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Journal of the Lepidopterists' Society 57(2), 2003, 86-91
TWO NEW SUBSPECIES OF PEREUTE LINDEMANNAE AND ONE OF PSEUDOPIERIS VIRIDULA
FROM PANTEPUI, VENEZUELA (PIERIDAE)
Jurg De Marmels, Jose A. Clavijo and Maria E. Chacin
Museo del Instituto de Zoologia Agricola "Francisco Fernandez Yepez", Facultad de Agronomia, Universidad Central de Venezuela, Apartado 4579, Maracay 2101-A, Venezuela
ABSTRACT. Pereute lindemannae pemona, new subspecies, Pereute lindemannae piaroa, new subspecies, and Pseudopieris viridula mimaripa, new subspecies from the Venezuelan Pantepui region are described, diagnosed and illustrated. Their biogeography is outlined and their distribution mapped. A probable sound-producing organ in the male genitalia of Pereute is presented.
Additional key words: biogeography, sound-producing organ, taxonomy.
In recent years the exploration of the remote and almost inaccessible region of Pantepui (Guyana highlands) has been significantly intensified, especially due to the effort of numerous expeditions, partly carried out by non-governmental organizations such as "FUDECI," and "Fundacion TERRAMAR" (both in Caracas). Three new subspecies of Pieridae from this interesting region are described in the present paper. All types are deposited in the Museo del Instituto de Zoologia Agricola "Francisco Fernandez Yepez" (MIZA) of the Universidad Central de Venezuela, at Maracay
Pereute lindemannae pemona De Marmels, Clavijo & Chacin, new subspecies
(Figs. 1-4, 16)
Description. MALE: FW length 31.0 mm. Dorsally: deep black; FW with pale postdiscal band fading from pale yellow between Sc and SR+M1 to white in the medio-cubital space (Mg-CuJ, this portion widely separated from the rest of the pale cross-band by black scales. Small groups of scattered blue scales subapically between R3 and M, and between Mx and M2, as well as between Cuj and Cu2. Base of wing pale steel blue between discal cell and anal margin, along which this color reaches to level of origin of Cuj. Scattered blue scales also within proximal half of discal cell; fringe dark brown. HW black with steel blue area extending from base to about outer margin of median area; small groups of scattered blue scales also along outer margin of wing, between ML and M2, and between M2 and Cur Subcostal space broadly white along costal margin. Ven-trally: FW dark brown, darker in discal cell, paler beyond yellow postdiscal band; an additional, but ill-defined marginal spot of scattered yellow scales between Cux and Cu2. Extensive red scaling in basal third of discal cell; scattered white scales in costal space and on Sc near base of wing. HW dark brown, veins darker; red spot within humeral expansion, an annectent red spot in the angle between Sc and discal cell, penetrating somewhat into the latter; distally of red spot a yellow streak, which is more than two thirds as wide as subcostal space and reaches distally to about level of end of discal cell; a third red spot near wing base, between A1 and A2, but entering space between Cu2 and A}; a white spot at wing root and a small patch of scattered white scales at wing margin, between Rs and Mr Labial palps black with complete white latero-ventral, and shorter dorsolateral line of same color. Antennae white. Thorax with tuft of orange hairs laterally at base of FW and ventrally between and behind second and third pair of legs; a tuft of white hairs between first and second pair of legs. Thorax dorsally with canescent pubescence. Abdomen narrowly black on dorsum, steel blue laterally and white ventrally. Legs sparsely beset with white scales. Genitalia: Tip of uncus
obtuse or scarcely emarginated (Fig. 16). FEMALE: FW length 31.3-32.3 mm; FW of more rounded shape. Dorsally: Color pattern as in male, but all-yellow postdiscal band broader, with few white scales at wing margin between Cu1 and Cu2; steel blue area of HW slightly smaller than in male; blue scales at outer margin of wing absent. Ventrally: yellow streak in subcostal space about three fourths as wide as this space, reaching to about level of distal end of discal cell. Types. Holotype 6: VENEZUELA, Bolivar State, Sierra de Lema, Road El Dorado-Santa Elena de Uairen, km 125, 1090 m, 18 May 1985 (J. De Marmels; Expedition MIZA). Paratypes: Same locality, date and collector, 1 d, 2 9.
Etymology. The "Pemon" are a local Indian tribe.
Diagnosis. Postdiscal band on FW dorsally yellow; extensive red scaling in basal third of discal cell, on FW ventrally Yellow streak between Sc+R2 and discal cell on HW ventrally two thirds to three fourths as wide as that space at level of origin of Rs; a patch of scattered yellow scales at wing margin between Rs and Mp on HW ventrally In conventionally spread specimens a white area is visible near Sc+R1 in space between that vein and discal cell, on HW dorsally Genitalia: Uncus in dorsal view (Fig. 16) very similar to that of P. telthusa (Fig. 18); tip of uncus obtuse or shallowly emarginated.
