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SOUTHERN LEPIDOPTERISTS’ NEWS VOLUME 44 NO. 4 (2022), PG. 487
BREPHIDIUM EXILIS (LYCAENIDAE) COLONIZES THE
TAMPA BAY AREA AS A NEW FLORIDA STATE RECORD
BY
JOHN V. CALHOUN A ND RON SMITH
The year 2022 saw three bu tterfly taxa recorded in
Florida for the first time. M inno et al. (2 022)
announ ced the d iscovery in the Florida Keys o f A nteos
clorin de (Godart) (wh ite angled-sulphur), und oub tedly
of Cub an origin [som etimes recognized as the
subspecies A. c. nivifera (Fruhstorfer)], and Ma rpesia
eleuchea bah amensis Mu nroe (Antillean daggerw ing)
of the Bahamas. It is su rp rising that these butterflies
had not previously been found in Florida, though the
nominotypical sub sp ecies of M . eleuchea has been
recorded at least three times in the K eys (Czaplak and
Calhoun 2018). Th e third bu tterfly new to Florid a is
much less conspicu ous and cam e as a com plete
surprise.
On 23 July 2022, w hile con ducting a butterfly co unt at
Fort De Soto P ark, Pinellas County, F lorida, several
participants encountered about 2 0 0 pygmy-blue
butterflies in the campgroun d on St. Christop her Key.
Some were photographed by Delia Smith and Ra che l
Sanchez (Fig. 1). R on Sm ith (RS) returned to th e area
on 25 July to photograph more of the butterflies. He
posted one of his im ages (Fig. 2) on iN aturalist an d
initially identified it as Brephidium pseudofea (eastern
pygmy-blue), which is known to occu r in th e park.
Harry Pavulaan, who has studied Brep hidium in
Florida an d elsew here, review ed the image on
iNatu ralist and commented that it was “ a typical
Western Pygmy-Blue by all characters.” Because this
would be a significant new Florida state record an d
ran ge extension, John C alhoun (JC ) requested
ad d ition al im ages fro m R S to c onfi rm the
identification . U pon receip t of the images, th ere was
no dou bt that B rephid iu m exilis (western pygmy-blue)
had reached th e Tamp a Bay a rea o f Florid a, and it
seemed to be established , at least temporarily. The
butterflies were mo rphologically consistent with the
nominotypical subspecies (B. e. exilis) , rather than the
West Indian B. e. iso phth alma, w hich occurs in Cuba
and the Bah am as.
We soo n discovered that B. exilis was present in the
Tamp a Bay area at least three month s earlier. On 29
Ap ril 2022, Liz Childress photograph ed what sh e
believed was a sin gle B . pseudofea nectaring at saw
palmetto (Serenoa repens) near the visitor center at
Weedon Islan d Preserve in southeastern Pin ellas
County. She posted an image (Fig. 3) on iNaturalist
(2022), which JC identified as B. exilis, representing
the first known record of this species in Florid a. This
suggested that B. exilis was more widely distributed in
the area. Additional b reeding populations were
subsequently found, b ut tw o hurrican es an d the
seasonal senescence of food plan ts took a to ll. B y mid-
October, B . exilis seemed to have disap peared from all
but one known locality, which w as still activ e in early
Decemb er.
T
The circumstances surrounding the su dden arrival of
B. exilis in the T amp a Bay area remain a m ystery, and
its con tinu ed p resence is uncertain. It h as possibly
been estab lished in central F lorida for several years,
bu t w ent unn oticed u ntil it colonized F ort De Soto
Park, where bu tterfly counts are routinely conducted.
Winter w eather in the Tampa Bay area has been
relatively mild over the past several years, and it has
not dipped below freezin g in P inellas C oun ty sin ce
Jan u ary 2018 (W eath e r S p a rk 2022). L o c a l
popu latio ns of B. exilis would likely perish if exp osed
to freezing temperatures.
Ad ditional records. On 26 July 202 2, Donald Fraser
fou nd 15 -20 B. exilis at the northern tip of H oneymoon
Island State Park, Pinellas County, alm ost 30 mi (48
km ) north of Fort D e Soto P ark (D. F raser p ers.
Comm.). On 2 A ugust 2022, JC located a sizable
Fig. 1. Fe ma le Bre ph id ium e. exilis, 23 July 202 2, Ft.
