Chrankia altivolans, (BUTLER)

Medeiros, Matthew J., Davis, Don, Howarth, Francis G. & Gillespie, Rosemary, 2009, Evolution of cave living in Hawaiian Schrankia (Lepidoptera: Noctuidae) with description of a remarkable new cave species, Zoological Journal of the Linnean Society 156 (1), pp. 114-139 : 119-125

S CHRANKIA ALTIVOLANS (BUTLER)

Scoparia altivolans Butler, 1880: 9 , in part.

Hypenodes altivolans (Butler) . – Butler, 1881: 325, in part. – Meyrick, 1899: 155, 1904: 347, in part.

Schrankia altivolans (Butler) . – Zimmerman, 1958: 413. – Poole, 1989: 898.

Hypenodes altivolans var. simplex Butler, 1881: 325 , syn. nov.

Schrankia simplex (Butler) ; Zimmerman, 1958: 419. – Poole, 1989: 899.

Hypenodes oxygramma Meyrick, 1899: 154 , pl. 4, fig. 14, syn. nov.

Schrankia oxygramma (Meyrick) . – Zimmerman, 1958: 419. – Poole, 1989: 899.

Hypenodes sarothrura Meyrick, 1899: 155, 1904: 347 , syn. nov.

Schrankia sarothrura (Meyrick) . – Zimmerman, 1958: 419. – Poole, 1989: 899.

Hypenodes arrhecta Meyrick, 1904: 347 , syn. nov.

Schrankia arrhecta (Meyrick) . – Zimmerman, 1958: 417. – Poole, 1989: 898.

Schrankia altivolans is a predominantly epigean Hawaiian species that closely resembles the following, Schrankia howarthi , in general morphology. However, although also possessing a relatively variable forewing pattern, adults of S. altivolans tend to exhibit a generally darker ground colour and larger size. The males may be distinguished from S. howarthi by the broader valva and more pronounced subapical constriction of the cucullus. Females of S. altivolans are distinguished by their more elongated ostium tube and ductus bursae.

Description

Adult ( Fig. 3A–D ): Head: Vestiture similar to S. howarthi except generally darker in colour, more fuscous. Antennal cilia of male between ~ 1¥ and 2¥ diameter of flagellomere in length, that of female similar to that of howarthi . Thorax: Wing venation as shown ( Fig. 4 ). Forewing: Length 6–10 mm; pattern variable, ground colour generally dark to light brown; median band variable, sinuate, sometimes black, white, or orange, and usually interrupted; reniform may be absent or present and black, if present, distal to medial band; antemedial line and postmedial lines sinuate, may be black, white, or orange, vary from strongly visible to indistinct, and are usually interrupted; subterminal line sinuate and pale, if present; black adterminal spots often present near ends of veins; fringe brown. Hindwing: uniformly pale brown to pale grey; base of fringe paler. Abdomen: uniformly pale brown to pale grey.

Male genitalia ( Fig 5A, B ): Similar to S. howarthi except cucullus of valva slightly broader and more abruptly narrowed near apex to form a uniformly slender apical lobe.

Female genitalia ( Fig. 6A ): Similar to S. howarthi except ductus bursae much longer than that of S. howarthi ; length of ductus bursae ~7–9¥ the length of eighth segment; ostium tube also longer than that of S. howarthi , usually ~1.0–1.2¥ length of posterior apophyses. Swelling at juncture of ductus bursae with corpus bursae often pronounced.

Remarks: Previously described Hawaiian Schrankia were separated into five species by Meyrick (1899, 1904) and Zimmerman (1958); however, we could find no morphological characters that varied discretely or consistently in the specimens we examined. All characters, including length of male antennal cilia, thickness of ampulla in the male genitalia, and wing pattern, showed a continuum of variation, even within the type series. For example, the length of the antennal cilia in S. altivolans varies considerably, from ~2¥ to ~1¥ the diameter of flagellomere in length. This observation, along with the lack of male genitalia in three of the five holotypes, and small amount of genetic divergence, renders recognizing individual epigean Hawaiian Schrankia as more than a single species impossible. We have therefore synonymized four of the species under the senior name, S. altivolans . In his review of the Hawaiian Schrankia, Zimmerman (1958) wrote:

‘Although the male genitalia are complex organs, little differences have been found in them. It would appear that they should be expected to display excellent specific characters, but such, disappointingly, is not the case. Moreover, a considerable amount of individual variation has been found in the genitalia of both sexes, and this further compounds the confusion. This group appears to be yet another which is in the process of variation and species formation and clearcut cleavage is not yet established.’

