Phausis reticulata ( Say, 1825 ), 1852

Hodson, Alicia M. & Branham, Marc A., 2024, Revision and Phylogeny of the Genus Phausis (Coleoptera: Lampyridae) with the Description of Three New Species, Zootaxa 5458 (1), pp. 1-52 : 32-35

Phausis reticulata ( Say, 1825)

(Male: Figs 13 ; 18C ; 20C ; 22Y –Aa; 24I. Female: Fig. 21A–F )

Lampyris reticulata Say 1825: 163 .

Phausis reticulata (Say) : LeConte 1852: 337.

Lamprohiza reticulata (Say) : Wentzel 1896: 294.

Diagnosis. Male. Side margins of pronotum more arcuate and less strongly convergent before the rounded anterior margin compared to Lamprohiza splendidula . Scutellum spatulate with posterior apices emarginated with a medial longitudinal groove. Elytra with single prominent costa, longitudinal and transverse lines. Surface strongly reticulated ( Fig. 18C ), but not as defined as in Phausis luminosa or Phausis christineae . Aedeagus with medial lobe slender ovate dorsally, and lateral line present ( Fig. 22Z ). Parameres with dorsal proximal flanges present, restricted medially ( Fig. 22Y ). Flanges are extensions of parameres that curve or extend outward from inner margin. Females are paedomorphic and apterous. Differ from female Lamprohiza by the absence of anterior vitreous pronotal spots and elytra.

Redescription. Male ( Figs. 18C ; 20C ; 22Y –Aa). Length 5.0–9.0 mm. Width 1.0– 3.2 mm. Color. Body dark brown; eyes black; mouthparts (excluding mandibles) brown-yellow; mandibles brunneous; antennae yellow to light brown; central pronotal disc brownish-black; scutellum pale brown to flavus; thorax overall light brown, with margins and upper portions pale yellow; coxae, trochanters, femora, and tibiae brunneous; tarsi paler than other leg segments; lateral margins of elytra paler than elytral disc; abdomen brown with posterior and lateral margins paler; photic organs on the VI and VII abdominal segments yellow with white margins; pubescence brunneous. Anterior vitreous areas (pronotal spots) present.

Head large and transverse (frontal view); vertex longitudinally concave with fine punctures dorsally. Eyes large and approximate, partially obstructing labium postventrally. Mandibles long (more than apices of mandibles crossing frontally), falcate, begins tapering at base; incisor region elongate (greater than half of total length of mandible); outer margin convex versus straight as in other taxa ( P. marina , P. nigra , P. skelleyi , and P. milleri ); basal angle approximately 90 degrees dorsally. Maxillary palps with four palpomeres, last palpomere securiform. Labial palps reduced, with three palpomeres.

Antennae filiform, long, extending to posterior end of metathorax ( Fig. 18C ). Scape 2.0×longer than pedicel. Pedicel approximately 1/3 the length of antennomere 3. Antennomere 3 ~1.5×longer than antennomeres 4 to 11. Antennomeres 4 to 11 approximately same size and length. Antennomere 11 rounded at apex. Flagellomeres feebly compressed, but not as compressed as in Lucidota or Ellychnia ; when viewed laterally, flagellomeres are wider than in ventral view. Sensorium present on the distal end of antennomere 11, semi-transparent, small, and subulate.

Pronotum transverse, subelliptical ( Fig. 18C ); longitudinal median carina present, extending from base to apex of pronotum; lateral margins more arcuate and less convergent before the rounded anterior margin compared to Lamprohiza splendidula ; all margins reflexed, very strongly anteriorly along the side and frontal margins, becoming flattened toward the hind angles; basal margin undulate; hind angles denticulate. Central pronotal disc present; surface shiny and sparsely punctured. Anterior vitreous areas (spots) present, large (less than one width of spot between them), falcate, and punctate along outer margins.

Scutellum spatulate; sides curving inward and posterior apex emarginated with longitudinal groove; surface dull and finely punctate ( Fig. 18C ).

Elytra well developed ( Fig. 18C ); coarsely punctate to reticulate; punctate basally, becoming finely reticulate apically. Single costa present medially on each elytron; arising from basal half of elytra and extending approximately to elytral apices. Longitudinal elevated and transverse lines present; transverse lines somewhat irregular in the interstitial spaces giving elytra a reticulated appearance.

