BROWN:  New Tortricidae from Panama TROP. LEPID. RES., 29(2): 67-73, 2019 67
Descriptions of four new species of fruit-feeding Tortricidae from Panama 
(Lepidoptera: Tortricidae)
John W. Brown
Department of Entomology, National Museum of Natural History, Washington, DC 20013-7012, USA (e-mail: tortricidae.jwb@gmail.com)
Date of issue online: 28 October 2019
Zoobank Registered: urn:lsid:zoobank.org:pub:105DA4A4-FB44-4074-B56C-6E30684F7666
Electronic copies (ISSN 2575-9256) in PDF format at: http://journals.fcla.edu/troplep; https://zenodo.org; archived by the Institutional 
Repository at the University of Florida (IR@UF), http://ufdc.ufl.edu/ufir; DOI: 10.5281/zenodo.3515363
© The author(s). This is an open access article distributed under the Creative Commons license CC BY-NC 4.0 (https://creativecommons.org/
licenses/by-nc/4.0/).
Abstract: Four new species of Tortricidae reared from fruit in Panama are described and illustrated: Histura panamana sp. nov. 
(Chlidanotinae: Polyorthini); Spinipogon triangularis sp. nov. (Tortricinae: Cochylini); Ricula croceus sp. nov. (Olethreutinae: 
Grapholitini); and Ricula lacistema sp. nov. (Olethreutinae: Grapholitini). Additional field-collected specimens of Spinipogon 
triangularis from Costa Rica are included in the type series of that species.
Key words: Central America, Cochylini, Grapholitini, host plants, Polyorthini, seed-feeder
INTRODUCTION commonly referred to as the DNA barcode, using standard 
 Sanger sequencing procedures employed at the Biodiversity 
Although our knowledge of the systematics and species Institute of Ontario, University of Guelph (Hebert et al., 2003; 
richness of Neotropical Tortricidae has increased dramatically Craft et al., 2010; Wilson, 2012). Barcode Index Numbers 
since the publication of the checklist part of the Atlas of (BINs) using the RESL algorithm were used to help delineate 
Neotropical Lepidoptera (Powell et al., 1995a, b), our groups that potentially represent species (Ratnasingham & 
knowledge of larval host plants has lagged behind and remains Hebert, 2013).
exceedingly poor. A recent study of fruit-feeding insects in 
Panama (Gripenberg et al., 2019) yielded about 450 specimens Collecting in Costa Rica. An ongoing inventory by Daniel 
of tortricid moths representing 20 or so putative species, Janzen and Winnie Hallwachs of the caterpillars of Área de 
providing new host plant records for virtually every species Conservación Guanacaste in northwestern Costa Rica began 
reared. However, because over half of the species are likely in 1977 (Janzen & Hallwachs, 2018). Currently, the bulk of 
undescribed, the contribution to our knowledge of species- collecting and rearing is done by local parataxonomists (Janzen 
level host plant usage is somewhat compromised. Hence, the & Hallwachs, 2011). Caterpillars discovered in the field are 
purpose of this paper is to describe four of the species for taken to “rearing barns” where they are placed singly in plastic 
which adequate material is available and the taxonomic status bags or bottles with cuttings of the host plant species upon 
relatively unambiguous. Where conspecific examples from which they were discovered. As adults emerge, they are frozen, 
Costa Rica are recognized, these specimens are included in the pinned, and labeled. Field-collected caterpillars are labeled 
type series. The intention is to make these names available for with a unique voucher number in the form of YY-SRNP-X…… 
subsequent publications focused on the insects reared during (e.g., 09-SRNP-15328), where the prefix is the last two digits of 
this survey. the year (e.g., 2009), “SRNP” refers to the project “call letters” 
assigned in 1977 (when the initial project site was referred to 
as Santa Rosa National Park), and the suffix is a unique number 
MATERIALS AND METHODS assigned within the year. Over the past few years, collecting 
also has been conducted at a sheet with a black light, and these 
Collecting in Panama. Between July 2010 and November specimens are vouchered similarly. All of these specimens are 
2013, freshly fallen seeds and fruit of trees, shrubs, and barcoded following the protocol mentioned above.
lianas were collected primarily on Barro Colorado Island, 
Canal Zone, Panama. All seeds and fruits were brought to the Identifications and Images. Identifications were made using 
laboratory where they were stored in pots under conditions both morphology and DNA barcodes (BIN numbers are 
resembling those of the ambient forest understory. During the included for each species). Methods for dissecting the genitalia 
rearing period, each pot was checked approximately twice a of adult moths followed those summarized by Brown & Powell 
week during which all adult insects were removed. Specimens (1991). Terminology for morphological structures follows 
of Lepidoptera were pinned, spread, and dried. Tissue samples Horak (1998, 2006). Forewing measurements include the 
(one leg of an adult moth) were used to amplify a ~650 bp fringe. In descriptions of the forewing, “hind margin” refers to 
region of the mitochondrial gene cytochrome oxidase I (COI), the trailing edge of the wing, which frequently is referred to 
68 TROP. LEPID. RES., 29(2): 67-73, 2019 BROWN:  New Tortricidae from Panama
as the “dorsum” in tortricid literature. Slide mounted genitalia the costa to the hind margin, and the length of the labial palpi is 
were examined using dissecting and compound microscopes. usually 2‒3 times the diameter of the compound eye. The male 
Images of adults and genitalia were captured using a Canon genitalia of H. panamana can be distinguished from all other 
EOS 40D digital SLR camera (Canon U.S.A., Lake Success, congeners by the short, rounded socius bearing a dense fascicle 
NY) mounted on a Visionary Digital BK Lab System (Visionary of long, slender, hair-like scales (Fig. 2).
