Photuris walldoxeyi Faust, 2019

Faust, Lynn Frierson & Davis, Jeff, 2019, A New Species of Photuris Dejean (Coleoptera: Lampyridae) from a Mississippi Cypress Swamp, with Notes on its Behavior, The Coleopterists Bulletin 73 (1), pp. 97-113 : 100-110

Photuris walldoxeyi Faust , new species

Zoobank.org/ urn:lsid:zoobank.org:act:A7F5CEF5-6828-4196-AD25-979F8E35ED3E ( Figs. 1–3 )

Type Material. HOLOTYPE labeled as “ Holotype P. walldoxeyi ³. USA, Mississippi, Marshall Co., Wall Doxey State Park, Spring Lake , 34.664781°N, 89.465717°W at 114 m, 14–15. V.2017. Netted while flying and flashing in cypress. L Faust. ” ALLOTYPE labeled as “ Allotype P. walldoxeyi ♀, USA, Mississippi, Marshall Co., Wall Doxey State Park, Spring Lake , 34.664781°N, 89.465717° W at 114 m, 28.V.2018. Perched on swamp vegetation with courting paratype. L. Faust .” Holotype ( Figs. 1A, B, F , 2A ) and allotype ( Fig. 1D, E ) are deposited in the US National Museum of Natural History, Washington, DC . PARATYPES: 5 ³ labeled with same data as holotype except as “2 ³ paratype P. walldoxeyi , collected 14–15 May 2017 ” and “3 ³ paratypes collected 16 and 28 May 2018 ” at the type locality, L. Faust . Paratypes ( Figs. 1G, H , 2B–F , 3 ) are deposited in the Mississippi Entomological Museum, Starkville, MS (2 ³, 14–15.V.2017), University of Georgia, Stanger-Hall Laboratory , Athens, GA (1 DNA voucher ³, 16.V.2018), Florida State Collection of Arthropods , Gainesville, FL (1 ³, 28.V.2018), and Lynn Faust Private Collection , Knoxville, TN (1 ³, 28.V.2018). All specimens are preserved in 95% EtOH and kept in a freezer .

Description. Male: Length 11.9–12.6 mm; width 4.4–5.0 mm at middle of elytra, 3.1–4.0 mm at base of elytra. Head: Frons and lower vertex pale yellow ( Fig. 1F ) with a small, brown, triangular to oblong mark medially on upper vertex, extending under pronotum. Ventral keel a thin, dark line, contouring the eye margin of gena ( Fig. 3B ). Maximum distance of keel to eye margin greater than least distance of gular margins. Clypeus connate to frons, pale to brown at the curved frontoclypeal suture. Anterior margin tridentate and dark brown ( Fig. 1F ). Mandibles amber basally, heavily sclerotized and dark brown apically; maxillary palpi dark brown laterally, pale medially; labial palpi sickle-shaped, dark brown ( Fig. 3C ). Antennal length 7.4 mm, 0.61X body length ( Fig. 3A ); antennal sockets pale yellow ( Fig. 3C ); scape pale to dark brown, palest at base, pedicel brown, pale basally, barely shorter than antennomere 3, antennomeres 4–11 brown with pale base and each twice as long as antennomeres 2 or 3 ( Fig. 3A ). Pronotum: Length 2.6–2.9 mm, posterior width 2.8–3.2 mm ( Figs. 1A , 2D, E , 3A ). Cream-colored, dark brown vitta with narrowed or broken stem just posterior to apex, widening gradually to base, width of which varies 0.16–0.33X pronotal width ( Lloyd 2018 vittagram terms), vitta bordered laterally by reddish orange in rounded or angular spot. Scutellum: Color entirely pale (holotype) to variably brown anteriorly, pale posteriorly ( Fig. 2A, C, E ). Mesonotal plates with dark brown margins medially and posteriorly; remainder of plates subtriangular, pale to brown. Elytra: Length 8.7–9.8 mm, width 2.2–2.6 mm at widest point near middle ( Figs. 1A , 2D, F , 3A ). Dark brown to black, evenly colored, with dense pubescence covering elytra, becoming less pubescent at humeri; elytral vittae absent; lateral margins yellow, widest at midlength, distinct at apices and confluent with yellow sutural margins that are continuous from apices to scutellum, somewhat narrower than yellow on lateral margins. Epipleura with dusky brown, longitudinal, basal patch, otherwise variably pale. Thorax: Mesosternal and metasternal sternites shiny dark brown ( Figs. 1B , 3A ), typical of P. versicolor species-group; metanotal tergites mottled pale and gray. Abdomen: Tergites 1–5 gray-brown, 6–8 yellow ( Fig. 1B , 2B , 3D ). Sternites 2–4 brown, sternite 5 crossed by pale band posteriorly, arched towards middle, with brown lateral margins or 3–5 rating of splash ( Lloyd 2018); sternites 6 and 7 with typical P. versicolo r species-group lanterns, with 6 more broadly emarginated than 7; sternite 8 pale yellow, with equilateral triangular cauda enclosing pale aedeagus. Legs: Color intensity variable depending on darker or paler specimen ( Figs. 1B , 3A ). Procoxae bicolored, generally pale with diffuse gray-brown lateral surface. Mesocoxae pale medially, darker laterally, slightly darker than procoxae. Metacoxae dark brown; Trochanters pale yellow. Profemur with basal half pale, apical half dark brown. Meso- and metafemora pale with dark brown band on apical fourth and apical third, respectively. Articulations of femur and tibia pale. Tibia pale basally with dark coloration and dark setae on apical 3/4; 2 amber meso- and metatibial spurs, one larger than other. Tarsus less dark than tibia; paler basally with dark setae and dark apical joint articulations. Hind legs 0.75X body length. Aedeagus: Length 2.2 mm from phallobase to lateral lobe tip; 0.7 mm at widest midway ( Figs. 1C , 2B ). Filaments extending 0.2–0.3 mm past tips of lateral lobes (parameres). Ventro-basal lateral lobe processes truncate, 0.12 mm long. Typical P. versicolor species-group form (Barber and McDermott 1951).

