Phyllocnistis drimiphaga Kawahara, Nishida & Davis, 2009

Kawahara, Akito Y., Nishida, Kenji & Davis, Donald R., 2009, Systematics, host plants, and life histories of three new Phyllocnistis species from the central highlands of Costa Rica (Lepidoptera, Gracillariidae, Phyllocnistinae), ZooKeys 27 (27), pp. 7-30 : 12-17

Phyllocnistis drimiphaga Kawahara, Nishida & Davis , sp. n.

urn:lsid:zoobank.org:act:48662DAE-2362-4959-A0B2-C9339B31BB1F

Diagnosis ( Table 1 ). Phyllocnistis drimiphaga is similar to P. maxberryi , but is larger and has slender, sharply angled costal fascia, V-shaped transverse fascia, three costal strigulae, and dissimilar signa. Phyllocnistis drimiphaga differs from P. tropaeolicola in having broad longitudinal fascia, genital valva that are only ̴ 1.8× the length of the vinculum, and paired signa. The pupa has curved, flattened frontal processes, which are reduced in P. maxberryi and conical in P. tropaeolicola .

Adult ( Fig. 2A ). Forewing length 2.9–3.5 mm. Head. Vestiture consisting of smooth, broad, silvery-white scales that overlap anterior margin of eye. Antenna ̴ equal to length of forewing, scape and pedicel enlarged laterally and covered with lanceolate scales, a single row of fine short scales completely encircling each flagellomere. Labial palpus long, slender, ̴ 1.0 mm in length, covered with lustrous white scales. Thorax. Forewing silvery white; with a long pale yellowish-orange longitudinal fascia with dark-gray margins extending 2/3 length of forewing slightly diagonal from base of costa to strongly oblique, costal fascia of similar color across distal third of wing; apex of forewing with three slender, fuscous, costal strigulae; apical to subapical pale yellowish orange bordered by gray; three apical, fuscous strigulae arising from small black apical spot, and one tornal, fuscous strigula also from apical spot; ventral surface mostly dark brown. Hindwing creamy white. Legs mostly silvery white; foretibia fuscous dorsally; fore- and mid-tarsomeres lightly suffused with cream scales dorsally. Abdomen. Length ̴ 2.0 mm, covered in long silver scales. Coremata present on segment VIII of male, consisting of a pair of elongate, inflatable tubular extensions bearing a terminal cluster of long slender scales ( Fig. 4A ). Male genitalia ( Figs 4 A–C). Uncus absent; tegumen complex, consisting of a narrow, sclerotized dorsal arch, continuing caudally, often slightly beyond apex of valva, as an elongate, mostly membranous, basally spinose cylinder that encloses the anal tube; vinculum well developed, ̴ 0.6× length of valva, U- to V-shaped with relative narrow anterior end; valva ( Fig. 4B ) relatively long, ̴1.8× length of vinculum, generally slender with a moderately broad base, very slender for most of its length, then broadening apically to form a prominent dorsal lobe and a smaller ventral lobe ( Fig. 4A ); transtilla arising from mesal base of valva as an elongate, acute process, and continuing mesally to articulate at midline with process from opposite valva. Aedeagus ( Fig. 4C ) slender, weakly sclerotized, externally finely wrinkled cylinder, ̴ equal to length of valva; cornuti absent; phallobase greatly extended as a membranous tube ̴ 1.7–2.0× length of aedeagus; terminal hood of phallobase abruptly inflated and curved at right angle to phallobase. Genitalia slide USNM 33208. Female genitalia ( Figs 4D, E ). Oviscapt greatly reduced; posterior apophyses very short, ̴ 0.8× length of papillae anales; anterior apophyses slightly longer, ̴ 1.3× length of posterior apophyses; ostium bursae opening in membrane between sterna 7 and 8; ductus bursae completely membranous, slen- der, elongate, over 7.5× length of papillae anales and terminating near caudal fifth of corpus bursae; corpus bursae greatly enlarged, ̴ 0.7× length of ductus bursae; walls of corpus bursae membranous except for a pair of ligulate and very dissimilar signa; longest signum ̴ 3× length of shorter member and with 5 short, acute to rounded, flattened spines projecting from one side of signum; shorter signum with a single, blunt, flattened, rounded spine projecting from middle; length of spines ̴ equal to width of signa; ductus seminalis extremely slender, elongate, ̴ 2.3× length of corpus bursae and arising from anterior end of corpus bursae. Genitalia slides USNM 33207, 33273.

