Early stages of the enigmatic Prodidactis mystica (Meyrick) with comments on its new family assignment (Lepidoptera: Prodidactidae) Author Epstein, Marc E. Author Brown, John W. text Zootaxa 2003 247 1 16 journal article 51306 10.5281/zenodo.156596 572e6dd5-7d46-4929-9ff6-5a717d36124a 1175­5326 156596 Prodidactidae , New Family The following description is based on the single species, Prodidactis mystica (Meyrick) and is intended also to function as a redescription of the genus and species. Hence, it includes a combination of familial­, generic­, and species­level characters. The description includes the states for each of the 24 key characters listed in Heppner’s (1998: Table 2) “Family Characters States.” The importance of certain family­level characters are detailed in the discussion following the description; a comparison of these among the taxa discussed is outlined in Table 1 . Choice of characters and definition of character states are based on Epstein (1996) , Heppner (1998) , and Kristensen (1999). Of the character states described below, the extremely reduced labial palpus and the elongate membranous lobe of the male hindcoxa appear to be of greatest significance, and both are considered autapomorphies for the family. Adult. Head ( Figs. 1-2 , 5-7 ): Smooth scaled or only slightly roughened, frons red; vertex pale gray; ocellus absent; chaetosema absent; maxillary palpus well developed, 3-segmented, covered and tufted with large pale scales; labial palpus greatly reduced, 3- segmented, with crimson-red scales; haustellum well developed, unscaled. Area dorsad of haustellum base with a vertical row of pale scales. Antenna filiform, finely ciliated, not differentiated between sexes; antennal pecten absent. FIGURES 1-4. Adult and larva of Prodidactis mystica . 1) Spread adult moth; 2) Adult in natural resting posture; 3-4) Last instar larva feeding on Nuxia congesta (photographs in Figures 2-4 by Neville Duke). FIGURES 5-6. Adult head and thorax of Prodidactis mystica . 5) Frontal view of head (scales removed from frons) (pr=proboscis (haustellum), mp=maxillary palpus, lp=labial palpus); 6) Dorsal view of head and prothorax. TABLE 1. Comparison of character states of egg, larva, pupa, and adult among families. [Where two states occur, the more common is listed first. * Differs from Heppner (1998)]

Yponomeutidae	Tortricidae	Zygaenidae	Limacodidae	Immidae	Crambidae	Prodidactidae
EGG
egg type	upright	flat	flat	flat	upright	upright	flat

LARVA
L­group on T1 trisetose	trisetose	bisetose	bisetose*	trisetose	bisetose	bisetose
pinaculum w/ absent out seta	absent	absent	absent	absent	absent/ present	present

crochets	multiserial rows	uni­triordinal circle	mesoseries	mesoseries	mesoseries	bi­triordinal circle	biordinal circle
prolegs	A3­6, 10	A3­6, 10	A3­6, 10	A2­7, 10*	A3­6, 10	A3­6, 10	A3­6, 10
L1 and L2 on abd	separate pinaculum	same pinaculum	same pinaculum	same pinaculum	separate pinac.	same pinaculum	same pinaculum
anal fork	absent	present	present	absent	absent	absent	absent
PUPA
general form	fusiform	fusiform	compact	compact	fusiform	fusiform	compact
haustellum	present	present	present	present	present	present	present
labial palpus	small	small	small	small	small	small	small
maxillary pal­ pus	absent	present	absent	absent	absent	absent	present
eye	unscuptured	unsculptured	sculptured	sculptured	unsculptured	unsculptured	unsculptured
dorsal spines	absent	present	present	present	absent	absent	absent
cremaster	absent	present/ absent	vestigial	absent/present	present	present/ absent	absent
cocoon	present	present/ absent	present	present	present	present	present
protrudes at em.	no	yes	yes	yes	?	no	no
ADULT
vertex of head	smooth­scaled	rough­scaled	unmodified	rough­scaled	smooth­ scaled	unmodifed	smooth­scaled
labial palpus	3­segmented	3­segmented	3­segmented	2­3­segmented	3­segmented	3­segmented	3­segmented

TABLE 1 (continued)