De S ot o Park, Pine lla s Co., Florid a (R . Sa nchez).
SOUTHERN LEPIDOPTERISTS’ NEWS VOLUME 44 NO. 4 (2022), PG. 488
colony of B. exilis, consisting of several hun dred
in d i v idu a ls, al ong the s outh east ern sh or e o f
Ho neym oon Island, just east of the entran ce to the
state park (Figs. 4, 14). T he next day, JC found large
numbe rs of th e butterfly at several shoreline spots
along the D unedin C auseway, east of Honeym oon
Island (Fig. 15). In these areas, as well as at Fort De
Soto Park, the butterflies were closely asso ciated with
cre sted saltbu sh (Atriplex pentand ra ) and esp ecially
sea blite (Suaed a linearis) (Figs. 14-17 ). Both of these
plant genera, in the A m arantha ceae (form erly
Ch en opodiaceae), serve as food plants of B. exilis in
oth er parts of its range (Sh apiro 1973, Robinson et al.
2002).
Du ring A ugu st 2 022, we conducted ex haustive
searches of sho reline habitats in P inellas Cou nty, as
well as those in Citrus, Hillsborough, Levy, Manatee,
and Pasco counties. As a result, we docum ente d B .
exilis at fo ur add ition al location s in Pinellas County
and two in no rthern M anatee Cou nty (Figs. 18-21 ).
This inclu des several shoreline sites along th e
Su nshine Skyway Bridge crossing low er Tampa Bay.
As news of the b utterfly’s discovery spread, other
observers recorded B . exilis at additional spots in Fort
De Soto Park. Sp ecimen s collected by JC confirm ed
that these popu lations represen ted the subspecies B. e.
exilis.
Figs. 2-13. Breph idium e. exilis in F lo rida in 2 02 2 (n os. 2 -1 2 from P inellas C o.). 2, ma le , F ort De Soto Pa rk , 2 5 Ju ly
(R . S mith ). 3 , f em ale (?) n ec ta ring o n sa w palmetto , W eedo n Island P re serve, 29 A pril; first kn own F lo rida r ec ord
(L. C hild re ss). 4, fem ale perching, Hone ym oon Isla nd , 2 Aug us t (J. C alh oun ). 5 , female ne ct arin g on h airy pod
cowpe a, D un edin C ausew a y, 8 A ug ust (J. C alhou n). 6, m ale ba sking, B ellea ir C ausew ay, 11 Aug us t (C . Ev ans). 7 ,
fema le necta ring on seaside he lio tr op e, Du nedin Ca us ew a y, 8 A ug ust (J. Calhou n) . 8, fem ale oviposit in g on sea blite,
Ga ndy B ridge, 11 A u gu st ( R. Smith). 9, fema le o vipos it ing on crested saltb ush, H oneymoo n Isla nd State Pa rk, 15
Au gust (R . Sm ith). 10, male pe rc hing , Fort De So to Pa rk, 5 A ug ust (R. Smith ). 11, fem ale b as king , Su nshine Skyw a y
Br idge , 8 Au gust (R. Smith). 12 , fem ale o vipo sitin g on sea blite , S unsh in e Skyw a y Br id ge , 8 A ugust (R . S mith ). 1 3,
ma le baskin g, T er ra C eia, M a natee Co., 2 9 Aug us t (J. C alho un).
SOUTHERN LEPIDOPTERISTS’ NEWS VOLUME 44 NO. 4 (2022), PG. 489
Distribution. Brephidiu m e. exilis is a resident of
low land , arid, and disturbed hab itats from Texas to
Californ ia, south to Venezuela. It is known to
periodically exp an d its range in spring and summer,
reaching as far as W ashington, Wyom in g, Nebraska ,
and Missou ri. Such immigran ts are probably driven
from their permanent breeding areas by stro ng winds
and weather fron ts (waif dispersal) (Pittaway et al.
20 06). T he species was accidentally introduced into
Hawaii in the 1970s and it is now firmly established
there (Riotte and U chid a 197 8, Jamieson and Denny
2001 ). In the 19 90s, B. exilis was inadvertently
introduced into the Persian Gulf (Arabian G ulf)
region, w here it h as since becom e locally comm on,
feeding on both native and introdu ced plants (P ittaw ay
et al. 2006, Pope and N ithyanandan 2014 ).