Female Allotype. Length 11.1 mm; width 3.9 mm at middle of elytra, 3.1 mm at base of elytra. Length slightly less than all male paratypes but overall appearance similar. Alate like male. Pronotum: Length 2.7 mm, posterior width 2.7 mm. Vitta stem constricted ( Fig. 1D ). Head: As in male, except smaller eyes typical of Photuris females. Robust, tridentate dark clypeus. Abdomen: Sternite 5 brown, lacking any central pale splashing as in males. Sternite 6 and 7 with typical P. versicolor species-group female lanterns but anterior lantern almost twice width of posterior lantern ( Fig. 1E ). Both lanterns flying saucer-shaped. Sternites surrounding lanterns paler centrally, brown laterally. Sternite 8 pale centrally with brown mottling laterally.

Flash-Train 1 Glow. Flashing began 45 minutes after sunset at 8:40 CDT on 14 May 2017 with the highest numbers of males flying 9:00–10:45 pm but continuing later in lesser numbers. The primary flash of a courting male at 17–19° C consisted of an average of 5.5 green-yellow pulses of bright intensity (range 4–9) (n = 15 FT+Gs with 82 pulses), each pulse ca. 200 milliseconds in duration and given at ca. 0.4 second intervals in a flash-train lasting ca. three seconds followed (332 milliseconds after cessation of final pulse) by a 1-second (range 0.7–1.2) glow of medium intensity. The entire FT+ G lasted 3–5 seconds and was given in a syncopated rhythm of fast, bright flashes followed by a slow diffuse glow. FT+Gs with more pulses lasted longer than those FT+Gs that had fewer pulses ( Figs. 4 , 5 ). Hundreds of these emissions being given by 50+ males were observed during 14–27 May 2017 at Wall Doxey State Park.

In 2018, with six weeks of observations in three different states, temperatures varied greatly from the 2017 season, illustrating how temperature affects the rate and speed of firefly flashes. At 17–19° C, the average FT+G lasted 3.3 seconds (SD = 1.111, n = 16). At 23° C, FT+G duration decreased to 2.75 sec (SD = 0.401, n = 9). During the warmest evening, 14 May 2018 when temperatures ranged 25–28°C, the average FT+G lasted 1.86 seconds (SD = 0.425, n = 45). At these high temperatures, discreet pulses within the FT+G were difficult to accurately count with the human eye when compared to the relatively cool 2017 emissions that had easy-todistinguish individual components. As Wolf River Conservancy’ s Cathy Justis (personal communication to first author) reported in 2018 after a warm night (>25° C), there was “kind of a strobe effect –4–9 flashes– followed by a longish ascending glow.” Because of the natural drop in temperature after sunset, particularly on clear nights, FT+Gs could go from this rapid strobe-like appearance as darkness falls to easily countable pulses by midnight.