Remarks. All specimens were caught feeding on a bush with small, white flowers, at the border of a narrow stream in primary forest.
Pereute lindemannae piaroa De Marmels, Clavijo & Chacin, new subspecies
(Figs. 5-8, 17)
Description. MALE: FW length 29.5-34.7 mm (the latter figure of holotype). Dorsally: FW deep black with pale postdiscal band, which fades from yellow between SC and Rs+Mj to white in the medio-cubital space (S3), pale mark here widely separated from rest of pale cross-band, by black scales; a group of scattered blue scales near tip of wing between R3 and Mr Wing base pale steel blue between discal cell anal margin along which this color reaches to level of origin of Cu:; only few blue scales within discal cell near base of wing; fringe dark brown. HW black with steel blue area extending from base to about outer margin of median area. Ventrally: FW black proximally of yellow postdiscal band, dark brown beyond it; the cross-band itself prolongued into space between Cu3 and Cu2; scattered yellow scales in costal space and on Sc near base of wing. Only single or no red scales in discal cell
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Fics. 1-12. 1-4, Pereute lindemannae pemona, new subspecies: 1, Holotype 6 (dorsally); 2, Same (ventrally). 3, Paratype 9 (dorsally); 4, Same (ventrally). 5-8, Pereute lindemannae piaroa, new subspecies: 5, Holotype 6 (dorsally); 6, Same (ventrally). 7, Paratype 9 (dorsally); 8, Same (ventrally). 9-10, Pereute lindemannae lindemannae Reissinger: 9, d from Cerro Neblina, Venezuela (dorsally); 10, Same (ventrally). 11-12, Pereute telthusa Hewitson: 11, c^ from Tingo Maria, Peru (dorsally); 12, Same (ventrally). Scale bar = 10 mm.
near base of wing. HW dark brown, veins darker; red spot within humeral expansion, an annectent red spot in angle between Sc and discal cell; distally of it a yellow streak which is less than half as wide as subcostal space at level of origin of Rs and reaches to about level of origin of M2; a third red spot near base of wing between A1 and A2, penetrating also into space between CuJL and Cu2; a patch of white scales at wing root. Labial palpi black with complete white ventrobasal and shorter dorsolateral line. Antennae pale yellow. Thorax with tuft of orange hairs laterally at base of FW, and ventrally between and behind second and third pair of legs; a tuft of white hairs between first and second pair of legs. Thorax otherwise with canescent dorsal pubescence. Abdomen on dorsum narrowly black, steel blue laterally and white ventrally. Legs sparsely beset with white scales. Genitalia: Tip of uncus obtuse or slightly emar-ginated (Fig. 17). FEMALE. FW length 30.0-34.5 mm; FW of more rounded shape. Dorsally: Color pattern as in male, but FW postdiscal band all yellow, and steel blue area in HW distally scarcely surpassing end of discal cell. Ventrally: FW with few (10-30) red scales at base of discal cell.
Types. Holotype 6: VENEZUELA, Amazonas State, Cerro Yu-taje, 1750 m, 5°45'N, 66°08'W, 12-17 February 1995 (J. Clavijo; Expedition TERRAMAR). Paratypes: same locality, date and collector
7 6, 5 9; same locality, 17-24 February 1995, 2 6, 1 9, (J. L. Garcia; Expedition TERRAMAR).
Etymology. The "Piaroa" are a local Indian tribe. Diagnosis. Postdiscal band on FW dorsally yellow. Only few or no red scales in basal third of discal
Figs. 13-14. Pseudopieris viridula mimaripa, new subspecies: 13, Holotype 6 (dorsally); 14, Same (ventrally). Scale bar = 10 mm.
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Journal of the Lepidopterists' Society
Figs. 15-18. Uncus (dorsal view) in Pereute of the "telthusa group": 15, P. 1. lindemannae (Mt. Neblina); 16, P. I pernona (paratype); 17, P. I. piaroa (paratype); 18, P. telthusa (Tingo Maria).
cell, on FW ventrally; yellow streak between SC+R2 arid discal cell on HW ventrally less than half as wide as that space at level of origin of Rs; same space in HW dorsally appearing entirely blue in conventionally spread specimens; no patch of scattered yellow
scales at wing margin between Rs and Mx on HW ventrally. Genitalia: Tip of uncus obtuse or slightly emarginated.
Remarks. All specimens were caught in primary forest flying through sun-hit spots.
Fir.. 19. Map of north western South America showing localities of Pereute telthusa (empty circles) (after Lamas pers. com., 7 April 1998); P. /. lindemannae (solid circles); P. I. pernona (circle with left half white); P. I. piaroa (circle with right half white); Pseudopieris v. mimaripa (same as jR /. pernona).