Brephidiu m exilis was first recorded in coastal
Louisiana in 1 970 (Mather 197 1), an d it has been m ore
frequ en tly rec orded there in recent years (M arks 201 8,
iNatu ralist 2022 ). O n 3 0 July 201 8, a male B . exilis in
good cond ition was photograp hed at Mob ile, Alabam a
(iN aturalist 2 022), but no others were seen at the
locality (C . Stem pien pers. com m.). Although this
species is considered to be a stray in Alab am a (ABA
2022 ), a popu lation was probably presen t in 2018.
Figs. 14-22. Brep hidium e. exilis habitats in Florida in 2022 (all bu t no. 20 from Pinellas Co.). 14, H oneym oon Island,
2 A ugu st (J . C alhoun). 15, Dunedin Causeway, 3 August (J. C a lh ou n). 1 6, abund an t s ea blite, F ort De Soto P ark, 1
Octo ber (R. Smith). 17 , c re sted saltb ush, Fort D e Sot o Par k, 8 August (R. Smith ). 18 , salt flat (note se a blite in
foregrou nd), Belleair C a us ew a y, 4 A ug ust (J. C a lhou n) . 19 , sea b lite pa tc h, G andy Br idge , 4 A ugu st (J. Calhoun ).
20 , sa lt flat ( with abun dan t s mall se a bl it e), Te rr a C eia, M anatee Co., 1 9 A ugu st (J. C alho un). 21, sea b lite along
shoreline , S unsh in e Skyw a y Br idge , 8 A ug ust (J. Calh oun ). 22, Hur rica ne Ia n impa ct to sa me locality sh own in
prev io us im age, 1 0 October (J . Ca lh oun).
SOUTHERN LEPIDOPTERISTS’ NEWS VOLUME 44 NO. 4 (2022), PG. 490
In Florida, JC searched many coastal parks and
shoreline s north to Ced ar K ey (L ev y County) and
sou th to H olmes Beach (Manatee County), but n o
oth er pop ulation s of B. exilis were found, despite the
presence of seemingly suitable habitat in several areas.
RS also explored along the c oast in Taylor County, in
the Big Bend region of Florida, w ithou t success. To
date, there are no kn own Florida records of B. exilis
north of H oneymoon Island (Pinellas Coun ty) or south
of Terra Ceia (Manatee County). The species seemed
to be confin ed to abou t 50 miles (8 0 km) of the west-
central coast (F ig. 23).
Th e apparent lack of populations of B. exilis alon g the
northwestern coast of Florida, and general scarcity of
suitab le habitat there, suggests that th e species did not
gradually spread aroun d the Gulf of Mexico to the
Tamp a B ay area. M ore likely, strong winds associated
with a storm system or weather front from the
northwest conveyed exilis adults across th e northern
Gu lf of Mexico. O ther small species of Lepid optera
are known to travel great distan ces via wind -borne
dispersal (e.g., D antart et al. 20 09). T emporary
populations may regularly be established in th e
sou theast, includ ing Florida, but they are overlooked
due to their localized nature and the inconsp ic uous
size of the adult butterflies.
Habita t. In the Tamp a Bay area, E. exilis was fou nd
only in coastal habitats, particu larly salt flats and
disturbed shorelin es wh ere sea blite and crested
saltbu sh were abundant (Figs. 14-21). Th ey preferred
open p atches of th ese plants that grew in th e direct sun
and w ere not shaded during the day by su rrou nding
veg etation. This was observed in several areas where
the butterflies swarm ed aroun d open patch es of sea
blite, while non e w ere seen just a few feet away where
the p lants w ere partially shaded by man grov es. The
butterflies favored m ore mature plants, 2 -3 feet in
height, though they were also foun d aroun d shorter
plants if they grew abundantly in o pen spaces.
Although sea blite is an ann ual in the temperate z one,
it is a lo ng-lived ann ual or w eak p erennial in the
subtrop ics. It is a pioneer, halophytic species,
inh ab iting a variety of coastal ecosystems that are
seldom flooded by h igh tides (Lonard et al. 2016). In
Florida, sea blite is frequent in salt marshes, strands,
and salt flats of the peninsula, n orth to the central
panhandle. Crested saltbush occurs in similar habitats
along mu ch of coastal Florida (W und erlin et al. 20 17).