Larva ( Figs 10F, G ). Mature sap-feeding larva ̴ 6.5 mm long, yellowish white, head capsule translucent pale brown ( Fig. 10F ). Last instar (cocoon-spinning) larva yellowish white, head capsule yellowish white; ̴ 6.2 mm long ( Fig. 10G ).

Pupa ( Figs 7 , 10I ). Dark brown, up to ̴ 3.8 mm long, diameter ̴ 0.75 mm. Vertex with a stout, triangular frontal process (cocoon-cutter) transversed by a pair of shorter, curved spines ( Figs 7 A–E), and single pair of long setae at base of frons ( Fig. 7C ). Dorsum of A2–A7 with a pair of curved, large spines, arranged roughly in the shape of a V, in between which is a concentration of smaller spines projecting posteriorly ( Figs 7 F–H); each segment with a pair of long, lateral, sensory setae ( Fig. 7K ). A10 prominently furcated ( Figs 7I, J, L ), with a pair of slightly divergent acute processes from caudal apex. Pupal slide USNM 34034.

Types. Holotype ( Fig. 2 A ): ♀, COSTA RICA: Prov. Heredia, 6 km ENE Vara Blanca, 2050 m, 10°10'34"N, 084°06'41"W, 27 Jan 2004, adult emergence, IN- Bio-OET-ALAS transect, col./rear Kenji Nishida , pupa collected 30 Dec 2003, host plant Drimys granadensis . Leaf miner on underside ( USNM) . Paratypes: Immatures: Prov. Cartago: Cerro de la Muerte, La Cañón, Genesis II Cloud Forest Preserve, 2422 m, 09°42'23.4"N, 83°54'36.1"W: 2 sap-feeding larvae, 1 pupa, 12 Sep 2008, Kenji Nishida, host Drimys granadensis ; Prov. San José: Cerro de la Muerte, Paraíso del Quetzal : 2 pupae, USNM 34034 . Adults: same locality as holotype: 1♂, 26 Jan 2004, USNM 33208 ; 1♀, 26 Jan 2004, USNM 33207 ; 1♂, 1♀ ( USNM 33273 ), 28 Jan 2004. 1♀ adult paratype at INBio and UCR, the remaining paratypes at USNM .

Life history ( Fig. 10 ). Mines are narrow, long, and serpentine, with a brown median frass line ( Figs 10A, C, D ) covering most areas of the leaf on small leaves (<6 cm) or half the area in larger leaves. Mines were found on relatively young leaves near the apex of branches, from branches close to the ground up to ̴ 3 m on young trees, along shaded areas of forest trails ( Fig. 1C ) or in the understory. We observed 43 of 48 mines on the abaxial side of the leaf ( Fig. 10A ), and the remaining mines on the adaxial side ( Fig. 10D ). Most mines were singly found on a leaf; however seven of 38 mined leaves carried two mines, either two on the abaxial side or one on both sides. All but one adaxial mine began near the mid-vein and extended along it ( Fig. 10D ). Mature mines are yellowish green in color ( Fig. 10C ). Mining on small, soft, youngleaves frequently caused the leaf margin to curl. We were unable to study the upper canopy for leaf mines.

Early stage mines were typically in the shape of a whorl ( Figs 10 A–C). Flat, oval egg shells were found attached to the leaf surface in the middle of an early mine whorl ( Fig. 10B ). A pupal cocoon fold (̴ 6.5 mm long), typical of Phyllocnistis , was found along leaf margins ( Figs 10A, H, J ) both on the adaxial ( Fig. 10H ) and abaxial sides ( Figs 10A, J ).

In 70 examined mines, only 20 had a live larva or pupa. Remaining mines either were empty or contained dead, early to middle stage sap-feeding larvae. Mortality of sap-feeding stages was most likely caused by desiccation after rupturing of the epidermal layer and by a cf. Ceraphron (Ceraphronidae) parasitoid wasp. In some pupal folds, a pupal shell of an entedonine wasp ( Eulophidae ) was found with a shrunken P. drimiphaga pupal shell. In others, cocoons of Ageniaspis sp. ( Encyrtidae ) were found in a last instar (cocoon-spinning) larval pelt ( Fig. 10K ).

We also discovered active mines of Marmara sp. ( Gracillariidae ) on the abaxial side of same host along the road to El Paraíso del Quetzal. Compared to those of P. drimiphaga , mines were much narrower, whiter, less serpentine, and were typically found near leaf margins.