Venter. Thorax transverse and rhomboidal. Metathorax with posterior apex notched; medial line present; surface impunctate. Tergal apicolateral angles strongly lobed. Abdominal segments VI and VII bearing photic organs, forming a central stripe approximating the lateral margins of segment. Photic organ absent on abdominal segment VIII. Abdominal segment VII elongate (longer than wide), apical margin deeply emarginated. Abdominal segment VIII length and apical margin same as abdominal segment VII.

Male Genitalia. Basal piece (dorsal view; Fig. 22Y ) wider than long, quadrate, segments contiguous, and moderately sclerotized (light brown). Median lobe (dorsal view; Fig. 22Y ) approximates tips of parameres, overall slender-ovate and rounded (greater than 45 degrees) at apex; median lobe position parallel to parameres ( Fig. 22Z ); sensilla absent; dorsal inner-basal margins cordate; dorsal opening absent; longitudinal dorsal carina absent; lateral line present (lateral view; Fig. 22Z ), comprising of the entire length of median lobe ventral opening present, curvate, comprising of the apical fourth of medial lobe ( Fig. 22 Aa). Parameres (dorsal view; Fig. 22Y ) elongate, slender, sub-parallel; inner lateral margins convex; outer lateral margins undulating; apex roundly acute (lateral view; Fig. 22Z ); proximal flanges (extensions of parameres that curve or extend outward from inner margin) present (dorsal view; Fig. 22Y ), ensiform and restricted medially. Sensilla absent.

Redescription. Female. Length 5.0– 11.5 mm. Width 1.0– 3.5 mm. Apterous and appear physogastric with 8 true abdominal segments. Color. Body white to pale-yellow, parts transparent; eyes black; mouthparts (excluding mandibles) brown-yellow; mandibles brunneous; antennomeres pale-yellow to light brown. Anterior vitreous areas (pronotal spots) absent.

Head ( Fig. 21E–F ) large and transverse (frontal view); vertex not concave. Eyes compound, small and separated, not partially obstructing labium postventrally as it is in males. Mandibles long (more than apices of mandibles crossing frontally), falcate, and begin tapering at base; incisor region elongate (greater than half of total length of mandible); outer margin convex; basal angle approximately 90 degrees dorsally. Maxillary palps with four palpomeres, last palpomere securiform. Labial palps reduced, with three palpomeres.

Antennae ( Fig. 21D–F ) with 7 antennomeres instead of 11 as in males; monofiliform (except for scape, which is cylindrical); short, does not reach to metathorax. Setae present on all antennomeres, simple, and of variable in length. Scape 2.0× longer than pedicel. Pedicel less than 1/3 the length of antennomere 3. Antennomere 3 as long as antennomeres 4 to 6. Antennomere 7 longer than previous antennomeres, rounded at apex. Flagellomeres not feebly compressed. Sensorium present on the distal end of last antennomere, semi-transparent, small, and subulate.

Pronotum ( Fig. 22A ) transverse, subelliptical; longitudinal median carina present, extending from base to apex of pronotum; sides less arcuate and less strongly convergent before the rounded anterior margin compared to Lamprohiza splendidula ; all margins reflexed, very strongly so anteriorly ( Fig. 21B ), becoming flattened toward the hind angles; basal margin weakly undulate; hind angles rounded; anterior margin emarginated. Central pronotal disc absent. Anterior vitreous areas (spots) absent.

Scutellum obstructed or absent, difficult to see.

Elytra absent.

Venter ( Fig. 21C ). Thorax elongate, oblong. Metathorax with posterior apex notched; medial line absent; surface impunctate. Abdomen with 8 visible true abdominal segments. Tergal apicolateral angles not as strongly lobed as in males; rounded apically. Abdominal segments VII and VIII bearing photic organs, consisting of a luminous spot along the lateral margins of each segment. Paired photic organs also present on abdominal segment I that are similar shape to other photic organs approximately reaching the lateral margins of abdominal segment. Observations of light emissions from other abdominal segments have been seen, but could not define a cuticular photic organ on these segments Photic organs can only be seen if the female is alive and emitting light. Females emit light from a single or multiple photic organs located along the lateral margins of abdominal segments I, VII, and VIII. The number or total photic organs seen emitting light can vary from 3 to 9 on different specimens (A. Hodson personal observation; DeCock et al. 2014). The paired photic organs on abdominal segment VIII are thought to be retained larval lights that lose function ( Lloyd 1965; Green 1948). Abdominal segment VII transverse (wider than long). Abdominal segment VIII elongate (longer than wide) with posterior apical margin emarginated.