These plants often grow directly alon g the shorelin e, at
the u ppe r edge of the wrack zone (Figs. 15-17, 21).
Bigleaf sump weed (Iva fru tescens) sometimes occurs
in the sa me areas and can be mistaken for sea blite,
especially at a distance.
Popu lations of B. exilis were extrem ely localized, yet
the tiny butterflies could be abund ant where found ,
fluttering in small clouds around patch es of sea blite
and crested saltb ush. Males spent m ost of day in
search of females, which were less com m on and
usually kept to th e perip hery of the food plants, where
they ovip osited on leav es and flowering terminal
spikes (Figs. 8, 9 , 12). Both males and females often
perch ed and basked on the food p lants (Figs. 4, 6, 10,
11, 1 3), and visited nearb y flowers (Figs. 5 , 7).
Nectar sources w ere always p resent around colonies of
B. exilis. W e observed adu lts nectaring at shoreline
seapu rslan e (Sesu vium portulacastrum), tu rkey tangle
fo g fru it ( P hyla nodiflo r a ), se a side h e l iotr o p e
(He liotrop iu m cu ra ssavicu m ) (Fig. 7), silverhead
(Blutaparon verm iculare), beggarticks (B id en s alba),
and hairypod cowpea (Vigna luteola) (F ig. 5).
Shoreline seapurslan e is reported as a food plant of B .
exilis elsewh ere (S cott 1986, Warren et al. 2022 ).
Although it grows abundantly in many areas where B.
exilis was found in the Tamp a Bay area, we did not
observe oviposition on this p lant. At one site in late
November and early D ecem ber, males of B. exilis were
seen p atr oll in g fo r f emal es a rou n d sh o rel ine
seapu rslan e, and both sexe s were perching and
nectaring on the plants. This may be a sign that this
plant is fed upo n if sea blite and crested saltbush are
dim inish ed or unavailable. However, so me green sea
blite was still availab le at that site, and no B. exilis
Fig. 2 3. M ap sh ow ing localities of B. e. ex ilis in F lo rida in
20 22. Nam ed coun ties wit ho ut record s are tho se sea rched
un su cc essfully for the species.
SOUTHERN LEPIDOPTERISTS’ NEWS VOLUME 44 NO. 4 (2022), PG. 491
were seen around seap urslane at other localities w here
sea blite an d crested saltbush h ad died b ack. Sh oreline
seapu rslane serves as a food plant of the Hawaiian beet
webw orm (Spolad ea recurvalis) and this m oth can b e
very com mon around patches of the plant. It is easily
mistaken for B. exilis in flight.
Flight period. Brep hidium exilis is a year-round
resident in the southw estern U nited States, w here it is
cap able of producing up to eight or nine broo ds, one
about every three weeks (Shapiro and Manolis 2007,
James and Nunn allee 2011). In southern Louisiana, it
has been recorded all year and is tho ught to produ ce
multiple broods (Marks 2018). The total num ber of
gen erations pro duced in Florida is u n kno wn, but fresh
adults were present from late July throug h early
December, suggesting m ultiple, overlap ping broods. In
September, the number of B. exilis was down
considerably from that observed in A ugust. Despite
searches into early D ecemb er, th e last known record at
Fort De Soto Park was 15 October, when RS and
others photograph ed a few individ uals. It w as no t seen
along the Dunedin Cau seway after 16 October, when
JC counted eight in dividuals arou nd a coup le o f
surviving sea blite plants. No B. exilis were found by
JC anywhere on Hon eymoon Island on 25 Novem ber.
On ly in a small, sheltered salt flat along the Belleair
Causeway in Pinellas C ounty (Fig. 18), where some
green sea b lite p lants remain ed , was B. exilis still
flying in early December.