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■'w ■■■ t
■jrftr.
^k^y jf ■* j| ;"*■■■'■'
Figs. 20-24. 20, Lateral view of left valva of P. lindemannae piaroa. 21, Detail of the hole of the valva of P. lindemannae piaroa. 22, Lateral view of left valva of P. charops. 23, Detail of the hole of the left valva of P. charops. 24, Ventral view of the genitalia of P. charops. 25, Detail of the gnathos of P. charops. Note: Scale of the bar = 10 mm.
Pereute lindemannae lindemannae Reissinger
(Figs. 9, 10, 15)
The nominate subspecies was described from the Brazilian (southern) slope of Mount Neblina (Reissinger 1970) and is here recorded from Venezuela (northern slope) for the first time. The female is still
unknown. D'Abrera (1981) does not mention the species, as there are no specimens in the Natural History Museum, London.
Diagnosis. MALE: Forewing (FW) length 31.6-34.2 mm. Narrow, mealy-white postdiscal band on FW dorsally. Only single or no red scales in basal
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Journal of the Lepidopterists' Society
third of discal cell on FW ventrally Yellow streak between Sc+R: and discal cell on hindwing (HW) ventrally about half as wide as that space at level of origin of Rs; same space on HW dorsally with white scaling reduced to region near costal margin and not visible in conventionally spread specimens (costal space appearing entirely blue). No patch of yellow scales at wing margin between Rs and Mp on HW ventrally Antennae white. Genitalia: Tip of uncus slightly bifid, ending in two small, laterad directed points (Fig. 15). FEMALE. Unknown.
Material examined. VENEZUELA, Amazonas State, Cerro Neblina, 1800 m, 0°50'40"N, 65°58'10"W, 1870 m, 2 6, 30 November 1984 (A. Chacon and E. Osuna; Expedition FUDECI); 1 6, 2 December 1984 (R. L. Brown; Expedition FUDECI).
Discussion. As already stated by Reissinger (1970,), P. lindemannae is most related to P. telthusa (Hewitson 1860). Following Croizat's (1976:565) view of the bio-geography of Pantepui, a common ancestor of these two species was already distributed across the whole Guyana plateau and from there south westwards to the eastern foothills of the (present) Andes, prior to Andean orogeny. As a consequence of the Andean uplift, which began in the Eocene (Schubert & Huber 1989), the Guyana plateau also rised, probably through isostasy. The plateau population of the ancestral form became separated from that of the lowland and progressively adapted to high elevation life, evolving into the lindemannae stock (primary vicariance and specia-tion event). The populations at the western edge of distribution of this group were "captured" by the uplifting Andes, but were raised to a lesser extent and did not subspeciflcally differentiate at a noticeable degree. The supposed occurrence of some isolated lowland populations of telthusa in the Amazon region, e.g., near Obidos (Rober in Seitz 1924) is emphatically denied by Dr G. Lamas, Lima, Peru (in litt., 19 Jan 1999). The Guyana plateau became at once fractioned by the uplifting movement, the pieces further dissected by erosion and reduced to what can be seen today as isolated remnant mounts and table-top mountains known as "tepuis" (see also Chapman 1931, Tate 1938). It is this secondary vicariance and (sub-)specia-tion event, which explains best the presence in Pantepui of the so far three disjunct subspecies of P. lindemannae. There is, however, also good morphological evidence supporting separation at the species level of lindemannae and telthusa: Male FW is narrower in telthusa than in lindemannae, with C almost straight in the former, while in lindemannae C is visibly arched after first fourth to third of its length. White areas are totally absent from FW (both sexes) in lindemannae, but conspicuous in telthusa. The red spot in humeral area of HW ventrally of telthusa is much reduced in
comparison with same spot in lindemannae. At least in male FW, branching of R4+5 from R2+3 occurs always distally of yellow postdiscal band (ventral view) in telthusa, but within this band in lindemannae. In ventral view, yellow postdiscal band (FW) is sharply limited externally against dark ground color in telthusa, but blurred in lindemannae. Finally, distal margin of valva of lindemannae with its shallow subapical sinuosity resembles the valva of P. charops more closely than that of telthusa, which has a straight outer border.
Although we did not study the genitalia in detail due to lack of large series of specimens, we noticed a very interesting hole located in the middle of the valvae of P. telthusa, P. charops and the three subspecies of P. lindemannae examined. Shape of these holes apparently varies depending on species and possibly can be used as a taxonomic character (Figs. 20-23). The presence of these holes, together with the inflated structure of the valvae, their shape and the way how the valves are exposed in live specimens, as well as structure of gnathos with its numerous denticles (Figs. 24, 25), and the strongly sclerotized aedeagus, all suggest that the genitalia may be used for sound production. Analogous structures are found in other body areas of many Lepidoptera (Scoble 1992). This, of course, needs to be studied in detail, indeed a very promising field for future research.