Larva. 1 st instar. About 1.5 mm. Color. Pale yellow, transparent; head darker in pigmentation that rest of the body (DeCock et al. 2014).

Eggs. Oval. About 0.7 mm long. Color. Yellow-tan. Clutch ranges from 27 to 37 eggs. Hatch between 34–35 days (DeCock et al. 2014).

Etymology. Unknown.

Specimens Examined. Type Material. Neotype (here designated; Figs. 18C ; 24I ). GEORGIA: MADISON Co., 8 mi NE Danielsville jct. Broad R. & Hannah Cr., 8.V.2006. Coll. B.J. Freeman, 1 ♂ ( UGCA).

Other material. FLORIDA: LIBERTY Co., Torreya State Park , malaise trap, 9–17.V.1968, Coll. H.V. Weems Jr. 3 ♂, ( FSCA). GEORGIA: CLARKE Co., Athens , insect flight trap, 30.V–2.VI.1978, Coll. R.H. Turnbow Jr., H.N. Greenbaum, 1 ♂ ( FSCA); same except insect flight trap, 2–5.VI.1978, 2 ♂ (FSCA); same except black light trap, 21–26 May 1973, Coll. R. Turnbow, Det. J.E. Lloyd 1975, 1 ♂ ( FCSA); MADISON Co. , 8 mi NE Danielsville jct. Broad R. & Hannah Cr. (Bud Freeman’s Cabin), 34°13’57.81’ N, 83° 9’39.76’ W, 345 m, in flight, 6.V.2011 Coll. A.M. Hodson, C.H. Scott, B.J. Freeman, Det. A.M. Hodson 2011, 20 males, 2 ♂ (AMHC); same except 5.V .2005 , Coll. B.J. Freeman, Det. A.M. Hodson 2010, 18 ♂ (UGCA); same except 3–5.V .2005 , 22 ♂ (UGCA); same except 8.V .2006 , 10 ♂ (UGCA); FRANKLIN Co., Dovetown Rd & HWY 29 S of Royston ( Mark Dalusky’s Cabin ), 7.V.2011, Coll. A.M. Hodson, C.H. Scott, B.J. Freeman, 6 ♂, 2 ♀ (AMHC); MONROE Co., Forsyth , 1.VI .1970 , malaise trap, Coll. F.T. Naumann. 1 ♂ ( CNCI) . ILLINOIS: UNION Co., Pine Hills , 30.IV.1976, Coll. L. R . Davis Jr., 3 ♂ ( FSCA) . NORTH CAROLINA: AVERY Co., Cranberry, 9–19. VI , 8 males, 1 female [2 ♂, 1 ♀ from MCZC determined incorrectly as Lamprohiza reticulata ] ( CAS, MCZC, SEMC); JACKSON Co., Balsam, Nantahala National Forest , 7–17. VII , 7 ♂ ( SEMC); same except unknown county, Highlands ( Nantahala National Forest ), VI .1988, 1 ♂ [specimen determined incorrectly as Lamprohiza reticulata ] ( MCZC); SWAIN Co., Great Smoky Mountains National Park , along Beech Gap Trail near Straight Fork Road, 35°37.598’ N, 083°12.613’ W, 1004 m , 17.VI .2010 Coll. A.M. Hodson, K. Thomas, Det. A.M. Hodson 2010, 9 ♂ (AMHC); same except Beech Gap Trail, Straight Fork Road, up Beech Gap Trail close to the trailhead, 35°39.700’ N, 83°12.592’ W, 915 m, 13.VI.2005, Coll. A.J. Mayor, B. Nichols, and T. Grannan, Det. A.J. Mayor, 12 ♂ ( GSNP); same except Straight Fork Road heading S towards large bridge, 35°37.365’ N, 083°12.676’ W, 972 m , 17.VI .2010 , Coll. A.M. Hodson, K. Thomas, Det. A.M. Hodson 2010, 7 ♂ (AMHC, GSNP); same except Straight Fork Road near Beech Gap Trail head, 35°37.436’ N, 083°12.91’ W, 978 m, 18.VI .2010 , Coll. A.M. Hodson, K. Thomas, Det. A.M. Hodson 2010, 17 ♂, 1 ♀ (AMHC); same except 35°36.646’ N, 083°12.958’ W, 9525 m, 2 ♂ ( AMHC); same except Deep Creek, Indian Creek trail below wetland, pine forest, 551 m , 23.VI.2005, Coll. A.J. Mayor, J. Jaynes, T. Grannan, Det. A.J. Mayor, 3 ♂ ( GSNP); same except Deep Creek at base of Indian Creek Trail, 35°27.878’ N, 083°26.034’ W, 572 m , 25.VI .2010 , Coll. A.M. Hodson, K. Thomas, 2 ♀ (AMHC); same except Deep Creek on Indian Creek Trail , 35°27.930’ N, 083°25.946’ W, 571 m, 8 ♂ ( AMHC); same except Deep Creek Parking lot, near horse trail , 35°27.862’ N, 083°26.040’ W, 573 m, 3 ♂ ( AMHC); same except