In central California, B . exilis does not ap pear to
diapause during the win ter. It continues to reprodu ce
in lim ited areas where food plants remain available. In
spring, population levels in crease until adults begin to
disperse into additiona l hab itats (Thacker 2004,
Shapiro an d Manolis 200 7). T he butterfly may employ
a similar strategy to survive the win ter in F lorida,
con tinuing to breed in small numb ers where sea blite
maintains a sufficient amount of green growth. W ith a
burst of new food plant grow th in spring an d sum m er,
brood s of the butterfly may gradually increase in size,
producing hun dred s of ind ividuals that spread to new
areas on coas tal win ds. Numbers peak in late July-
Au gust, only to decline as food plan ts begin to die
back in autum n. Of course, th e p resence of B. exilis in
the Tampa Bay area may prove to be tem porary.
Popu lation sites will continue to b e monitored .
Hu rrican e impa ct. O n 28 September 202 2, Hurricane
Ian made landfall in so uthw est Florida as a strong
Categ ory 4 storm. A lthoug h the highest winds
rem ained south of th e Tampa B ay area, gusts of up to
75 mph (121 kph) were reported as rainb ands passed
over the reg ion. Many of th e p lants along the shore
were damaged, including sea blite growing at the h igh
tide line. Communities of sea blite are p rone to burial
or removal during storm surges (Lonard et al. 2016 ).
Afte r the hurricane, most of the sea blite at so me
localities was blown d own or buried in debris, and
very little remain ed g reen (F igs. 21 , 22 ). On
Ho neymoon Island, where a large colon y of B. exilis
existed along the southeastern shoreline, food p lants
were heavily impacted by the storm and no butterflies
were seen there on 7 October. Along the northe rn side
of the Du nedin Causeway (where winds an d tides had
less imp act), ab out a dozen B . exilis were ob served on
that date. No B. exilis w ere seen at several localities on
10 October, inclu ding alon g the eastern sho res of the
Sunshine Skyway Bridge an d F ort De Soto P ark,
wh ich were most affected by winds that blew un abated
from the east/northeast across the open waters of
Tampa Bay. Th e butterfly was still present in small
numbers on the western sh ore of the park, where
winds were less d irect. W e suspect that adults and
early stages of B. exilis were forcibly extricated fro m
patch es of the plants by strong winds, driving rain, and
tidal inun dation.
Remnants of existing populations w ere possibly also
affected by H urrican e N icole, which passed over th e
Tampa B ay region as a trop ical storm on 10 N ovember
after making landfall on the east coast of Florida as a
Category 1 hu rricane. W inds gusted from the
north /northwest in excess of 5 0 mph (80 kp h) in some
coastal locations, with strong west winds and rain as
the storm continued up the peninsula. This coin cided
with the season al senescen ce of the food p lants,
leaving no living crested saltbush and very little sea
blite with green growth.
Ta xonom ic notes. The perceived relationship b etween
the two N orth Am erican species of pygmy-blues has
shifted over the years, resulting in confusion over th eir
scien tific names. The western pygm y-b lue w as
origin ally described as Lycaena exilis by Boisduval
(1852) based on one male specimen (w hich B oisduval
thou ght was a female) collected in 1850 or 1851 by P.
J. M. Lorquin in the vicinity of San Francisco,
Californ ia (Em m el et al. 1998). Two decades later, the
eastern pygm y-blu e was described as Lyca en a
pseudofea by M orrison (1873 ) based on three
specimens captured in 18 69 at K ey West, F lorida.
Ho wever, S cudder (187 6), believed that pseudofea
was the same species as Lycaena isophthalm a, which
had been described eleven years earlier by Herrich-
Sc häffer (18 62) from specim ens collected in Cu ba by
J. G und lach. Scudder (1 876 ) also placed these species
in their ow n genus, Brephidium . As a resu lt, the
eastern pygmy-blue was then know n as B reph id um
isoph th alma.
SOUTHERN LEPIDOPTERISTS’ NEWS VOLUME 44 NO. 4 (2022), PG. 492
M o rpholo g ical d i ffe r en ces led C o m sto ck an d
Hu ntington (1943) to treat isophthalm a as a West
Ind ian sub sp ecies of B . exilis, while recognizing the
eastern pygm y-blu e as a separate species named B.
pseudofea. Most subsequent authors (e.g., K lots 19 51,
Brown and Heinem an 1972, Riley 1975 , Sm ith et al.