Each FT+G was completed in a threedimensional yet restricted area in vertical and horizontal space ( Fig. 4 ). The male hovered almost stationary within an imaginary aerial box or vertical column <0.5 m. Males moved more during the final single glow than the entire initial multipulse flashtrain. The glow could be described as a “flourish or swoosh” as it was just as likely to be an arc or J-shape as a wavering sinuous line. Males displayed each individual flash-train in their own concise area, not cross-flying other displaying males. The observer’ s camera position and the angle of flight and changing, sinuous aerial positions of the firefly influence the apparent perspective of the spacing of the FT+Gs in the photographs ( Figs. 4C, D , 5 ). Fig. 4B and E show less looping emissions and more clearly illustrate the typical predictable and even spacing of the pulses and terminal glow.

A secondary “extra 2 pulse + Glow” emission ( Fig. 5 ) often occurred 0.7–1.7 seconds after the primary FT+G, especially during the time of peak numbers of courting males and peak flight times (ca. 9:00–11:00 CDT). These “2 pulse + Gs” were identical to the primary flash emissions except only two bright pulses followed by the glow occur in the secondary emissions that lasted 1.1–1.8 seconds at 17–23° C. Both FT+Gs, primary and secondary, were given as one combined continuous showy emission by the male. When the primary emission consisted of five or fewer flashes, these secondary emissions were likely to occur. When the primary emission consisted of 7–9 flashes, the secondary “2 pulse + G” emission was usually not given. These short “2 pulse + Gs” were never given in isolation. They were only seen immediately following a primary emission ( Fig. 5 ). Other than the identical though shorter “2 pulse + Glow” pattern added at times of high competition to the primary FT+G, P. walldoxeyi does not appear to have any additional patterns, unlike many of the P. versicolor speciesgroup members that often have several different male courtship flashes (Barber and McDermott 1951; Faust 2017; Lloyd 2018).

Males displayed from less than a meter above the water up to the lower and midway limbs of the cypress and tupelo. At most sites, the majority of males flew <6 m aboveground. One site at Cypress Swamp in Mississippi has enormous trees with few lower branches remaining. Here the males did fly higher but still remained out of the uppermost crowns of the large trees. At an old slough, 100 m distant from the swamp proper, with shorter vegetation, males flashed low over the water and around the lower branches. Whether intentionally or not, these low flying males achieved the doubling effect of their flash trains by having the water reflect the FT+Gs back up into space. When a secondary “2 pulse +G” was added to the primary FT+G by a low flying male, he was much more noticeable than any high-flying males because of these additional apparently reflected flashes.

Because of the high numbers of courting males gathered in a relatively small area, it was difficult to follow an individual for the timing of its repeat FT+ G. During the dark, non-flashing period, males quickly moved horizontally some distance and were lost in the dark and among other displaying males before repeating their next train ca. 7–12 seconds later.

As the number of males decreased by season’ s end 4–5 weeks after the first males appeared, there were long periods of no flashing followed by a few seconds of <10 males giving one or two FT+G emissions in loose concert. This was followed by more darkness instead of the near constant display evident during peak times.

Etymology. Photuris walldoxeyi is named for both the type locality and the Mississippi Senator for whom Wall Doxey State Park is named. Wall Doxey, 1892–1962, was a lawyer politician from nearby Holly Springs, Mississippi, who championed forest preservation throughout his career as both Congressman and later US Senator representing northern Mississippi. The common name cypress firefly has already been in use among trained observers and local conservation and wildlife agencies in three different states prior to publication of this description. We also propose Wall Doxey’ s firefly as a more formal common name since other firefly species are seen in the cypress areas and surrounding forest.