Yponomeutidae	Tortricidae	Zygaenidae	Limacodidae	Immidae	Crambidae	Prodidactidae
maxillary pal­ pus	1­2­segmented	2­4­seg­ mented	1­2­seg­ mented	1­3­segmented	1­2­seg­ mented	2­4­seg­ mented	3­segmented
haustellum	naked	naked	naked	naked	naked	scaled	naked
male antenna	filiform	filiform/ variable	bipectinate/ filiform	bipectinate/ filiform	filiform	variable	filiform
antennal pecten	absent/present	absent	absent	absent	absent	absent	absent
ocellus	absent	usually present	usually present	absent	absent/ present	present/ absent	absent
chaetosemata	absent	present	usually present	absent	usually present	present/ absent	absent
venation	heteroneurous	heteroneur­ ous	heteroneur­ ous	heteroneurous	heteroneur­ ous	hetero­ neurous	heteroneurous
wing coupling	frenulate	frenulate	frenulate	frenulate	frenulate	frenulate	frenulate
FW pterostigma	present/absent	absent	absent	absent	absent/ present	absent	absent
FW chorda	present/absent	present/ absent	absent	absent	absent	absent	present
FW anal veins	A1+2 stalked	A1+2 stalked	A1+2 stalked	A1+2 stalked	A1+2 stalked	A1+2 stalked	A1+2 stalked
FW cell vein	absent	present/ absent	usually absent	present	absent	absent	absent
HW pterostigma	absent	absent	absent	absent	absent	absent	absent
HW anal veins	A1+2 A3 stalked	A1+2 A3 stalked	A1A2A3 stalked	A1+2 A3 stalked	A1+2 A3 stalked	A1+2 A3 stalked	A1+2 A3 stalked
HW cell vein	present	usually absent	present/ absent	present	present	absent	absent
epiphysis	present	present	present/ absent	absent	present	present	present
tibial spurs	0­2­4	0­2­4	variable	variable	0­2­4	0­2­4	0­2­4
pretarsal pul­ villus	unmodifed	unmodified	unmodified	unmodified	unmodifed	setiform outgrowth	setiform outgrowth
abdominal tympana	absent	absent	absent	absent	absent	present	absent
abd. apodemes	tineoid­type	tortricoid­ type	tortricoid­ type	tortricoid­type	tortricoid­ type	unmodified	tortricoid­type