1994 ) agreed with this arrangement. Ho wever,
checklists by dos P assos (1 964) and M iller and Brown
(198 1) treated the eastern pygmy-blue as the
subs pecies B. isophthalma pseudofea. Scott (198 6)
went so far as to conside r the eastern pygmy-blue as a
subspecies of the w estern pygmy-blue, using the
combination B. exilis pseud ofea. This treatment w as
not widely accepted, as there is little evidence to
suggest that they are conspecific (O pler and W arren
2002 ). In fact, genom ic studies b y Zhang et al. (201 9)
supp ort their segregation. M ost au thors now reco gn ize
the eastern and western pygmy-blues as discrete
species (B . pseudofea an d B. exilis, respectively), as
reflected by Pelham (2008, 2022). No netheless, some
recent references (e.g., Glassberg 2017) identify the
eastern pygmy-blue as B. isophthalm a.
In an attemp t to com ply with the rules of zoolog ical
nomenclature, some au thors prefer an alternative
spelling of the nam e exilis. A rticle 31.2 of IC ZN
(1999) dictates that adjectival Latin species names
must agree in gender with the generic names with
wh ich they are associated . In Latin, Brephidium is
neuter, but exilis is feminine. As noted by Ferris
(1989), the nam e exilis should therefore be am ended
to the neuter exile. However, this nomenclatural
provisio n is extrem ely controversial and rou tin ely
ignored. In this case, the Latin exilis is an ad jective
meaning sm all or meag re, which certainly is
appropriate for one of the world’s smallest bu tterflies.
We prefer the original spelling of exilis for historical
reaso ns and because exile is too rem in iscent of th e
English word “exile,” though this also seems fitting
given the species’ propensity for being forcibly b lown
from its home.
The story behind th e n am e pseudofea is also
notew orthy. Several butterfly specimens captured by
G. W. Belfrage near Waco, Texas, were described by
Edward s (1871) as a new species n am ed Lycaena fea.
Edward s n oted that fea is “allied to Exilis . . . of
Califo rnia.” Two years later, Morrison (1873)
described Lycaena p seud ofea , noting that it was
sim ilar to Edwards’ fea. Morrison therefore nam ed his
new butterfly pseudofea because it was a false (i.e.,
pseudo) fea. We n ow know that the specim ens that
Edward s d escribed as fea were actually B. exilis
(Brown 1970), which still occurs around Waco, Texas
(Wauer 2006). The butterfly we know today as
Brephidium pseu dofea is, qu ite literally, a false
Brephidium exilis.
Ackn owled gemen ts. We th ank Brian Ahern , Liz
Ch ildress, Christina Evans, Don ald F raser, John
Lamp kin, Harry Pavulaan, Rachel Sanchez, and Delia
Sm ith for kindly sharing im ages an d information.
Literature Cite d
AB A [ Alab ama B utte rf ly A tlas]. 20 22. W ebp ag e: h ttp s://a labama.butterflyatlas.usf.edu/.
Bo isdu val, J. B . A. D . de. 1 852. Lepido ptères de la C alifo rnie. A nn. Soc. E ntomo l. Fr. (2)10:275-32 4.
Br own , F. M. 1970 . Th e typ es of lyca en id butterflies name d by William Henry Edw ards . Part III. Ple bejinae. With four
neotypes estab lis he d by John C . Do wney. Trans. Amer. En to mol. S oc. 96 -353 -433 .
Br own , F. M ., and B . Hein em an. 19 72. Jam aica and its butterflies. E. W . Classey Ltd., L ondo n. xv+47 8 pp., 10 pls.
Co mstock, W. P., and E . I. Hu ntin gton. 19 43. Lyc aenidae of the Antilles (Lep id opte ra , Rho palocera). Ann . New Yo rk
Ac ad . Sci. 45 :49-13 0.
Cz apla k, D . an d J. V . C a lhou n. 201 8. A ntillean daggerwin g in the F lo rida Keys. S. Le pid. Soc . N ews 40:87.
Da ntart, J., C . S tefa ne scu, A. Avila, an d M . A la rcón . 2 009 . Lon g-distance win d-bo rn e dispersal of the moth Cornifro ns
ulceratalis (L ep id opte ra : C ra mbidae: Ev ergestinae) into the n or thern M editerran ea n. E ur. J. En to mol. 106:225-229.
dos P a ssos, C . F. 196 4. A s yn onymic list of the Nea rc tic R hopalo cera . Lepid. S oc . M em. No . 1. v+ 145 p p.