Diagnosis. The presence of reddish orange spots bordering the pronotal vitta places this species in the P. versicolor species-group in the key by McDermott (1967). Its small-medium size (for Photuris ) and absence of elytral vittae further separates this species from many in this group. The described flash train of an average of five to six evenly spaced pulses followed by a 1-second glow (at 17° C) with and entire emission lasting 3–5 seconds is unique among North American Photuris and easily recognizable in the wild. According to flash pattern categories defined by Lloyd (2018), P. walldoxeyi falls into the “HH” group that is characterized by more complex combinations of different elements, e.g., pulses in flash trains combined with sustained glows, as opposed to the more typical single pulses, flickers and simple flash-trains. Though P. walldoxeyi , P. barberi , and P. forresti occasionally emit flash trains, especially at temperatures>23° C, that can appear similar to the human observer depending on aspect and flight path of the firefly and position of the observer, each species retains a predictable and unique flash pattern of its own, different from the other two species. Photuris walldoxeyi emits a primary FT+G often followed immediately by a secondary (2 pulse + G) flash with up to nine pulses in a looping three-dimensional compact space and always ending with a glow ( Figs. 4 , 5 ). Photuris forresti emits a series of pulses and more prolonged flashglows in varying order also in a looping manner and compact 3D space. Photuris barberi gives an “electric wink” ( Lloyd 2018) with fewer pulses in a more horizontal manner. Differences in habitat, elevation, manner of flight, and display are detailed below.

The recently proposed species P. barberi described by Lloyd (2018) has the most similar Photuris flash pattern to that of P. walldoxeyi . Lloyd’ s single voucher was caught “where they flew at the top of tall pines” with a “twenty foot tropical net” on 3 June 1994 near Steinhagen Lake in East Texas in dense second growth habitat of low elevation (27 m). Photuris walldoxeyi has only been found in inundated or swampy cypress or tupelo habitat at seven sites. It has not been seen flying in second growth thickets along woodland trails. Photuris barberi displays are described as high in the air around the treetops, “where they flew at the top of tall pines” ( Lloyd 2018), whereas P. walldoxeyi often displays low over the water and at trees’ midlevel at highest. The “HH” flash pattern of P. barberi has a different, more irregular rhythm than P. walldoxeyi ’s very predictable pattern. In P. barberi , the initial few pulses within the flash-train are regularly spaced, after which the rhythm changes to a shorter duration final pulse immediately followed by a more prolonged dash lasting 0.5 seconds at 22° C. Lloyd calls this feature “an electric wink”, since the final shorter pulse in the train is contiguous to the terminal dash. The entire emission of P. barberi has few average pulses (1–5), including the “electric wink” pulse ( Lloyd 2018), than P. walldoxeyi ’s 4–9 pulses. Compare, also, P. barberi ’s changing pulse rhythm and abrupt horizontal flight and “dash” to P. walldoxeyi ’s hovering, evenly spaced pulses and sinuous, diffuse, less abrupt terminal glow lacking the “electric wink” of P. barberi .

Photuris barberi does not display the secondary “extra 2 pulse + Glow” that P. walldoxeyi frequently displays ( Fig. 5 ). The distinctly sinuous, looping, and always hovering nature of the FT+Gs of P. walldoxeyi were not mentioned by Lloyd (2018) in his brief description of P. barberi . He instead noted the linear manner of flight and emissions of P. barberi that also maintained a constant intensity, whereas P. walldoxey ’s glow emissions appear diffuse and less intense than the flash-train pulses.

Although there is only one specimen of P. barberi (current deposition uncertain), true abdominal segment five of P. barberi has a larger central area of pale coloration, the “splash” ( Lloyd 2018), than all the known P. walldoxeyi specimens. According to Lloyd’ s abdominal segment pattern code, P. barberi has abdominal segment five rated a 7 in coloration, having a larger pale area reaching the height of the segment, whereas P. walldoxeyi , in all specimens, are rated 3–5 on Lloyd’ s scale, having a smaller pale central area. In habitat, elevation, splash pattern, flash pattern, flight manner, and height and appearance of display, P. walldoxeyi and P. barberi differ.