Zimmerman also wrote (1958) of S. altivolans , ‘This is an extremely variable species, and its true limits are not known.’

Type material examined: Schrankia altivolans , lectotype. – USA: HAWAII: 1 ♂, 1880, slide BMNH 1692 (BMNH), here designated . Paralectotype. – USA: HAWAII: 1 ♂, 1880, slide BMNH 1691 (BMNH) .

Schrankia simplex , lectotype. – USA: HAWAII: 1 ♀, 1881 (BMNH), here designated . Paralectotypes. – USA: HAWAII: 1 ♂, 1 ♀, 1881 (BMNH). Schrankia oxygramma , holotype. – USA: HAWAII: Kauai Island: Koholuamano : ♀, iv.1895, slide BMNH 1702 (BMNH). Schrankia sarothrura , Holotype. – USA: HAWAII: Hawaii Island: Olaa : 1 ♂, vii.1895, slide BMNH 1690 (BMNH). Schrankia arrhecta , Lectotype. – USA: HAWAII: Kauai Island : 1219 m, 1 ♀, 1896, slide BMNH 1703, (BMNH) . Paralectotype. – USA: HAWAII: Kauai Island: Mts. Waimea , 1219 m: 1 ♀, vi.1894, Perkins (BMNH) .

Additional material examined: USA: HAWAII: 1 ♂, v.1896 (BMNH). Hawaii Island: Hawaii Volcanoes National Park : 1160 m : 1 ♂, 3 ♀, 13.viii.1977, F. G. Howarth (BPBM); Crater Rim Trail near Volcano House , 1189 m : 1 ♂, 23.iii.1979, G. K. Uchida (BPBM); Escape Road , 1128 m : 4 ♂, 5 ♀, 26.v.2006, slide and DNA 06A52, M. J. Medeiros (BPBM); Field Research Center : 1 ♀, 20.viii.1979, F. G. Howarth ; 1 ♀, 20–21.viii.1979, F. G. Howarth (USNM); Kipuka Puaulu : 2 ♂, 27.v.2006, slide and DNA 06A70, M. J. Medeiros (BPBM); Napau Crater , 1 km south, 800 m : 1 ♂, 5 ♀, 22.viii.1979, F. G. Howarth (USNM); near Thurston Lava Tube , 1189 m : 1 ♀, 13.viii.1977, G. K. Uchida (BPBM); near Thurston Lava Tube , 1189 m : 1 ♀, 15.viii.1977, F. G. Howarth & G. K. Uchida (BPBM); near Thurston Lava Tube , 1189 m : 1 ♂, 3.i.1979, F. G. Howarth (BPBM); Puna, E. Rift , 730 m , Boundary Puu: 2 ♂, 2 ♀, 7.ix.1977, F. G. Howarth (BPBM). Hilo , 610 m : 2 ♂, 3 ♀, xii.1895 (BMNH). Hilo, Puueo Point : 3 ♂, 2 ♀, 10–11.viii.1977, F. G. Howarth (USNM). Hilo, Puueo Point : 1 ♂, 8 ♀, 10–12.viii.1977, F. G. Howarth (BPBM). Honokaa Forest Reserve, Honokaa Cave , 610 m : 1 ♂, 25.iv.1972, F. G. Howarth (USNM). Kaumana, Kaumana Cave : 2 ♂, 26.iv.1972, D. R. Davis, slides 1632′1, 16462 (USNM); 290 m: 1 ♂, 26.ii.1980, slide USNM 33571; 1 ♂, 27.ii.1980, slide USNM 33624, F. G. Howarth (USNM); dark zone : 1 ♂, 30.x.1972, slide USNM 4328, F. G. Howarth (USNM). Kealakekua, Kealakekua Ranch , 1310 m : 1 ♂, 10.xii.1972, slide 33625 USNM, F. G. Howarth, W. C. Gagné, J. Jacobi (USNM) . Keauhou , 600 m : 1 ♂, 7–8.vi.1969, B. Hocking (BPBM). Kilauea : 1 ♀, viii.1896, slide BMNH 1593 (BMNH). Kilauea : 2 ♂, 4 ♀, 12.viii.1958, J. W. Beardsley (BPBM). Kipuka Puaulu, Bird Park Cave , dark zone, 1250 m : 1 ♂, 6 Jul 1973, slide USNM 33088, F. G. Howarth (USNM). Kipuka Puaulu, Bird Park Cave #2, entrance, 220 m : 2 ♂, 29.vii.1976, F. G. Howarth (BPBM). Kipuka Puaulu, Bird Park