Butterflies — Made in America

A new paper in Nature Ecology and Evolution suggests that butterflies evolved in what is now the Americas, where they originally fed on plants in the legume family.

The research team, led by Akito Kawahara at the University of Florida’s McGuire Center for Lepidoptera and Biodiversity, sequenced 391 genes from nearly 2,300 butterfly species, sampled from 90 countries and 28 specimen collections, to reconstruct a new phylogenomic tree of butterflies representing 92% of all genera. 

Kawahara et al focused on three main questions: (1) did butterflies originate in the northern (Laurasia) or southern (Gondwana) hemisphere; (2) what plants did the ancestor of butterflies feed on; and (3) are host repertoires (that is, diets) of butterfly species and clades constrained by host phylogeny?

In sum, they write, “Our data support the hypothesis that butterflies originated in the Americas in the late Cretaceous, 100 million years after the origin of angiosperms, and that they first fed on legumes. Butterflies dispersed from the Americas to the Eastern Palaearctic across Beringia ~75 Ma before diversifying in the Palaeotropics. Although our analyses point to a Nearctic origin, evidence for a North American versus a Central American origin is not strong, and we therefore tentatively conclude that a Laurasian origin is likely. Larval host plants played an important part in the evolution of butterflies, and some groups became host specific whereas others retained a wide host breadth.”

Butterfly global colonization following origin in Americas

Posted in evolution, general butterfly news, taxonomy | Leave a comment

Carolinas Butterfly Monitoring Program Kicks Off

I’m reproducing here a note (originally from the CarolinaLeps listserv) about a terrific new initiative headed up by Jeff Pippen to systematically survey the butterflies of the Carolinas using standardized methodology and investing in the ID and ecological skills of participants. This is an effort sorely needed not just for the Carolinas, and I hope the program is successful enough to spawn copycats across the US! And I hope our Carolinian readers will participate if you haven’t already signed up! And best of luck, Jeff.


I’m delighted to officially announce the CarBMP – Carolinas Butterfly
Monitoring Program, a citizen-science based program developed for you to be
able to regularly walk a route, record the butterflies you see, and contribute
to a national database to monitor butterfly populations.

Several volunteers from across the state are already participating.
Ultimately, our goal is to have hundreds of observers across the Carolinas
conducting hundreds of surveys every week or two, in order to get a big
picture of our butterfly populations and how they are faring with changes in
climate, land use, human population, etc.

To that end, we will be conducting workshops on butterfly identification and
how to set up and monitor a route in the coming months.  I’ll post another
announcement soon for the first few workshops, which are in the Raleigh-Durham
area starting next week.  Details for a Charlotte area workshop in early
August are also coming together.

From the CarBMP website
(https://urldefense.com/v3/__https://www.jeffpippen.com/butterflies/carolinasbmp/carolinasbmp.htm__;!!OToaGQ!soPPRhGTBYsy-Tg59P876L9Qxm-TGsCo8A1_PhjxAIYdknizDboOFzteVnaZYOZBMjcyWIIaFiHwnDAOZCTgfZI$ ):
Brief Description

The Carolinas Butterfly Monitoring Program (CarBMP) aims to track butterfly
populations using scientifically sound, standardized survey methods as well as
opportunistic sightings across both North and South Carolina. This Citizen
Science based program partners with non-profit and government organizations,
offers public outreach, encourages interaction between the general public and
professional biologists, and provides data to researchers evaluating the
health of habitats and insect populations in the Carolinas.

Our Mission

The Mission of the Carolinas Butterfly Monitoring Program is to:
Develop and train a team of professional and citizen scientists to conduct
regular surveys on permanent routes across both North and South Carolina
Develop a long-term picture of butterfly population trends while connecting
people to the outdoors

Partner with conservation and educational organizations to increase awareness
and enable synergistic efforts for protecting insect populations and natural

Foster public interest in and awareness of pollinators and natural habitats
and the importance of monitoring insect populations by providing education
outreach tools such as workshops and identification guides
Provide data to researchers evaluating the health of habitats and insect
populations and to assist with creating potential solutions to reversing or
slowing negative trends
A Little Background