A second newly described species, P. forresti (Faust 2017 (as “ Photuris F or Loopy 5”); Lloyd 2018), is similar to P. walldoxeyi in its manner of displaying an erratic, looping flash-train in a welldefined aerial vertical column over higher elevation (277–320 m) in swampy habitats in eastern Tennessee and northwestern South Carolina that lack cypress. Instead of the evenly spaced, predictable pulses of P. walldoxeyi that are always followed by a diffuse glow, emissions by P. forresti are remarkably modulated with combinations of pulses and short glows that occur in varying order in each flashtrain. In 2018, professional photographer Radim Schreiber took over 1,000 high quality photographs of P. forresti ’s more unpredictable flash-trains, thereby documenting and enabling more detailed analysis of what the first author has observed at both sites for the past seven years (Faust 2017 and unpublished data). We would suggest, however, that P. forresti belongs in the “HH” flash group ( Lloyd 2018) because of its erratic pulses and glow flashtrains and not in the “DD” group as Lloyd (2018) states. Photuris forresti lacks any secondary “extra pulse + glow” that is so characteristic of P. walldoxeyi . The first author has deposited DNA vouchers of three male P. forresti , collected 6 June 2018 in Jefferson County, TN at Eslinger Swamp, at the University of Georgia in the Stanger-Hall Lab for future genetic studies with P. walldoxeyi . The first author currently retains six P. forresti specimens from 2012–2018, male and female, in her private collection. In habitat, elevation, flash pattern arrangement and appearance, P. walldoxeyi and P. forresti differ.

Courtship, Lifespan, and Habits. Two courtships were observed by both authors 28 May and 5 June 2018 at two different sites. The female perched about one m above the ground on vegetation growing in the water yet within easy flying distance of drier land for eventual oviposition. She gave erratic, quick single, doublet, and triplet flashes. The courting male landed on swamp vegetation 0.5–1.0 m away. The male switched from his courtship flash to single flashes as he approached her, and both continued a private back and forth dialogue. This 28 May male and female pair and a male collected 16 May were kept alive for captive studies until 9 June. The males were never observed approaching the female for courtship when placed together in a mesh cage. The female never made predatory motions to either male, though their time together was closely monitored to prevent predation of either paratype.

When the first author had to preserve them while they were still alive on 9 June, each was a minimum of 12–26 days old, suggesting the typical approximate 3–5-week lifespan typical of many fireflies (Faust 2017). The captive female flashed nightly with both periodic and erratic multi-peaked flashes after capture but never oviposited. At the type locality, predatory females of another P. versicolor species-group member, much larger and with strong elytral vittae, were captured flying and signaling from the ground with erratic or three-pulsed flashtrains within the displaying P. walldoxeyi male’ s arena each year. Larvae of typical Photuris form were found on forest trails and near the lakeshore in small numbers. Because other Photuris species were present at Wall Doxey State Park throughout the year, these larvae were not collected. No larvae were seen submerged in the water of the cypress areas of Spring Lake itself. At all sites, newly eclosed adult P. walldoxeyi could easily fly out from slightly drier, higher land nearby to the cypress areas to display.

Type Locality Habitat. Northern Mississippi falls in the humid subtropical climate zone like much of the rest of the southern USA. Photuris walldoxeyi was found displaying in the bald cypress areas of 24-hectare Spring Lake. Wall Doxey State Park ( Fig. 6A ), established in 1938 and with 363 hectares (Land Conservation Assistance Network, 2017, www.landcan.org/local-resources/Wall-DoxeyState-Park/19962/) at 114 m elevation, consists of pine and upland second growth hardwood forests and some man-made open grassy areas surrounding the spring fed lake, dammed>150 years ago at the southern end. The lower lying cypress areas (Mississippi Museum of Natural Science 2015) are located closer to the source of the major spring, the outflow and multiple smaller springs that feed Spring Lake . All observed males displayed in and around the branch tips of the bald cypress from just above the surface of the lake to up to about six m above the water’ s surface. At times of highest display, a few males flew nearer the shore in and around shoreline cypress and Virginia sweetspire shrubs ( Itea virginica L., Iteaceae ) and hazel alder ( Alnus serrulata (Aiton) Willdenow , Betulaceae ) overhanging the water of the swamp. This flash pattern was not observed away from the cypress area and surrounding shoreline, although other species of fireflies were observed elsewhere in the pine and upland type forest and open areas surrounding the lake. The bald cypresses appeared to be maturing second growth and were of medium size, averaging 25–40 cm diameter at breast height (as measured 50 cm above the butt swell). This entire area was heavily impacted by humans in the past 175 years but is now recovering since establishment of the state park. Photuris walldoxeyi was found around the shore of Spring Lake wherever there were cypress stands, but the fireflies were absent elsewhere.