Forney Trail, above Andrews Bald , 26.VI .2005 , Coll. A.J. Mayor, 4 ♂ (GSNP); same except Cherokee, Soco Valley , 20.VI .1954 , Coll. J.W. Green, 1 ♂ (CAS); WATAUGA Co., near Boone, at Price Memorial , 2.VII .1967 , Coll. W.B. Biggley, Det. J.E. Lloyd 1972, 2 ♂ (JELC); YANCEY Co., Black Mountains , VI, Det. J.E. Lloyd 1988, 1 ♂ ( AMNH, CAS); same except VI , Det. J.E. Lloyd 1988, 1 ♂ ( JELC); same except 28. VI , 5 ♂ ( AMNH); same except 22. VI , 3 ♂ ( AMNH); same except 22. VI , Det. K.M. Fender, 1 ♂ ( AMNH); same except 12. VI , 1 ♂ ( AMNH); same except 3. VI , Det. J.E. Lloyd 197/88, 2 ♂ ( AMNH); same except V , Det. J.E. Lloyd 1988, 2 ♂ ( AMNH); same except no date, Det. J.E. Lloyd 2002 1 ♂ ( UCMC); same except 1.VI .1902 , no determination, 13 ♂ (CAS); same except Valley of the Black Mountains , 20–23.VI .1906 , Coll. W. Beutenmuller, Det. J.E. Lloyd 1988, 3 ♂ (AMNH); same except Crabtree Meadows , 9.VI.1960, Coll. L. Mason, Det. R. S. Miller 1999, 3 ♂ ( SEMC); same except Crabtree Meadows Campground , Coll. G.W. Bayers, D. Rinkel, and L. Mason, no determination, 1 ♂ ( SEMC); same except VI .1902, no collector, no determiner, 1 ♂ [specimen determined incorrectly as Lamprohiza splendidula with label attached “ Bradley says not splendidula”] ( MCZC) . SOUTH CAROLINA: GREENVILLE Co., Cleveland, Ryan Drive, Firefly Forest-Site 1, 35°03.315’ N, 082°32.448’ W, 345 m , 30.IV.2010, Coll. A.M. Hodson, Det. A.M. Hodson, 23 ♂, 1 ♀ ( AMHC); same except Cleveland, Ryan Drive, Firefly Forest-Site 2 , 35°03.422’ N, 082°32.477’ W, 316 m, 9 ♂ ( AMHC); same except Cleveland, Ryan Drive, Firefly Forest-Site 3 , 35°03.421’ N, 082°32.483’ W, 346 m, 1.V .2010 , 3 ♂ (AMHC); OCONEE Co ., 23.VI .1956 , Coll. Dunavan, Det. J.E. Lloyd 1979, 8 ♂ (CUAC); same except light trap, 1.VI .1956 , 1 ♂ (CUAC). TENNESSEE: BLOUNT Co., Great Smoky Mountains National Park, Cades Cove Intersection of Forge Creek Road and Parsons Branch Road , 17 S, 284414 E, 3953339 N, 588 m , 22.VI .2005 , Coll. A.J. Mayor, J. Jaynes, 5 ♂ (GSNP); KNOX Co., Faust Woods , on hillside, 35°39.178’ N, 083°34.804’ W, 267 m, 21.VI .2010 , Coll. Hayden, Det. A.M. Hodson 2010, 5 ♂, 1 ♀ (AMHC); same except on hilltop, 1012pm, 35.94456 N, 084.20583 W, 9.V .2003 , Coll. L. Faust, Det. M.A. Branham 2003, 2 ♂ (MABC); same except Hardin Valley on Beaver Creek , 35°56’38.13 N, 84°12’15.97 W, 317 m, 3.V .2010 , 2 ♂ (MABC); MORGAN Co., Deer Lodge , 4.VI .1952 , 1 ♂ (CAS); same except Greasy Creek, Log Landing , 36°11.985’ N, 084°35.205’ W, 379 m, 22.VI.2010, Coll. A.M. Hodson, K. Thomas, L. Faust, Hayden, Det. A.M. Hodson 2010, 10 ♂, 2 ♀ (AMHC); same except Greasy Creek ( Emery River ), 36°11’30.24’ N, 84°35’16.51’ W, 348 m, 30.IV.2010, Coll. L. Faust, 2 ♂ ( MABC / AMHC); SEIVER Co., Great Smoky Mountains National Park, Elkmont , 35°39.169’ N, 083°34.814’ W, 688 m , 21.VI.2010, Coll. A.M. Hodson, K. Thomas, L. Faust, Det. A.M. Hodson 2010, 7 ♂ (AMHC); same except 24.VI .2009 , Coll. L. Faust, Det. L. Faust, 2 ♂ (LFPC); same except Elkmont, on Little River , 35°39’20.00’ N, 83°35’4.00’ W, 659 m, 13.V .2010 , 3 ♂ (LFPC) same except Lindsey Creek , wet ravine, 17S, 0284550 E, 3958510 N, 1400 ft., 29.VI .2005 , Coll. B. Nichols, A.J. Mayor, 4 ♂ (GSNP); same except Chimneys Picnic Area Nature Trail, 17 S, 274325 N, 3946251 E, 891 m, 21.VI.2005, Coll. A.J.