The potential dire consequences of declining insect populations across the
globe have become a focal issue in both scientific circles and the general
public. This recent increased attention has not only highlighted the
consequences of losing the ecosystem services insects provide, such as
pollination, but has also revealed a dearth of rigorous, long term population
studies. Despite rising concern among scientists, the scientific evidence for
global and even regional declines is still weak; many of the key studies are
based on results with very limited spatial and temporal replication, so it is
difficult to support conclusions. In order to better document how widespread
these declines may be, we critically need more scientifically produced data on
insect populations. Establishing biodiversity monitoring programs like the
CarBMP is a key to understanding the future of natural resources, and this is
especially true in the Carolinas where rapid population growth and urban
expansion are accelerating at a very high rate. We welcome you to join the
CarBMP, enjoy some time outside, and become a part of our growing citizen
science program.

Good Butterflying!


Jeffrey S. Pippen
Mebane, NC & Georgetown University
Director, Carolinas Butterfly Monitoring Program
>   |   N. Am. Butterfly Monitoring Network
>   |   Jeff’s Nature Pages

Posted in checklists, Field Trips/Annual Counts, general butterfly news, sightings, state butterflies | Leave a comment

And the winner is ….!

The winner of the 2023 Falcate Foray is Barry Marts!

Barry submitted an identifiable photo of a male Falcate Orangetip from Green Ridge State Forest a few weeks ago, and this week followed up with a perched female Falcate to complete the set. Congratulations!

The contest, as you may recall, was to be the first observer to photodocument BOTH a male and a female Falcate in 2023. Barry was sorting through various small whites flying in GRSF for Olympia Marbles, without much success, so we hope this is a nice consolation prize.

As winner of the challenge, Barry will receive a copy of of Jessica Speart’s “Winged Obsession: The Pursuit of the World’s Most Notorious Butterfly Smuggler.”

Posted in general butterfly news, maryland, sightings | 1 Comment

In Defense of Plants: Butterflies in a Changing World

This week’s drop of a new podcast from In Defense of Plants (a terrific weekly listen if you don’t already know it) features University of Arizona’s Katy Prudic (an eButterfly co-director) as she discusses work she has been doing with community scientists surveying butterflies and managing butterfly habitats in urban areas, especially in the Southwest.

Listen to the podcast here.

You can also visit her lab and see what kinds of work she has been doing.

Posted in climate change, conservation, endangered species, evolution, general butterfly news | Leave a comment

A Tale of Two (or More) Crescents

2018 May 25 northern crescent 1 in situ_MD-Indian Springs WMA-Indian Springs Pond copy

Field or photo ID for the non-tharos crescents is only accurate for the males as the females of all our species in MD are indistinguishable in the field except by association with known-species males. [this one photographed 2018 May 25, Indian Springs WMA, so likely one of the bivoltine cocyta-group crescents if it sports an antennal club that is fully orange underneath — this one did, see below]

Oh for the simple days when in the mid-Atlantic region if you saw a crescent you could safely assume it was the common, indeed almost ubiquitous, Pearl Crescent (Phyciodes tharos).  The only doubt to contend with would be if you saw windows in the black spots on the trailing edge of the hindwing to make it a Silvery Checkerspot.

But no, we had to go poking around looking at flight behavior, timing of broods, and minute differences in color of the antennal clubs and go confusing ourselves with multiple crescent possibilities.

So what are the crescents we can expect in the area?

Of course Pearl Crescent is still the go-to orange-and-black small butterfly of choice for most of Maryland most of the time.  Like many crescents, the wing pattern is highly variable, as is the amount of orange on the antenna.  Many older accounts say there is one continuous, rolling flight of Pearl Crescent from early spring until frost, and in fact you can find crescents in the field almost anytime during the summer season. Many good authorities still think all our crescents are Pearls, Phyciodes tharos.

Harry Pavulaan begs to differ.

I asked him recently to give us the lay of the land, crescent-wise, for the MD-VA-WV region and help us figure out what to look for in the field in 2023.  He and colleagues have been working on the complicated relationships of the, um, complex (in every sense of the word). Here’s his capsule summary of what we know — and don’t know — about our crescents.