Cave #3, twilight zone, 1250 m : 1 ♂, 3.iii.1973, F. G. Howarth (BPBM); twilight zone, 1250 m : 1 ♂, 3.iii.1973, F. G. Howarth & S. L. Montgomery (BPBM). Kohala Forest Reserve, White Road , 792 m : 2 ♂, 1 ♀, 30.v.2005, slides and DNA 05A68 & 05A69, M. J. Medeiros (BPBM). Kona , 1219 m : 1 ♀, 8.ix.1892, Perkins (BMNH). Manuka State Park , 400 m : 1 ♀, 12.vii.2004, slide and DNA 04A98, P. T. Oboyski (BPBM). Mauna Loa Bird Park , 1220 m : 1 ♂, 9–11.xii.1976, D. Davis & M. Davis (USNM). Mauna Loa Bird Park Cave #1, entrance, 1220 m : 3 ♂, 9.xii.1976, D. Davis & M. Davis (USNM). Mauna Loa Bird Park Cave #3, entrance, 1250 m : 2 ♂, 1 ♀, 9.xii.1976, D. Davis & M. Davis (USNM). Mauna Loa Strip Road , 2000 m : 6 ♂, 1 ♀, 5.vii.2004, slides and DNA 04A91 & 04A95, slides 04A94, 04A96 & 04A97, P. T. Oboyski (BPBM). Mauna Loa Strip Road , 2000 m : 2 ♂, 28.v.2006, slide and DNA 06A77, M. J. Medeiros (BPBM). Mountain View, Kazumura Cave , entrance sinkhole : 1 ♂, 8.xii.1976, F. G. Howarth, D. Davis, & M. Davis (USNM). Mountain View, Kazumura Cave , twilight zone, 400 m : 1 ♀, 11.iii.1973, F. G. Howarth (BPBM). Olaa , 610 m : 1 ♀, ix.1896 (BMNH). Olaa Forest Reserve, Wright Rd , 975 m : 1 ♂, 14.v.1978, emerged (em.) 1.vi.1978, W. P. Mull (BPBM). Puu Waawaa Forest Bird Sanctuary : 2 ♂, 5.vi.2007, M. J. Medeiros. Saddle Road & Tree Planting Road intersection, 1280 m: 2 ♀, 17.vii.2004, slide and DNA 04A49; 2 ♂, 2 ♀, 25.v.2006, slides and DNA 06A35 & 06A39, M. J. Medeiros (BPBM). Stainback Highway & Tree Planting Road intersection, 853 m: 4 ♂, 2 ♀, 18 Jul 2004, slides and DNA 04A54, 04A56 & 04A57; 2 ♂, 25.v.2005, slide 05A24, M. J. Medeiros (BPBM). Two miles east of Pahoa , 145 m : 1 ♀, 7.ii.1987, K. Arakaki, W. Gagné, G. Nishida, & D. Preston (BPBM) . Upper Waikea Forest Reserve, Stainback Highway , 930 m : 1 ♀, 27.vii.1976, F. G. Howarth (USNM). Volcano , 1160 m : 1 ♂, 6 ♀, 12–29.vii.1976, F. G. Howarth (BPBM). Volcano , 1160 m : 4 ♀, 23–26.ii.1980, F. G. Howarth (BPBM). Volcano , 1160 m : 1 ♂, 2.i.1979, F. G. Howarth & W. P. Mull (BPBM). Volcano , 1160 m : 1 ♂, 1 ♀, 9.ix.1979, S. L. Montgomery (BPBM). Volcano , 1220 m : 1 ♀, 10.xii.1976, D. Davis, M. Davis, & F. G. Howarth (USNM). Kauai Island : 1219 m : 2 ♀,. x.1895 (BMNH). Hanalei Ridge west-north-west of Waikoko Val. 125 m : 2 ♂, 2 ♀, 3.iii.2005, slide and DNA 05A03, P. T. Oboyski (BPBM). Koholuamano , 1219 m : 1 ♂,. iv.1895 (BMNH). Koholuamano , 1219 m : 1 ♀,. iv.1895, slide 1592 BMNH, Perkins (BMNH). Koholuamano , 1219 m : 1 ♂, iv.1896 (BMNH). Kokee : 3 ♀, 6–10.vii.1937, E.C. Zimmerman (BMNH). Kokee State Park, near Kahuamaa Flat , 1202 m : 1 ♀, 27.ii.2005, P. T. Oboyski. Kokee State Park, Kaluapuhi Trail , 1219 m : 4 ♂, 3 ♀, 1–4.vi.2005, slides 05A73, 05A90 &05A94, slides and DNA 05A77, 05A78 & 05A92; 1 ♂, 24.v.2007, M. J. Medeiros (BPBM). Kokee State Park, near Pihea Trailhead , 1158 m : 2 ♀, 27.v.2007, M. J. Medeiros. Kokee State Park , near 17 mile marker of park road : 1 ♂, 1 ♀, 2.vi.2005, M. J. Medeiros. Waimea Mtns , 914 m : 1 ♂, v.1894, Perkins (BMNH). Waimea Mtns , 1219 m : 1 ♂, 1 ♀, v.1894, Perkins (BMNH). Waimea Mtns , 1219 m : 1 ♀, v.1894, slide 1594 BMNH, Perkins (BMNH). Maui Island: E. Maui, Hana, twilight zone, Offal Cave , 90 m : 3 ♂, 26.viii.1977, F. G. Howarth & G. K. Uchida (BPBM) ; 1 ♂, 26.viii.1977, slide USNM 32928, F. G. Howarth & G. K. Uchida (USNM) ; 1 ♂, 26.viii.1977, slide USNM 32928, F. G. Howarth & G. K. Uchida (USNM). E. Maui, Koolau Forest Reserve, Waikamoi Flume Trail , 1300 m : 2 ♂, 24.viii.1977, F. G. Howarth & G. K. Uchida (USNM). E. Maui, Koolau Forest Reserve, Waikamoi Flume Trail , 1300 m : 1 ♀, 24.viii.1977, F. G. Howarth & G. K. Uchida (BPBM). E. Maui, Upper Hana Forest Reserve , 1675 m : near Mid Camp Aaa Kukai Cave twilight : 1 ♂, 5.viii.1973, slide 16951 USNM, F. G. Howarth (USNM); near Mid Camp Trapline Cave twilight : 1 ♂, 5.viii.1973, F. G. Howarth (USNM). Haleakala National Park, Holua Cabin : 1 ♂, 29.vii.2004, slide and DNA 04A86, M. J. Medeiros (BPBM). Haleakala National Park, Hosmer Grove : 5 ♂, 3 ♀, 23.vii.2004, slides 04A69, 04A71, 04A78 & 06B59, slides and DNA, 04A70, 04A73; 2 ♂, 18.ix.2006, M. J. Medeiros (BPBM). Haleakala National Park, near Hosmer Grove , 2075 m : 1 ♀, 17.vi.1976, F. G. Howarth (BPBM). Kula , 975 m : 1 ♀, 28.xii.2003, slide 03A17; 2 ♀, 14.vii.2004, slides and DNA 04A26 & 04A27; 2 ♀, 4.vi.2006, M. J. Medeiros (BPBM). Makawao Forest Reserve , 1280 m : 2 ♂, 8 ♀, 23.v.–3.vii.2006, slide and DNA 06B09, slide 06B27, DNA 06B12, M. J. Medeiros (BPBM). Paliku, Haleakala Crater , 1945 m : 2 ♂, 22.vi.1976, F. G. Howarth (BPBM). Puu Kukui Trail, Haela’au , 914 m : 5 ♂, 6 ♀, 5.vi.2006, slides and DNA 06B45, 06B46, & 06B55, slide 06B43, M. J. Medeiros (BPBM). W. Maui, Kaulalewelewe, Puu Kukui Trail , 910 m : 1 ♂, 3 ♀, 22–23.viii.1977, F. G. Howarth & G. K. Uchida (BPBM) ; 1 ♂ 22–23.viii.1977, F. G. Howarth & G. K. Uchida (USNM). W. Maui, Puu Kukui Trail, Haela’au Cabin , 290 m : 1 ♀, 18.v.1974, F. G. Howarth (USNM). Molokai Island: Kamakou, Puu Kolekole Cabin : 2 ♀, 21.v.2007, M. J. Medeiros. Mtns. , 914 m : 1 ♀, 13.vi.1893, Perkins (BMNH). Oahu Island : 610 m : 1 ♂,. iv.1893, slide 1698 wings BMNH, Perkins (BMNH). 610 m: 1 ♀,. iv.1893, slide 1595 BMNH, Perkins (BMNH). Koolau Mtns, Konahuinui Trail : 1 ♀, 1.v.2006, W. P. Haines. Manana Stream Trail , 305 m : 2 ♀, 18.v.2007, M. J. Medeiros. N.W. Koolau Range : 1 ♂,. v.1901 (BMNH). S.E. Koolau Range : 1 ♀,. vii.1900 (BMNH). Waianae Mtns , 610 m : 1 ♂, 1 ♀,. iv.1892, Perkins (BMNH). Waianae Mtns., Palikea Trail ,> 914 m : 4 ♂, 16.v.2006, slide 06B00, slide and DNA 06B01, W. P. Haines (BPBM) .