Mayor, B. Nichols, J. Jaynes, Det. A.J. Mayor, 6 ♂ (GSNP); same except 35°39.199’ N, 083°29.538’ W, 826 m, 21. VI .2010, Coll. A.M. Hodson, K. Thomas, L. Faust Det. A.M. Hodson 2010, 5 ♂, 3 ♀ ( AMHC); same except Greenbrier , Plemons Cemetery, woods below cemetery, 17 S, 284414 E, 3953339 N , 1843 ft., 2. VI .2005, Coll. A.J. Mayor, 4 ♂ ( GSNP); same except Old Sugarlands Trail at Le Conte Creek, 17 S, 0274834 E, 3950615 N , 762 m, 16. VI .2005, A.J. Mayor, B. Nichols, Det. A.J. Mayor, 5 ♂ ( GSNP); same except Porters Creek Trail, 17 S, 0283072 E, 3951861 N , 579 m, 15. VI .2005, A.J. Mayor, J. Jaynes, Det. A.J. Mayor, 12 ♂ ( GSNP); same except Chilhowee Mountains , 28.V .1950 , Coll. A.C. Cole, 52 ♂ (ECUT); SMITH Co., Elmwood, Coll. J. Corse, 2 ♂ ( CAS) .

Distribution. Known throughout eastern regions of the United States (Arkansas, Florida, Georgia, Illinois, Indiana, Kentucky, North and South Carolina, ( Faust et al. 2019), and Tennessee). Fender (1966) claims that they have also been collected in Texas ( Fig. 13 ).

Phenology. Occurs in late April through early July. Specimens from the Great Smoky Mountains National Park and proximate surrounding areas have two major seasonal peaks. The first peak begins in late April to early May. Specimens collected during this time are called “early Phausis ”. The second peak begins mid to late June. The specimens collected during this time are called “late Phausis ”.

Remarks. This species has been historically associated with the genera Lampyris and Lamprohiza based on shared or similar male morphological characters. These characters include the presence of anterior vitreous areas (spots or windows) on the pronotum; 11 antennomeres; photic organs present on abdominal segments VI and VII; and general appearance in coloration and shape. However, P. reticulata like other Phausis species differs from these genera by having a sensorium on the last antennomere and a strictly New World distribution.