Ed war ds, W. H. 1 871. De scrip tions of new spe cies of diur na l Lep id op te ra found within the United States. Trans. Amer.
En to mol. S oc . 3:2 05-216.
Em mel, J. F., T. C . E mmel, a n d S. O. M atto on. 1998. The typ es of California bu tte rflies nam ed b y Jean A lp ho nse
Bo isduval: desig nation of lectotyp es and a neo type , an d fixa tion of type localities. Pp. 3-36 . In Emme l, T. C. (ed .) ,
Systema tic s of we stern N o rth A merican B utte rflies. Maripo sa P r., G aine sville, Florida .
Fe rris, C. D . 1 989. Supplem en t to: a catalog ue/checklist of th e butterflie s of Ame rica north of M ex ico. Le pid. So c. M em.
No . 3. vii+103 pp .
Glassberg, J. 2 017. A swift gu id e to butterflies of North Ame rica. Seco nd editio n. 420 pp.
Herric h-Schäffer, G. A . W. 1 862. Schm etter lin ge aus Cuba. Corr es p.-Bl. zo ol.-m in . V er R egenb urg 16:141-143.
ICZN [Inter nation al Commission on Zoo lo gica l Nome nc la tu re]. 1999. Intern atio nal code of zoolog ic al nom en clature.
Fo urth ed ition . The In ternational Trust for Zoo l. Nom en., Londo n, UK. xxix +306 pp .
iNatur alist. 2 022 . Online data base . Webpa ge : https://ww w.ina turalist.org .
SOUTHERN LEPIDOPTERISTS’ NEWS VOLUME 44 NO. 4 (2022), PG. 493
Ja me s, D. G ., and D . Nu nna lle e. 2011. L ife h istorie s of Ca sc adia butte rflies. O regon St. Un iv. P r., C orva llis. 44 7 pp.
Ja mieson , D., and J. De nny. 2001. H a wai’ i’s butterflies and mo th s.: an iden tification guide to easily observe d specie s.
Mutual Publ., Hon olulu, Ha wa ii. xv ii+9 4 pp .
Klots, A. B . 1 951. A field guid e to the b utterflies of N orth A m erica, east o f the G rea t Plain s. H oughto n M ifflin C o., Boston,
Massac husetts. x vi+3 49 p p., 40 pls.
Lo na rd, R . I., F . W . Judd, R . Stalter, a nd J. J . B r own. 20 16. A revie w of the N orth Am erican halo phyte Su aeda linearis
(Ell.) Moq. Pp. 73-84. In Khan, M., B . Bo ër, M. z turk, M. Clü se ne r-G o dt, B. Gul, and S. W. Br ec kle (eds.) , Sab kh a
ecosysems . V o l. V: th e Am ericas. T asks fo r Veg. Sci. 4 8. https://d oi.o rg /10.10 07/9 78-3-3 19-27 093-7_5.
M ar ks, C. 201 8. Bu tte rflies of Louisian a: a gu id e to identificatio n and loc ation. Louisiana St. Univ. Pr., Baton Ro ug e.
viii+46 2 pp .
M at he r, B . 1971 . Annual field season sum ma ry fo r 19 70. Zo ne 6 – southe ast. Lo uisiana. N ew s Le pid. Soc. N o . 3 :4 .
M iller, L . D., an d F. M . Bro w n . 198 1. A ca ta lo gu e/ch ecklist of the butterflie s of America north of M exic o. Lep id . Soc.,
Mem. N o. 2 . vii+2 80 pp.
M inno , M . C., J. R. S lo tten , D. V ollm ar , an d S. K o lterma n. 202 2. Two C aribbe an b utter flies new to F lorida. New s Lepid.
Soc. 64:174-175.
M or rison, H . K . 1873 . Note s on N orth Americ an Lepidop tera. B ull. Bu ffalo Soc. Nat. Sci. 1:186-189.
Op ler, P . A ., and A . D . W a rr en . 2 00 2. Bu tte rflies of Nor th A m erica. 2. Sc ientific na mes list for bu tterfly s pe cies of N orth
Am eric a, north of M exic o. Con trib. C. P . G illette M us. Arth. Div., C olorado St. Un iv. 7 9 pp.