Harry writes…

Our present understanding, via published literature, is that there are two active species of Phyciodes present in this region:  P. tharos and P. cocyta (often known by other names).  P. batesii is certainly not present.  To the best of our knowledge, P. cocyta consists of two species in our region: a univoltine taxon (P. cocyta) that flies in the higher mountains in late June to early July; and a bivoltine species that flies in the mountain region as far east as the Frederick Municipal Watershed, in May and again in August.  This has not been published, but is the subject of considerable field observation, collection and rearing, and is now being analyzed by Nick Grishin’s genome research team at Univ. of Texas, Southwestern Medical Center. 

Distinguishing these three species is a considerable challenge.  The females are indistinguishable because females of all three species are identical in morphology and their antenna clubs all vary from black to orange.  In specimen series, tharos females are smallest, the bivoltine cocyta females are slightly larger, and univoltine cocyta females are the largest.  Photos won’t do any good and are only identifiable to genus (Phyciodes), though one can get them in the ballpark by flight date.  Females are best identified by association with males within population concentrations.

The males, on the other hand, can be distinguished only by a single character: the lower (ventral) side of the antenna club.  P. tharos males have black antenna clubs with some gray on the lower side.  Some P. tharos males have a slight orange tip on the upperside of the club, but the underside of the club is not orange.  P. tharos often lives in concentrations around aster hostplants and fly in April, July and September, though their flights can vary somewhat from year to year.

The males of the two P. cocyta species (univoltine and bivoltine) are indistinguishable except by size and flight date.  The male antenna clubs of both are orange beneath.  The orange may also appear on the upperside of the clubs, but extends across the entire lower side.

Several field guides suggest that P. tharos and P. cocyta (as “Northern Crescent”) can be differentiated by the extent of orange on the dorsal hindwing.  I found this to be unreliable.  While univoltine P. cocyta does, in all cases,  have reduced black postmedian maculations and a broader orange field on the hindwing, individuals of both P. tharos and bivoltine P. cocyta vary considerably in this character.  In fact, individuals of bivoltine P. cocyta generally have fully-developed hindwing maculations and appear identical to P. tharos.  Yet, some P. tharos have reduced hindwing maculations, showing an extensive field of orange.  Thus, one cannot rely on this character.

Another, also somewhat unreliable way to distinguish the species is by habitat.  P. tharos is essentially an open field species, using white aster species as their larval hostplants.  Bivoltine P. cocyta is essentially a forest species and has not been found in open habitats.  At George Thompson WMA in Fauquier County, VA, for example, P. tharos is common around Lake Thompson, on the dam and along the open trail on the north side.  They are showing wing wear at the time when bivoltine P. cocyta emerges inside the forest.  Bivoltine P. cocyta can be seen along the north side of the lake along the forest edge and likely use the woodland aster that grows along the uphill trail.  The two species segregate themselves.  Univoltine P. cocyta is more of a mystery.  I have been looking for these for many years from western Maryland to Bath County, VA and over on the WV side of Reddish Knob – without luck with but one sole exception, a female taken at Sleepy Creek WMA in Berkeley County, WV that laid eggs and produced giant adults, compared to the other two species.

The relationship of the two P. cocyta species should resolve itself in coming months or a year or two.

The notion that there are species or subspecies hidden in our Maryland Phyciodes complex is one that we as diligent field observers can do a lot to elucidate.  Take careful photographs, making sure you have a male and can clearly see the underside of the antenna.  Net them if you need to. And make even more careful notes about the date seen, the behavior, the habitat, and the size of the population you encounter.  And include that data when you post your picture to Flickr, or iNat, or eButterfly, or the Maryland Biodiversity Project.

If you see something, say something!

2018 May 25_MD-Indian Springs WMA_Indian Springs Pond copy

Here’s the money shot you need — straight on, looking at the full underside of the antennal club on a male Phyciodes crescent. You don’t see many views like these on iNat [2018MAY25, Indian Springs WMA, photo by REB]

Posted in conservation, evolution, general butterfly news, Identification tips, maryland, sightings, state butterflies, taxonomy | Tagged , | 1 Comment

Low-hanging Fruit for Maryland Lep Observers

A White M Hairstreak like this documented in Somerset Co would close out the list for Maryland counties where it is known to occur [2013, Linkwood WMA, Dorchester Co — close but no cigar! Photo by REB]

With the data from the Maryland Biodiversity Project in hand that I posted about recently, I dug a little deeper to see where a diligent butterfly observer might make some easy dents in our understanding the distribution of Maryland butterflies.