The holotype was described by Say (1825) from a single male specimen Lampyris reticulata from Arkansas given to him by Thomas Knuttel. The name has since been synonymized under the genus Phausis ( LeConte 1852) . Unfortunately, Say’s collection has been largely destroyed either by pests or fire, which makes it difficult, if not impossible to obtain type specimens ( Horn & Kahle 1934; Duval 1859; Mawdsley 1993). Say’s and LeConte’s collections are located in the Museum of Comparative Zoology; however, the type specimen was not contained in this material ( Mawdsley 1993; MCZ web 2010; P. Perkins, personal communication 2011). No other references to the type material have been found and an exhaustive search of collection holdings (Supplementary Table S1; Material and Methods) has not located the type. Therefore, a neotype is herein designated under Article 75 of the International Code of Zoological Nomenclature (ICZN) in order to maintain taxonomic stability and clarification of type locality. The neotype was determined by examining specimens that were identified by previous workers and matched previous descriptions for the species ( Say 1825; LeConte 1852, 1888; Lloyd 1965; Fender 1966; Frick-Ruppert & Rosen 2008).

The female was originally described under the name Lamprohiza reticulata by Wentzel in 1896, but has been redescribed and synonymized by later authors ( Lloyd 1965; Frick-Ruppert & Rosen 2008; De Cock et al. 2014) as well as this publication. LeConte (1878) also describes the female, but he mentions that females have shortened elytra as in the female of Photinus scintillans . This observation suggests LeConte had described a female specimen belonging to different species. Female P. reticulata are not known to have shortened elytra. They are apterous.

Variation. Identified. The apical pronotal margin can be either emarginated or rounded medially across different specimens. The shape of the photic organs in males varies depending on whether the male is alive (photic organs reniform) or dead (photic organ rectangular). Females vary in the number and placement of photic organs which can only be seen when the female is alive and bioluminescent ( Lloyd 1965; Frick-Ruppert & Rosen 2008; De Cock et al. 2014). The number of ommatidia differs between males and females. Males have approximately 1,000 ommatidia, whereas females have 40 ommatidia ( Frick-Ruppert & Rosen 2008).

Biology. The preferred habitat for P. reticulata consists of deciduous forests with thick leaf litter and high levels of moisture. Small streams or seepage areas are generally found proximate to where specimens have been collected. Rhododendron plants are also commonly associated with these habitats (Tennessee, Georgia). This species is a darkactive firefly, where males begin flight approximately one hour after sunset from late April until early July. Females are wingless. They are commonly found within or on top of moist leaf litter, close to the base of vegetation, or on slopes with similar leaf litter and vegetation.

Figures

FIGURE 13. Detail of North American distribution of Phausis reticulata.

FIGURE 18. Dorsal habitus of male Phausis (A–F). (A) Phausis milleri sp. nov.; (B) Phausis nigra; (C) Phausis reticulata; (D) Phausis rhombica; (E) Phausis riversi; (F) Phausis skelleyi.

FIGURE 20. Ventral view of adult male photic organs (A–C). (A) Phausis christineae sp. nov. (paratype); (B) Phausis luminosa; (C) Phausis reticulata.

FIGURE 21. Phausis reticulata Female (A–F). (A–C) Habitus of female (left to right: dorsal, lateral, ventral); (D) Close up of female mouthparts (ventral view); (E) Dissected female head (dorsal view); (F) Dissected female head (ventral view).

FIGURE 22. Phausis aedeagi (A–Jj) (from left to right: dorsal, lateral, ventral view). (A–C) Phausis branhami sp. nov.; (D–F) Phausis californica; (G–I) Phausis dorothae; (J–L) Phausis inaccensa; (M–O) Phausis luminosa; (P–R) Phausis marina; (S–U) Phausis milleri sp. nov.; (V–X) Phausis nigra; (Y–Aa) Phausis reticulata; (Bb–Dd) Phausis rhombica; (Ee–Gg) Phausis riversi; (Hh–Jj) Phausis skelleyi.

FIGURE 24. Type labels of Phausis (A–L). (A) Phausis branhami sp. nov.; (B). Phausis californica; (C) Phausis dorothae; (D) Phausis inaccensa; (E) Phausis luminosa; (F) Phausis marina; (G) Phausis milleri sp. nov.; (H) Phausis nigra; (I) Phausis reticulata; (J) Phausis rhombica; (K) Phausis riversi; (L) Phausis skelleyi.