Thorax: Whitish gray ( Fig. 6 ). Epiphysis well developed in both sexes. Distinct hairpencils on posterior aspect of hind coxae of male associated with free, elongate, membranous, saclike lobes that are asymmetrical ( Fig. 8 ); hindtibia with 4 spurs. Pulvillus of pretarsus with long setiform outgrowths on dorsal lobe ( Fig. 21 ). Forewing ( Figs. 1­2 , 9 ) moderately broad; costa weakly arched throughout; apex slightly falcate. Length 17­28 mm (females larger). Upper surface buff to yellow buff, faintly rosy tinged in some specimens; anterior 0.66 of costal region suffused whitish, with a few scattered black specks, especially towards dorsal and terminal areas; costa crimson at base; an elongate orange spot near middle of costa; variably developed antemedial, medial, and postmedial zig­zag fasciae of pale orange. No costal fold, upraised scales, or other male secondary structures. Length of discal cell approximately 0.55 forewing length, closed; all veins present, free and separate basally; discal cell between M1 and M2 weak, angled inward; M­stem present, weak, ending between M1 and M2; accessory cell present at distal end of discal cell; CuP weak; 1A+2A forked at base; pterostigma absent. Hindwing uniform pale yellow or pale orange yellow. All veins present, separate; discal cell approximately 0.65 length of hindwing, closed; discal cell between M1 and M2 weak; CuP well developed; tuft of long scales arising near base of CuP; pterostigma absent. Wing coupling frenulate, 1 spine in male, 3 in female. Under surface pale yellow. FIGURES 7­8. Adult head and hind coxa. 7) Laterofrontal view of cleared head (pr=proboscis (haustellum), mp=maxillary palpus, lp=labial palpus); 8) Hind coxa with saclike lobes (arrows point to lobes, st=sternum II of first abdominal segment, ap=apodeme). Abdomen: Apodemes of tortricioid­type ( Fig. 8 ); lacking tympanum. Pale orange buff. Male genitalia ( Fig. 10 ) with uncus stout, as long as width of tergite, attenuate distally, tapering to a rounded point, densely covered with stout hairs directed anteriorly; attached to tegumen at a partially membranous point of articulation. Gnathos arms moderately broad, united into a single narrow, mesal process, strongly up­curved in distal 0.33, ending in a foot­shaped sclerite with pointed toe. Socius apparently absent. Valva long, with broad basal portion, slender central portion, and broad apical region; costa strongly sclerotized; inner surface densely covered with long hairs. Aedeagus stout, bent near middle, slightly undulate; attenuate distally with short spine on left side near apex; cornuti absent. Female genitalia ( Fig. 11 ) with papillae anales broad, rounded ventrally, moderately sclerotized, densely covered with long hairs; not fused dorsally. Apophyses moderately long, slender. Sterigma a simple plate, ostium bursae simple. Ductus bursae nearly as long as abdomen; corpus bursae small, rounded; signum absent. FIGURES 9­11. Adult morphology of Prodidactis mystica . 9) Wing venation; 10) Male genitalia; 11) Female genitalia. FIGURES 12­14. Pupa and larva of Prodidactis mystica . 12) Venter of pupa; 13) Dorsum of pupa (without caudal segments); 14) Chaetotaxy of larva of Prodidactis mystica . Last Instar Larva (n = 3) ( Figs. 3­4 , 14 , 17­19 ). General: Length 22­26 mm , rather stout. Head red brown with no conspicuous markings. Prothoracic shield dark greenish brown with dorsal separation of halves indicated by a yellow line. Body greenish yellow, with a narrow dark green longitudinal middorsal line and a broad, dark green longitudinal subdorsal line extending ventrad to dorsal edge of spiracles; pinacula conspicuous, yellow; non­setal bearing pinacula on meso­ and metathorax red brown; secondary setae absent. Head: Frons extending about 0.7 to occipital foramen; stemmata arranged in an arc except for S5 ( Fig. 19 ); spinneret moderate in length, rounded distally ( Figs. 17­18 ) ; mandible with three large teeth, one smaller tooth, and a flattened region at inner portion ( Fig. 18 ). Thorax ( Fig. 14 ): Prothorax with bisetose L­group on common prespiracular pinaculum; SD2 seta approximately 0.4 times length of SD1; spiracle on prothorax surrounded by large, clear disk. SV­group bisetose on prothorax, unisetose on meso­ and metathorax. Meso­ and metathorax each with non­seta­bearing dorsal pinacula; mesothorax with three naked pinacula, two slightly dorsad of D pinaculum, one on each side, and a third anterad and slightly ventrad of SD pinaculum; metathorax similar, with hint of an additional naked pinaculum anterad and ventrad of first anterior pinaculum. Abdomen ( Fig. 14 ): Spiracles surrounded by inconspicuous unsclerotized disk; spiracle on A8 slightly larger than those on A1­7; D2s on separate pinacula on A9; SD1 well dorsad and slightly anterad of spiracle on A1­7, conspicuously more anterad on A8; L1 and L2 together, in horizontal row, on large common lateral pinaculum below spiracle on A1­8; L1+L2 pinaculum on A9 nearly vertical; L3 absent on A9; SV group on 1,2,3­6,7,8,9 usually 2:3:3:2:2:1; anal fork absent. Crochets 70­75 in uniform biordinal circle on A3­6; crochets 50­55 in a semicircle on A10. Pupa. General ( Figs. 12­13 , 15­16 ): Short and broad, compact, zygaeniform. Compound eye unsculptured; haustellum elongate, extending to distal end of front legs; maxillary palpi extending to near end of antennae; labial palpus a small triangle. Abdomen unmodified, without rows of spines on dorsum; spiracle not visible on A1; cremaster absent. Cocoon. General ( Fig. 20 ): Somewhat smooth on one surface and rough on opposite side; oval and flattened, 1.8 cm x 1.0 cm; opening along edge of narrow end, ca. 0.33 length of cocoon. Life history. Eggs are flat and scalelike (N. Duke, pers. comm.), similar to those of many Tortricidae ( Horak 1999 ) and Limacodidae ( Epstein 1996 ) , and are laid on the leaves of the host. Behavior of the early instars was not recorded. Fourth and fifth instars are solitary, apparently feeding from the upper surface on leaves of Nuxia congesta , as supported by photographic evidence ( Figs. 3­4 ). The larvae form a shelter in a rolled leaf, but spend more time feeding exposed on the leaf than do tortricids (N. Duke, pers. comm.). Pupation occurs on the host in a dense, tough cocoon spun on or between leaves forming a shelter; overwintering is apparently accomplished in this stage. The pupa does not protrude from the cocoon prior to eclosion. Adults have been collected from October through March ( Janse 1964: 22 ). Geographic distribution. According to Janse (1964: 22) , P. m y s t i c a is known from Durban Nggeleni (Pondoland), Nkandhla Forest (Zululand), Noordkaap, Barberton, Marieps Mountain, Graskop, Kowyn's Pass, Pretoria, Woodbush, Malta , and Zoutpansberg (all Transvaal). Specimens in the Transvaal Museum (M. Krüger, pers. comm.) provide the following distribution data: SOUTH AFRICA : TRANSVAAL: The Downs, Entabeni Forest, Kownyn's Pass, Pilgrim's Rest District, Uitsoek, Louis Trichardt, Piesangkop NE, Louis Trichardt Mariepskop Forestry, Malta Forest, Woodbush, Barbeton, De Hoek Forestry, Timbadiola Forestry, Pretoria North. NATAL : Ngoya Forest, Mtunzini District, Durban, Nkhandla Forest. EASTERN CAPE: Transkei, the Haven, Nggeleni. SWAZILAND : Mbabane. ZIMBABWE : Vumba Mountains, Lundi.