While there are still quads within the counties lacking records, these butterflies species have now been documented in every Maryland county:

  • Least Skipper
  • Sachem
  • Common Checkered-skipper
  • Silver-spotted Skipper
  • Horace’s Duskywing
  • Dun Skipper
  • Fiery Skipper
  • Zabulon Skipper
  • Common Sootywing
  • Peck’s Skipper
  • Little Glassywing
  • Red-banded Hairstreak
  • Summer Azure
  • Eastern Tailed-blue
  • Gray Hairstreak
  • Common Wood-nymph
  • Monarch
  • Variegated Fritillary
  • Common Buckeye
  • American Snout
  • Viceroy
  • Red-spotted Purple
  • Mourning Cloak
  • Pearl Crescent
  • Red Admiral
  • Painted Lady
  • American Lady
  • Black Swallowtail
  • Eastern Tiger Swallowtail
  • Spicebush Swallowtail
  • Sleepy Orange
  • Orange Sulphur
  • Clouded Sulphur
  • Cabbage White
  • Pipevine Swallowtail

But there are nearly a dozen butterflies for which only one county remains undocumented for that species. These are mostly common butterflies, so a little field work could easily mean we could polish off this dozen with quick dispatch! Clearly Baltimore City, with its largely urban landscape, will be the most challenging. We could do a butterfly blitz or two and end the day at Inner Harbor over crab cakes.

  • Eastern Comma (Somerset)
  • Zebra Swallowtail (Somerset)
  • Cloudless Sulphur (Allegany)
  • Tawny-edged Skipper (Caroline)
  • Hayhurst’s Scallopwing (Garrett)
  • White-M Hairstreak (Somerset)
  • American Copper (Baltimore City)
  • Hackberry Emperor (Baltimore City)
  • Falcate Orangetip (Baltimore City)
  • Little Yellow (Baltimore City)

I left Spring Azure off this one-county list. Technically, according to MBP, this species has been documented from every county in the state except Baltimore City. However, many of these records were established before we came to understand that the earliest azure in the spring for us is actually Summer Azure. It used to be thought that Summer Azure flew only later in the spring, so many of these MBP records for Spring Azure are almost certainly in error. In addition to that, the two are very hard to separate from online photos, and I am regularly reminding azure posters in the April timeframe on iNaturalist that their photo can’t be confidently identified to species. We probably should establish a new baseline for azure records that specifically checks the male upperside wing coloration (the diagnostic character) to figure out where Spring Azure is found. Summer Azure is on the list of species that has been found in all Maryland counties, and that is likely accurate. Spring Azure is almost certainly much more restricted in range, and even that range seems to be declining.

Only two species are missing just two county ticks, and both should be easy to score with a little summer fieldwork: Wild Indigo Duskywing (no records for Queen Anne or Somerset) and Clouded Skipper (absent from Queen Anne and Garrett).

The first thing to do would be to go back to your existing records and make sure you don’t already have documentation for a species in a location that hasn’t yet been recorded. I’m guessing some of us do. Submit your records to MBP or iNaturalist and log that checkmark now!

If not, you have your marching orders for 2023. Remember that you don’t need to be an ace photographer to contribute content to either MBP or iNaturalist (which also feeds into MBP). They just need to be identifiable; you can even net them and photograph them in hand. Let’s see if we can’t close out at least half of these in the next season.

Posted in checklist, checklists, general butterfly news, maryland, sightings, state butterflies, taxonomy | Leave a comment

2022 Was a Good Year — For New Butterfly Records

2022 was a good year for butterfly observations, even if it wasn’t a very good one for this Cabbage White snagged by an ambush bug in my back yard this summer.