Pseudopieris viridula mimaripa De Marmels, Clavijo & Chacin, new subspecies
(Figs. 13, 14)
Description. MALE: FW length 23.5 mm. Dorsally: FW white with broad, brown black, mesially zig-zagged apical margin, beginning rather abruptly at the costal margin after branching point of R2, from where it extends more or less diagonally to middle of space between M2 and M3, here sharply bending proximad, following M3 until about half its length (i.e., for about 5 mm), thence again sharply bending outwards, running diagonally towards external margin and following the latter analwards to slightly beyond Cu9, ending at anal angle. Costa and fringe brown black. HW fringe white; broad band of silvery scales along costal margin; a pale, salmon sex brand on costal side of discal cell, beginning proximally of origin of Rs + M1 and ending before branching point of these two veins. Ventrally: FW white with costal and apical region yellow; a long, salmon sex brand along cubital border of discal cell, from close to base of wing outwards to branching point of Cur HW yellow. Palpi white, dorsally and apically black; antennae brown black with ill-defined white an-nules on shaft. Head marbled dorsally, white ventrally. Thorax on dorsum covered with brown hairs anteriorly, with white hairs posteriorly; on venter bearing white and yellow scales and hairs; legs also beset with white and yellow scales. Abdomen mostly white. Genitalia: ill-preserved; uncus and gnathos do not seem to differ from those structures in P. v. viridula or P. nehemia. FEMALE (after A. Neild in litt., 6 Dec 1999): HW apex rounder. Color pattern almost identical to male. Dorsally: black apical region of FW little wider than in male, most notably at the outer margin of Cu^Cu^ wider margin also extending basad along anterior edge of Cu2 for slightly more than one millimeter. Ventrally: Suffusion of yellow scales in postdiscal/subapical region anterior to vein M3 barely discernible, except at costal margin.
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Types. Holotype 6: VENEZUELA, Bolivar State, road El Dorado-Santa Elena de Uairen, km 125,1090 m, 18 May 1985, J. Clav-ijo (Expedition Inst. Zool. Agricola); paratypes: 4 6, 3 9, same area, but km 131.7, 1400 m, 11-14 February 1999 (A. Neild; currently in his private collection).
Etymology. The dorsal color pattern of the new taxon is reminescent of Leptophobia aripa (Boisduval 1836) with which it does not coexist, however.
Diagnosis. Dorsally: FW white with broad, brown black area on tip and along external margin, and produced into a pointed jag mesially between M3 and Cur HW with pale, salmon sex brand along costal side of discal cell. Ventrally: FW white with broadly yellow apical region and pale salmon sex brand along cubital border of discal cell. HW yellow.
Remarks. The holotype specimen was baited with viscera of a cracid bird. The paratype males and females were collected feeding on white flowers of low-growing Eupatorium bushes at the road side (A. Neild in litt, 6 Dec 1999).
Discussion. Some Pseudopieris viridula Felder from Ecuador (Napo) and all from the Venezuelan Coastal Cordillera have tip of FW dorsally only extremely narrowly lined brown black, lacking also the jag between M3 and Cur In some specimens from Peru (Tingo Maria) and from Colombia (Valle del Cauca) the pattern is reminescent of P. nehemia populations, also from Tingo Maria. No differences have been found in the genitalia (uncus and gnathos) of the specimens of the two taxa examined. Therefore, we consider P. v. mimaripa a well-defined (by color pattern and allopatry) subspecies of P. viridula.
The occurrence of P. viridula mimaripa in Pantepui, in disjunction from the populations in the Andes and in the Venezuelan Coastal Cordillera, is seemingly a
consequence of the same geohistorical processes described above for Pereute lindemannae.
Acknowledgments
We wish to thank Mr. Andrew Neild, London (U.K.) for calling our attention on specimens of Pseudopieris viridula mimaripa collected by himself. His description and comments allowed us to include his specimens in the present paper as paratypes. Ing. Quintin Arias, Maracay, helped with the preparation of some genitalia and the solution of photographic and other technical problems. FUDECI (Caracas) organized the expedition to Mt. Neblina. Fundacion TER-RAMAR (Caracas) invited one of us (J. Clavijo) to participate in the expedition to Mt. Yutaje and neighboring tepuis, hence making the collection of one of the new taxa possible. Dr Gerardo Lamas, Uni-versidad Nacional de San Marcos, Lima (Peru), kindly forwarded distributional data of Pereute telthusa and made some usefull comments. Fundacion Polar (Caracas), Fundacite Aragua, Consejo Nacional de Investigaciones Cientificas y Tecnologicas and the Consejo de Desarrollo Cientifico y Humanistico (CDCH) of the Univer-sidad Central de Venezuela gave financial support.
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Received for publication 15 January 2000; revised and accepted 4 November 2002.