Additional Collection Sites. The 2018 season focused on searching for additional sites and elaborating on our 2017 studies. Ten observers in three states spent 40 nights searching for new populations of P. walldoxeyi in suitable tupelo-cypress habitat during these studies. Six additional major sites were located that stretch 608 km in a north-south axis from central Mississippi to southern Indiana ( Figs. 6 , 7 ) and average 105 m elevation. All seven sites in Mississippi, Tennessee, and Indiana where P. walldoxeyi was observed share similar characteristics. All are in cypress or tupelo-cypress swamp areas. All share a narrow band of elevation where they occurred at 95–115 m elevation. Except for the Indiana site, located in the Interior River Lowland ecoregion III (Omernik 1987; US Environmental Protection Agency 2018), five sites are located in the Mississippi Valley Loess Plains , ecoregion level III, which runs in a narrow south-north track from northern Louisiana to the Ohio River in western Kentucky. The type locality closely borders the Mississippi Valley Loess Plains (by four km) but technically lies within the Southeastern Plains. No populations have yet been found in the lower elevation Delta Mississippi (Mississippi Alluvial Plain ecoregion III) to the west, despite several nights of looking in promising sites in Warren County, Mississippi. Nor has the species been seen in the cypress-tupelo swamp areas of the Noxubee watershed (in Southeastern Plains ecoregion III) to the east. Future searches will continue.

With the exception of one site thus far, all P. walldoxeyi sites remain inundated year-round instead of experiencing seasonal flooding and dry-out. Other plants commonly noted at the sites were hazel alder, Virginia sweetspire, lizard’ s tail ( Saururus cernuus L., Saururaceae ), yellow pond-lily ( Nuphar advena (Aiton) W.T. Aiton , Nymphaeaceae ), arrowhead ( Sagittaria L., Alismataceae ), swamp rose ( Rosa palustris Marsh , Rosaceae ), arrow arums ( Peltandra virginia (L.) Schott, Araceae ), fetterbush ( Lyona lucida (Lam.) Koch , Ericaceae ), buttonbush ( Cephalanthus occidentalis L., Rubiaceae ), and tussoch sedge ( Carex stricta Lamarck , Cyperaceae ). Many sites that appeared to have all the proper attributes yet differed in elevation or other yet unknown variables lacked P. walldoxeyi , while nearby locations with near identical habitat did have this species. Moving south to north, P. walldoxeyi is confirmed to be present at the following sites:

· along Mississippi Natchez, Madison Trace,32.5795166 County, ° Cypress N, 89.870016 Swamp ° W at 96 m elevation .

· Park Mississippi, 34.664781, Marshall °N, 89.465717 County, Wall ° W Doxey at 114 State m elevation ( Fig. 6A ).

· water Mississippi River, Desoto , Old Hwy County 78, Bridge , 34.908172 over Cold-° N, 89.751070° W at 106 m elevation.

· Hwy Mississippi 267 Bridge , Marshall. Enormous County population, Coldwater stretching River on over a mile. 34.937117° N, 89.641560° W at 99 m elevation .

· Ghost Tennessee River, Fayette State Natural County Area , Mineral , 35.026719 Slough ° N in, 89.272287° W at 115 m elevation ( Fig. 6B ).

· Tennessee Conservation, Fayette Area on County Wolf River , William, 35.055036 B. Clark ° N, 89.540112° W at 95 m elevation.

· Indiana Preserve,, Posey 37.828215 County ° N,, 87.983554 Twin Swamps ° W at Nature 112 m elevation ( Fig. 6C ).

Timing, Seasonality, and Degree Days. Male courtship flashing began 35–45 minutes after sunset (7:54 PM) at about 8:30–8:50 PM, with maximum numbers displaying at 9:00–10:30 PM (65–155 minutes after sunset) followed by gradually reduced numbers continuing until pre-dawn hours. On peak nights, males began flashing as much as 10 minutes earlier than on nights with low competition.

In calendar days, P. walldoxeyi was observed from 12 May 2018 at Wall Doxey State Park, MS to 17 June at Twin Swamps, IN where they were still displaying in moderate numbers the final time observations were made. The most southern site, Cypress Swamp in Mississippi, north of Jackson, most likely had an earlier start date because of its latitude.