It appears that 2022 was a pretty good year for folks in MD prospecting for new county records. Thanks to Bill Hubick at the Maryland Biodiversity Project, we now know that 2022 gave Maryland lepsters three new county records, five new month records (observations for a month where there previously had been no sightings), and a whopping 635 new quad records!

The county records are:

Henry’s Elfin (Callophrys henrici) recorded by fm5050 via iNaturalist was new for Baltimore Co. (View)

Long-tailed Skipper (Urbanus proteus) recorded by marciarose via iNaturalist was new for Somerset Co. (View)

Zebra Swallowtail (Eurytides marcellus) recorded by Stephanie via iNaturalist was new for Baltimore City (View)

We’re very close to closing out counties for Zebra Swallowtail — only Somerset remains as a Maryland county where Zebra Swallowtail has not officially been reported. Three county ticks remain for Henry’s Elfin, and five counties remain for Long-tailed Skipper.

We should be able to make short work of these in the 2023 season!!

MBP also allows us to see new data records, although it’s hard to know what to make of them since it isn’t always clear if they represent new early or late dates without combing the data more thoroughly. I suspect that our phenology data still reflect more observation effort than they do actual early/late flight dates, but the dataset at MBP is getting so rich that eventually we will probably be able to tease out this information as well. But here they are in the meantime:

Bog Copper (Lycaena epixanthe) recorded by Josh Emm was new for MBP for June. (View)

Cabbage White (Pieris rapae) recorded by cwgoldy via iNaturalist was new for MBP for January. (View)

Cherry Gall Azure (Celastrina serotina) recorded by Josh Emm was new for MBP for May. (View)

Dion Skipper (Euphyes dion) recorded by rinwolfe via iNaturalist was new for MBP for August. (View)

Northern Azure (Celastrina lucia) recorded by rborchelt via iNaturalist was new for MBP for April. (View)

Because the list of new quad records is so long, I won’t post it here. But you can find it here to peruse to your heart’s content — and make a plan to do some quadbusting with me in 2023.

As always, we are indebted to the Maryland Biodiversity Project, which has replaced the laborious hand curating of observations each year undertaken by the late Dick Smith. MBP is now the default authority on butterfly phenology and occurrence in Maryland, but it’s only as good as the data *you* contribute. Unfortunately, the tendency is for contributors to populate MBP with photo-documented observations, and that can skew the dataset toward large, photogenic, or unusual species, not to mention butterflies on the wing in the most clement months of the summer! Plus, now that MBP ingests quality records from iNat — which is also biased toward photo documentation — we as a community need to careful not to conflate reported sightings with actual abundance or presence in the field.

It also means we can do more to make MBP, iNat, and other similar sites more robust — like documenting common species, and documenting butterflies across their lifecycles, across their geographic range, and across their flight seasons. Scientifically, those Cabbage White observations are just as valuable as that Great Purple Hairstreak you saw! So let’s take more opportunities to document *all* our species — in MD and elsewhere — in addition to the cool ones. That’s a good 2023 New Year lep resolution.

Posted in Calendar, general butterfly news, maryland, sightings, state butterflies | 1 Comment

Past Pleasures — Noel Humphrey’s British Butterflies

The still-gloomy end of winter, with its short days and cold temperatures, means we can spend time perusing some of the 19th-century lep-works without fear of missing out on what’s happening in the field.

With this dispensation in mind, I’ve been paging through the 1859 edition of H. Noel Humphreys’ British Butterflies, whose lavish illustrations of butterflies and their habitats set a new standard in “realistic” portrayal of butterflies in situ.

The hand-colored lithograph plates are stunning, and there is also a wealth of life history info on many of the butterflies illustrated. The subtitle tells what a treat you’re in for:

The Genera and Species of British Butterflies, described and arranged according to the System now adopted in the British Museum … illustrated by Plates in which all the Species and Varieties are represented, accompanied by their respective Caterpillars, and the Plants on which they feed.