Pelham , J. P. 2 008 . A catalo gue of the butterflie s of the U nited States an d Can ada: with a co m p le te bibliog rap hy of th e
descrip tive an d sy stem atic literature. J. Res. Lep id. 4 0:i-xiv, 1 -658 .
Pelham , J .P . 202 2 . A catalogue of the butterflie s of the United States and Canada. Rev ised 2 Februa ry 2 02 2. Webp ag e:
htt ps: //ww w.b utt erf lies ofa mer ica .co m/U S- Can -C at.h tm .
Pitta w ay, A . R., T . B. L a rsen , A. Leg ra in, J. Ma jer, Z. Weide nh offer, and M . Gillet. 2006. Th e establishm en t of an
Am erican butterfly in the A ra bian G ulf: Brephidium exilis (B oisd uval, 1 852) (Lycaenidae). Nota Lepid. 29 :5 -16.
Po pe, M ., and M . N ithy ana nd an. 2014 . R ec ord of the western pygmy blue butterfly Bre ph id iu m exilis (B o isduval, 1 852)
(Lepidoptera: Lycaenid ae ) fro m Sa bah Al Ahm ad S ea C ity, K uwa it. J. Thr ea tened Ta xa . 6:6723-6725.
Riley , N. D . 1 975 . A fie ld g uide to the butter flies of the West Ind ies. De mete r Pr, B osto n, M assachusetts. 22 4 pp ., 24 pls.
Riotte, J. C . E., a nd G . Uc hida . 19 78. Bu tterflie s of the H awaiian Islands. J. R es. Lepid. 17 :3 3-39 .
Ro bins on, G. S ., P. R . Ackery, I. J. K itchin g, G . W . B ecc alo ni, and L . M . Herná ndez. 2002. Ho stplan ts of the m oth and
bu tte rfly ca terpillars o f Am erica no rth o f Mexico . Mem . A mer. E ntomol. Inst. 69:1 -8 24.
Scott, J. A . 1 986 . Th e butte rflies of North Ame rica: a na tu ra l h istory and field g uide. S ta nford U niv. Pr., Stanfor d,
Ca lifornia . xiii+58 3 pp ., 64 p ls.
Scud de r, S. H . 187 6. Syno nymic list of the butterflie s of N orth A m eric a, nor th o f Mexic o. Part II. Rura les. Bull. B uffalo
So c. Nat. S ci. 3 :98-12 9
Shap iro, A. M . 1973 . Host records fo r Brep hidium exilis (Lycaenid ae ). J. Lepid. S oc. 27 :1 57 -1 58.
Shap iro, A . M ., and T. D. M an olis. 200 7. Field guide to butterflies of the San Francisc o Bay and Sacramento Valley
regio ns. Un iv. C alifo rnia Pr., Berkeley and Lo s Ange les. xiv+34 3 pp., 31 pls.
Smith, D . S., L . D . M iller, and J. Y. M iller. 1994. T he bu tterflie s of th e W est Indies an d So uth F lo rida. Oxfo rd Univ . P r.,
Ne w York. x+26 4 pp ., 32 p ls.
Th ac ke r, P. D . 20 04 . California bu tte rflies: at ho me w ith alie ns. BioS cience 54:182-187.
W arr en, A . D., K. J. D avis, E. M . S ta nge land , J. P . Pelh am, K. R. Willmo tt , and N. V . Grishin. 202 2. Butterflies of
Am erica. W eb page: http ://www.butterfliesofam er ica.c om/.
Wau er , R. H . 2006. Fin ding b utterflies in T ex as: a gu id e to th e best sites. Johnson B o ok s, Bou lder, C olor ad o. viii+3 27 p p.
Wea th er Spa rk . 2 022 . Intera ctive w ea ther da ta. W ebpage: https://we atherspark.co m/.
Wun de rlin, R. P., B. F. Ha nsen , an d A. R. Franc k. 2017. F lora of Florida. Volume IV . Dicotyledon s, Com br etaceae
thr ou gh A mara nthace ae. U niv. Pr. Florida, Gain esville. 400 p p.
Zh ang , J., Q. Con g, J . Shen, P .A . Opler, and N .V . Grishin. 2 019. Genom ics of a com plete butte rfly contin en t. bioR xiv
829887.
(John V. Calhoun; E-mail: bretcal1@verizon.net )
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