Host. Drimys granadensis L. f. ( Winteraceae ) ( Fig. 1D ). Drimys Foster & Forster is the only genus in the family Winteraceae found in the New World tropics ( Doust and Drinnan 2004). All other genera of Winteraceae are found in the Old World southern hemisphere with a center of diversity in Southeast Asia ( Gentry 1996; Hartshorn 1983). Drimys granadensis , commonly known as ‘chilemuelo’ or ‘quiebra muelas’, has been recorded from central Mexico (̴ 20°N) south through Central America to northern Peru (̴ 5°S) ( Missouri Botanical Garden 2009). Trees grow to nearly 15 m in height and are characterized by pepper-flavored leaves with white underside surfaces and aromatic bright, white flowers ( Fig. 1E ), found mostly in primary forest (Alfaro- Vindas 2003). In Costa Rica, the species has been recorded between 1100 and 3700 m elevations on both Pacific and Atlantic slopes. Large young leaves are pale green color, sized ̴ 10–15 cm long and 2–4 cm wide (KN, pers. obs.).

Distribution. Known only from cloud forests above 2000 m in Cordillera de Talamanca and Cordillera Volcánica Central. More specifically, specimens have been collected from Heredia Province, 6 km ENE of Vara Blanca; San José Province, Cerro de la Muerte, Paraíso del Quetzal; and Cartago Province, Cerro de la Muerte, Genesis II Cloud Forest Preserve. In February 2009, several additional old leaf mines were observed in Chirripó National Park along the main trail between 2200 and 2700 m elevation.

Etymology. The species name, drimiphaga , comes the host plant genus, Drimys , and the Greek word phaga, meaning “to eat”.

Figures

Figure 1. Habitats and larval host plants of Phyllocnistis species. A Cerro de la Muerte, Villa Mills region, 3000 m and below, in Cordillera de Talamanca B Volcán Barva, ALAS transect, 2000 m, in Braulio Carillo National Park C habitat of P. drimiphaga in Cerro de la Muerte, km 70 Pan-American Hwy, road to El Paraíso del Quetzal, 2700 m, arrow pointing to host plant where mines were found D young stem shoots and leaves of Drimys granadensis of C, growing from base of the tree E flowers and leaves of D. granadensis F habitat of P. maxberryi in Cerro de la Muerte, km 95 Pan-American Hwy, trail front of La Georgina in Villa Mills, 3100 m, arrow pointing to host plant where mines were found Gyoung growth of Gaiadendron punctatum in front, and mature trees with yellow fruits in behind, at ALAS transect in Vara Blanca, 2000 m H habitat of P.tropaeolicola in Cerro de la Muerte, on km 95 Pan-American Hwy, near La Gegina in Mills, 3100 m, arrow pointing to host plant where mines were found I Tropaeolum emarginatum, details of host plants that are shown in H.

Figure 2. Adults of three new Phyllocnistis species from Costa Rica. A Phyllocnistis drimiphaga sp. n., holotype female B P. maxberryi sp. n., holotype female (abdomen removed for dissection) C P. tropaeolicola sp. n., holotype male.

Figure 4. Phyllocnistis drimiphaga sp. n., genitalia. A Male, ventral view B right valva, mesal view C aedeagus D female, lateral view E ventral view of terminal segments. (Scale bar 0.5 mm except for figure B, 0.2 mm.)

Figure 7. Phyllocnistis drimiphaga sp. n., pupa. A Ventral view of head B ventral view of cocoon-cutter C frons D lateral view of head E lateral view of cocoon-cutter F dorsal view of fifth abdominal tergum G spines on fifth abdominal tergum H lateral view of spines on fifth abdominal tergum I view of abdominal tip Į dorsal view of A9–10 K lateral seta on seventh abdominal tergum L ventral view of A9–10. Scale bars 100 µm.

Figure 10. Life history of Phyllocnistis drimiphaga sp. n. A Leaf mines on abaxial side of leaf surface, white square enclosing early mine, arrow pointing to pupal cocoon fold B close-up view of early mine, arrow pointing to egg shell remains C same as figure B, but showing frass pattern (photo taken with sunlight projecting through the leaf from behind) D nearly mature old mine on adaxial side E nearly mature old mine on abaxial side (photo taken from adaxial side) F opened mine showing mature sapfeeding larva in situ G opened young pupal cocoon fold showing cocoon-spinning larva in situ H pupal cocoon fold on adaxial mine I opened pupal cocoon fold showing pupa in situ (dorsal view) Į protruded and attached pupal shell (arrow) on pupal cocoon fold of an abaxial leaf mine K opened pupal cocoon fold on adaxial mine showing Ageniaspis cocoons in situ.