Here’s a terrific plate of Peacocks and Camberwell Beauties (better known here in the States as Mourning Cloak) as an example of the rich biological detail the plates include:

You could spend several hundred dollars to obtain a copy for your bookshelf, or you could read it for free here in the LepLog library, courtesy of the Biodiversity Heritage Library:

Posted in European butterflies, general butterfly news, Identification tips, Review | 1 Comment

Urban Life for Leps is Driving Their Evolution

The Green-veined White, Pieris napi [Wikimedia Commons]

Much work has been done on the effects of urbanization, and especially light and sounds, on bird fauna.  In general, the effects have been fairly striking, including more singing at “night” (because there is artificial lighting) and changes to song to make them more audible in a very competitive urban soundscape.  Some work has also been on nocturnal insects, including some moths, focused especially on the deleterious impact of artificial lighting on finding mates. 

Comparatively little is known, however, about the impacts of urbanization on the phenology of Lepidoptera.  A recent (2021) paper in PNAS suggests a range of changes in the biology of some common leps in response to urban environments.  These changes even appear to be driving evolutionary pathways.

The researchers studied two common leps in Scandinavia, the Green-veined White (Pieris napi), and the Latticed Heath (Chiasmia clathrata, a geometer moth), and their response to urbanization.  The butterfly is a multivoltine species that flies throughout the season; the moth is normally bivoltine but long-flying. Both are common and regularly show up in citizen science surveys; the geometer flies by day as well as by night. 

From lab experiments on the leps’ photoperiodism and phenology, the team predicted that both would emerge earlier in the season and fly later, in response to both artificially induced longer daylight early in spring and late in the summer and to warmer ambient temperatures in urban areas as compared with rural locations.  In the field, these predictions held true. For P. napi for example this averaged out to urban whites flying some five days later in the season than their rural counterparts. Warmer urban sites also allow for faster development rates and longer periods of growth and reproduction than their cooler rural surroundings, the researchers noted, so the five-day difference is likely even more important than simply a longer time to fly.  The difference probably also is enough to drive evolutionary changes in flight period and development.

The research “shows typically longer flight periods for both study species that end later in the season in urban areas versus their rural surroundings, suggestive of a partial extra generation,” the authors write.  “From an evolutionary perspective, it is this increased voltinism that is the most important response, since an extra generation may accelerate population growth and facilitate further adaptation. Nonetheless, producing an extra generation at high latitudes also entails risks; entire cohorts may perish if winter arrives before individuals reach a stage that allows successful overwintering.”

“Altered seasonality of urban environments can lead to corresponding evolutionary changes in the seasonal responses of urban populations, a pattern that may be repeated in other species,” they conclude.

Read the full paper below in the LepLog library.

Posted in climate change, conservation, European butterflies, evolution, general butterfly news | Tagged | Leave a comment

New Year, New Challenge

[Updated to clarify sightings must be AFTER January 1, 2023. Cut-and-paste is the nightmare of editors]

It’s barely six weeks now in the mid-Atlantic until we can be confident of seeing early spring butterflies, so it must be time to launch our own East coast version of Art Shapiro’s Cabbage White Contest.

In the past we’ve had a challenge here at LepLog that recognized first butterfly, first azure, first Mourning Cloak, first anglewing trio, and other contests. This year the challenge is the Falcate Foray!

The Falcate Foray will reward the first set of BOTH male and female Falcate Orangetip. Rules are as usual for this contest:

1.      All butterflies must be live adults seen in the wild (i.e., not from chrysalids brought indoors to emerge).  Netted adults in hand are acceptable.

2.     All butterflies must be seen in Maryland, Delaware, DC or the Northern Virginia counties of Loudon, Fairfax, or Prince William.

3.     All butterflies must be photodocumented and submitted as a set to rborchelt@gmail.com.  This isn’t a photo contest, photos only need to be clear enough for positive ID.

4.     The order in which they are seen/documented doesn’t matter.

5.     All observations must be made after January 1, 2023.  The first complete set submitted (as certified by the time stamp on the email) will be declared the winner. 

The 2023 Falcate Foray winner will receive a copy of Jessica Speart’s “Winged Obsession: The Pursuit of the World’s Most Notorious Butterfly Smuggler.”

So, cameras to the ready! Males reliably take wing in our area the last week of March; females a week to 10 days later.

Posted in general butterfly news, sightings | Tagged , | 3 Comments