Host: Tree roots, probably of many tree species. Larvae occasionally are attracted to rotting mushroom baits used to collect native Drosophila (F. G. Howarth, pers. observ.).

Flight period: Adults and larvae are active throughout the year.

Distribution: This species normally occurs in epigean habitats on at least Kauai, Oahu, Molokai, Lanai, Maui, and Hawaii Islands. It also occasionally occurs in caves on at least Maui and Hawaii Islands.

Figures

Figure 3. Hawaiian Schrankia adults. A, Schrankia altivolans, ♀, Upper Waiakea Forest Reserve, Stainback Highway, 930 m, on tree trunk, Hawaii Island (8.8 mm). B, S. altivolans, ♂, Aaa Kukai Cave, twilight zone, Hawaii Island (8.6 mm). C, S. altivolans, paratype ♂, Kaumana Cave, dark zone, Hawaii Island (5.5 mm). D, S. altivolans, ♂, Kealakekua Ranch Shelter Cave, twilight-dark zone, Hawaii Island (8.0 mm). E, Schrankia howarthi, paratype ♂, Kealakekua Ranch Shelter Cave, twilight zone, Hawaii Island (7.5 mm). F, S. howarthi, paratype ♂, Kealakakua Ranch Stone Wall Cave, twilight zone, Hawaii Island (6.5 mm). G, S. howarthi, paratype ♂, Kazumura Cave, dark zone, Hawaii Island (7.0 mm); H, S. howarthi, paratype ♂, Kealakekua Ranch Shelter Cave, dark zone, Hawaii Island (5.5 mm); I, S. howarthi, holotype ♂, Keauhou Ranch, Keamoku Cave, dark zone, Hawaii Island (5.8 mm). J, S. howarthi, paratype ♂, Ulupalakua Cave, dark zone, Maui Island (5.3 mm). Forewing lengths shown in parentheses.

Figure 4. Schrankia altivolans, ♂, wing venation.

Figure 5. Schrankia ♂ genitalia. A, Schrankia altivolans, slide 04a78, Maui Island: Haleakala National Park, Hosmer Grove; B, S. altivolans, slide 33088, Hawaii Island: Bird Park Cave; C, Schrankia howarthi, slide 33081, Hawaii Island: Kazumura Cave; D, S. howarthi, slide 33082, Hawaii Island: Keamoku Cave; E, S. howarthi, slide 4327, Hawaii Island: Stonewall Cave; F, S. howarthi, slide 32929, Maui Island: Ulupalakua Cave.

Figure 6. Schrankia ♀ genitalia. A, Schrankia altivolans, slide 33867, Hawaii Island: Hawaii Volcanoes National Park, Field Research Center; B, Schrankia howarthi, slide 17505, Hawaii Island: Kealakekua Ranch Shelter Cave.