Digital, Palmyra, VA), and edited in Adobe Photoshop.  Description. Adult (Fig. 1). Head: Vertex and upper frons pale orange-yellow, 
Specimen deposition and abbreviations. The specimens lower frons cream colored; length of labial palpus (all segments combined) ca. 4.0 times diameter of compound eye, pale orange-yellow with a few brown 
examined are deposited in the National Museum of Natural scales, cream on inner surface and ventrally, third segment conspicuously 
History, Smithsonian Institution, Washington, D.C., USA upturned, extending well beyond scaling of second; ocellus small; antenna ca. 
(USNM). Additional reared conspecifics not included in the 0.5 times length of forewing costa, with two rows of pale-yellow scales per 
type series are deposited in the collection of the Smithsonian flagellomere, sensory setae extremely short in both sexes. Thorax: Nota pale orange-yellow, tegula yellow. Legs unmodified. Forewing length 6.5–7.5 mm 
Tropical Research Institute, Barro Colorado Island, Panama. (n = 5); pattern complex (Fig. 1), ground color mostly mottled orange-yellow 
Abbreviations used in the data of specimens examined are as with some pale violet and brown scales, a narrow, oblique, brown dash from 
follows: ca. = circa (approximately); r.f. = reared from. near middle of termen extending nearly to costa, arched basad at upper edge 
of discal cell, a similarly colored, irregular, interrupted band originating from 
hind margin near tornus, arching through lower basal part of wing, terminating 
RESULTS near hind margin near forewing base, an area of pale yellow subbasally 
between arch and hind margin (Fig. 1); costal strigulae conspicuous; fringe 
Chlidanotinae: Polyorthini mostly brown, slightly paler in apical region. Hindwing uniformly dark gray-
brown throughout; frenulum with one spine in male, three in female. Fringe 
Histura Razowski, 1981 cream colored. Abdomen: Male with long paired hairpencils from invaginated 
pockets of segment eight. Male genitalia (Figs. 2) with tegumen subrectangular, 
narrowed and slightly rounded dorso-posteriorly; vinculum large, with rounded 
 Histura was described by Razowski (1981) for the single dilation at junction with tegumen (i.e., near point of attachment of costa of 
species Polyortha hirsuta Walsingham, 1914, represented by valva); saccus not developed; uncus moderately long, strongly sclerotized, 
the female holotype from Guatemala and a female paratype rod-like, curved and flattened laterally in apical 0.3; socii short, rounded, with 
from Panama. Razowski & Becker (1981) subsequently added fascicle of long, fine, hair-like scales from a rounded patch near middle; gnathos arms slender, uniform in width throughout, united distally into a weak, mostly 
H. xanthotypa Razowski & Becker, H. doriae Razowski & membranous terminal process; transtilla with large membranous subrectangular 
Becker, and H. chlorotypa Razowski & Becker, all from Brazil. flap with narrow sclerotized margin; juxta small, shield-like; valva somewhat 
Razowski (1984) later described H. bicornigera Razowski membranous, split longitudinally along costa typical of other polyorthines (to 
from Colombia and H. boliviana Razowski from Bolivia, and accommodate hairpencil), broadest at base, slightly attenuate distally, rounded apically, sacculus fairly broad, subrectangular, confined to basal 0.33 of valva, 
transferred Peronea limosa Meyrick, 1912 to the genus. Powell without free projection. Phallus straight, semi-membranous, without cornuti, 
et al. (1995) treated all these species in Histura and added small area of tiny teeth subapically. Female genitalia (Fig. 9) with ovipositor 
Peronea cuprata Meyrick, 1917, a treatment followed by Brown short; papillae anales narrow, slightly arched outwards; apophyses very slender, 
(2005). Razowski and Pelz (2007) described H. brunneotypa apophyses anteriores ca. 1.2 times length of apophyses posteriores; sterigma mostly membranous, with slender, crescent-shaped sclerite at posterior margin 
Razowski & Pelz from Argentina, and Brown and Hoddle (2010) of lamella antevaginalis; ductus bursae slender, frail, membranous, from 
added H. perseavora Brown, from Guatemala. Most recently, shallow antrum, slightly and gradually broadened to corpus bursae; corpus 
Razowski and Becker (2011a) described H. berylla Razowski bursae narrow, membranous, with pair of tiny, plate-like signa.
