are more expansive than in floridensis in all

of the comparative seasonal variants (early spring, late spring, summer). To better visualize and compare

these diagnostic characters, composite images are shown below, with Fig. 7 showing the lectotype

(Floridian spring form) and Fig. 8 showing a typical northern spring form of marcellus. Fig. 9 shows a

typical Floridian summer form and Fig. 10 shows a typical northern summer form.

Fig. 7. Composite image of lectotype. Dorsal (left), ventral Fig. 8. Composite image of early spring form marcellus

(right). February 28 flight date is indicative of earliest annual in Virginia. Dorsal (left), ventral (right). Leesburg, VA.

Floridian brood. Photo © by permission of the Carnegie June 17, 2006. Leg. H. Pavulaan.

Museum of Natural History. Leg. W. J. Holland.

Fig. 9. Composite image of floridensis summer form. Fig. 10. Composite image of Virginia summer form. Dorsal

Dorsal (left), ventral (right). Marineland, FL. August 7, (left), ventral (right). Ex-ova, Leesburg, VA. Em: July 11,

2000. Leg. M. DeGrove. 2015. Leg. H. Pavulaan.

TYPE LOCALITY DATA

The exact locality where the type specimen was collected remains generally unknown. The

specimen data label (Fig. 11) simply reads “Florida” with a collection date of February 28, 1884. Calhoun

(pers. corr.) pointed out that the type label is, in fact, in

Holland’s handwriting. Holland (1898) states: “I find this

form prevalent in the spring of the year on the St. Johns

River, in Florida.” While the description of the location

“St. Johns River’ applies in a general sense to an

approximate type locality, it will have to suffice. As the

River winds over 300 miles from its headwaters in Indian

River County, to the mouth at Jacksonville (Duval County),

Holland’s exact travel routes and collecting site(s) in the

area remain unknown. In 1898, most of the west shore of

the river was accessible between Jacksonville and Palatka

(along what is now Route 17). South of Palatka, there were

few access points along the east side (off today’s Route 17)

until one travelled down as far south as the Deland and

Sanford area. Early rail transportation to the region was a

primary means for travelers, but with limited stops along

this route. However, the entire river lies well within the

range of subspecies floridensis. It would essentially be

pointless to assign an arbitrary type locality without more

detailed information.

[342] CMNH-IZ

aaa 724,189

Fig. 11. Original data labels (CMNH).

DISTRIBUTION, FLIGHT PERIOD AND HOSTS

Distribution: Primarily Florida and adjacent areas

of Georgia; then northward along coastal South Carolina,

where the floridensis phenotype is dominant (Fig. 12). The

floridensis phenotype occurs abundantly throughout

southern Alabama during the summer months, while

nominotypical marcellus and intermediates dominate the

spring brood statewide and northern half of the state in

summer. The floridensis phenotype also ranges west in

rapidly diminishing numbers along the Gulf Coast at least

to southern Mississippi (Mather, 1970), but there are

surprisingly few records of the floridensis phenotype in

Lousiana and Texas, among greater numbers of

nominotypical marcellus.

Review of 6000+ images from many sources

reveals a small percentage of floridensis intermediates and

variants throughout the inland range of marcellus, almost

exclusively as an extreme variant of the summer form,

thus consistent with the observations of Maudsley (1973).

Fig. 12. Blue counties indicate only floridensis present.

Green indicates intermediate forms present. Yellow

indicates only nominotypical marcellus present. Grey

county records indicate no images available.

These intermediates occur more frequently north along the Atlantic Coastal Plain, as far north as Maryland,

where it appears primarily as a late summer variant form among greater numbers of nominotypical

marcellus. 1 have taken a single wild specimen (out of hundreds) as far north as Loudoun Co., VA. on

7/27/2015 that perfectly matches the Floridian phenotype. Reared specimens of marcellus from Loudoun

Co. are normally of the nominotypical phenotype, but a very small percentage (<4%) of summer specimens

appear nearly indistinguishable from Floridian specimens, thus representing a minor variant form in the

nominotypical population. Conversely, a very small number of intermediates occur in northern Florida,

primarily in the spring brood.

Flight period: Multiple broods in Florida. Flight dates span February-December.

Hosts: All known hosts are members of the Annonaceae, mainly Asimina. On the Florida

peninsula, found on Asimina angustifolia (=longifolia), A. incana (=speciosa), A. obovata, A. parviflora, A.

pygmaea, A. reticulata, A. tetramera, Deeringothamnus pulchellus and D. rugelii. On the Florida

panhandle and in areas northward reported to use A. triloba, the sole host of nominotypical marcellus. Other

non-Annonaceae hosts are suggested in Heppner (2003) but require confirmation.

ACKNOWLEDGEMENTS

Thanks go to Crispin Guppy for manuscript review; John Calhoun for helpful comments regarding

the type locality; Blanca Huertas (Natural History Museum, London) for locating and photographing the

marcellus “syntype” specimen; Vanessa Verdecia (Carnegie Museum of Natural History, Pittsburgh) for

locating and photographing the original floridensis “type” specimen.

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