Using the “modified corn growing degree day or 86/50 Fahrenheit” (mGDD) with 1 March start date formula used in prior studies (Faust and Weston 2009; Faust 2010, 2012, 2017; Faust and Forrest 2017), P. walldoxeyi was not yet active at ca. 589 mGDD on 5 May 2018 at the type locality, yet had appeared in low to moderate numbers by 12 May at 739 mGGD and were still present in declining numbers 3 June at 1,270 mGDD. It was displaying in low numbers through 1,503 mGDD at the farthest southern site at Cypress Swamp on 30 May 2018. The Indiana site ( Fig. 6C , Fig. 7 ) at Twin Swamps had the latest observed display date of 17 June 17 at 1,460 mGDD. Judging from the lifespan of 3–5 weeks and seasonality of most adult Photuris species (Faust 2017) , males of P. walldoxeyi most likely would begin to emerge as adults in late April or early May in the southernmost sites near Jackson, Mississippi and display until mid- to late June as far north as Indiana during the range 700–1,550 mGDD. The highest numbers representing peak flight occurred in about 900–1,200 mGDD at all sites in 2018 and 2018.

For northern Mississippi, the first appearance of P. walldoxeyi in early May represents the earlier phase of the predictable seasonal progression of the local firefly species assemblage. Pyractomena borealis (Randall) (L. Faust, personal observation, 2004; T. Mann, personal communication, 2002–2018; K. Allred, personal communication, 2007–2011; P. Hartfield, personal communication, 2018) and Phausis inaccensa LeConte (Faust and Forrest 2017) in very early spring precede P. walldoxeyi ’s emergence in central to northern Mississippi. Photuris versicolor var. quadrifulgens Barber has a similar flight period as P. walldoxeyi in nearby open fields (Forrest and Eubanks 1995). Photuris walldoxeyi completes its life cycle earlier than the peak flights of slightly later emerging Photuris frontalis LeConte , Photinus pyralis (Linnaeus) , Photinus cooki Green (C. Barksdale, personal communication, 2011; J. Davis, personal observation, 2013; E. and P. Hartfield, personal communication, 2014–2017) and other late spring and summer firefly species in Mississippi.

Associated Fauna. In this brief snapshot of time in May 2017 and 2018, at least six other Lampyridae were active in low numbers at Wall Doxey State Park during the surveys: Lucidota atra (Olivier) , Lucidota punctata LeConte , Photinus macdermotti Lloyd , P. pyralis , P. frontalis , and Photuris tremulans Barber. It is known that additional firefly species are present during different times of the year in selected habitats within 40 km of Wall Doxey State Park and may likely be present in this state park (Forrest and Eubanks 1995; Faust and Forrest 2017).

Figures

Fig. 1. Photuris walldoxeyi. A) Holotype, dorsal view (all specimens lack pale elytral vittae), B) Holotype, ventral view, with splash, C) Paratype aedeagus (truncate basal lateral lobe projections enhanced to show details), D) Allotype female pronotal pattern with constricted stem, E) Allotype female lantern with no splash, F) Holotype, tridentate clypeus (left) with paratype variations from dark to pale. Scale bars = 1 mm. Image C by Luiz da Silveira.

Fig. 2. Photuris walldoxeyi. A) Holotype, scutellum and brown-edged, proximate mesonotal plates, B) Paratype, lantern detail, C and E) Scutellar color variation with postero-medial, brown-edged mesonotal plates, D and F) Pronotal vittagram variations (stem constricted or interrupted) Scale bars = 1 mm.

Fig. 3. Photuris walldoxeyi, male paratype. A) Habitus (pronotal pattern with wider base yet still constricted stem), B) Ventral keel (arrows), C) Head, frontal view. Scale bar = 1 mm. Photographs by Luiz da Silveira.

Fig. 4. Photuris walldoxeyi male flash-train+glow (FT+G) pattern at 17° C. A) Diagram illustrating 4–9 quick, bright pulses of the flash-train followed by a prolonged lower intensity glow, B–E) FT+G patterns in loops and arcs in a small aerial space. Photographs by Jeff Davis.

Fig. 5. Diagram showing primary flash-train+glow and secondary “2 pulse + G” rhythm and pattern, shown in straight line for clarity. Flash trains, primary + secondary, as they appear connected in space. Photographs by Jeff Davis.

Fig. 6. Habitats of Photuris walldoxeyi. A) Type locality at Wall Doxey State Park in northern Mississippi, B) Mineral Slough in Ghost River Natural Area in western Tennessee, C) Twin Swamps Nature Preserve in Indiana. Image C by Max Henschen.

Fig. 7. Distribution of known populations of Photuris walldoxeyi (stars) at ca. 105 m elevation, Photuris forresti (circles) at ca. 300 m elevation, and Photuris barberi (triangle) at ca. 27 m elevation.