& Becker from Mexico and H. luteochlora Razowski & Becker DNA barcodes. There are seven sequences available for this 
from Brazil. species in BOLD (BIN ABV2176), with an average distance 
 As currently defined, Histura includes the 12 described of 0.29% among them, and a distance of 7.92% to its nearest 
species listed above, distributed from Veracruz, Mexico to neighbor. 
Tucumán, Argentina. The only species of Histura for which a 
larval host has been reported previously is H. perseavora, which Types. Holotype ♀, Panama, Barro Colorado Island, r.f. Beilschmiedia pendula, 
is a fruit and stem borer in Peresea americana (Lauraceae) in 17 Mar 2011, em: 18 Apr 2001, I. Simón (USNM). Paratypes (2♂, 2♀). Same locality, host, and collector as holotype: no 
Guatemala (Brown & Hoddle, 2010; Hoddle & Brown, 2011). date (2♀) (USNM); 2 Mar 2011, em: 4 Apr 2011 (1♂) (USNM); 29 Mar 2011, 
During the study in Panama, a previously undescribed species em: 18 Apr 2011 (1♂) (USNM).
of Histura was reared from Beilschmiedia pendula (Lauraceae), 
and that species is described below. Distribution and Biology. Histura panamana sp. nov. is 
known only from a series of specimens from Barro Colorado 
Histura panamana sp. nov. Island, Panama. It was reared from the fruit of Beilschmiedia 
(Figs. 1, 2, 9) pendula (Lauraceae).
Etymology. The specific epithet refers to the country of the 
Diagnosis. Histura panamana can be distinguished from all type locality, Panama.
congeners by its yellow orange forewing maculation (Fig. 1) 
and its exceedingly long labial palpi (ca. 4 times the diameter Remarks. The assignment of the new species to Histura is 
of the compound eye) with an upturned third segment. In nearly somewhat provisional. It is easily excluded from Pseudatteria 
all other species of Histura the forewing is gray to brown with Walsingham, 1913, by its forewing shape and maculation 
a variably developed, dark basal patch, the outer margin of (see Clarke 1958). The absence of a long, slender, scobinate 
which extends obliquely outward from the subbasal region of signum in the female genitalia excludes it from Polyortha 
BROWN:  New Tortricidae from Panama TROP. LEPID. RES., 29(2): 67-73, 2019 69
Dognin, 1905, and the relatively shorter valvae exclude it from developed, with long, subrectangular, trough-shaped median process bifurcate 
Ardeutica Meyrick, 1913 (see Razowski, 1984). The slender, distally; valva wide at base, narrowing throughout, costa sclerotized, rounded 
mostly membranous corpus bursae, the shorter valvae, and the apically with a few long setae, sacculus a short triangular process, with a second triangular process near mid-venter. Phallus (Fig. 4a) large, ca. 1.2 times length 
straight, slender, membranous phallus are all most similar to of valva, abruptly narrowed near the middle, pointed apically; vesica without 
those of Histura (see Razowski & Becker, 2011a). However, the cornuti, but with patch of tiny scobinations. Female genitalia (Fig. 10) with 
extremely long labial palpi with an upturned terminal segment papillae anales simple, slender, setose; apophyses long and somewhat broad, 
is unlike that of any Polyorthini I have seen. length of apophyses anteriores ca. equal that of apophyses posteriors; sterigma a lightly sclerotized shield, ostium surrounded by sclerotized collar; ductus 
bursae slender, membranous, well differentiated from corpus bursae, with small 
Tortricinae: Cochylini accessory bursa from frail ductus ca. midway between ostium and junction of 
corpus bursae; corpus bursae large, round, surface covered with minute spicules; 
Spinipogon Razowski, 1967 a large, truncate, cone-like process emanating from near middle of corpus bursae with margins strengthened by slender, linear sclerites.
 Spinipogon was described for S. trivius Razowski and S. DNA barcodes. There are four sequences available for this 
spinifera Razowski from Brazil, and S. signata Razowski from species in BOLD (BIN AAA2890), with an average distance of 
Argentina (Razowski, 1967). Razowski and Becker (1983, 0.21% among them. They are the first sequences of this genus 
1986, 2002), Razowski (1986), and Metzler and Sabourin in BOLD.
(2002) together described an additional 10 species in the genus. Types. Holotype ♀, Panama, Barro Colorado Island, r.f. Guapira standleyana, 
The male genitalia of the type species, S. trivius (Razowski, 13 Jun 2013, S. Gripenberg (USNM).
1967: figs. 71, 72), deviate considerably from those of all other  Paratypes (5♂, 3♀). COSTA RICA: Cartago: Turrialba, 13‒17 Mar 1965 
congeners, hence the genus is somewhat difficult to define. (1♀), W. D. & S. S. Duckworth (USNM). Guanacaste: Área de Conservación 
However, Razowski (1994) recognized two “groups of species,” Guanacaste, Sector Orosí, Casa Rafa, 579 m, 10.95884, -85.49539, 1 Jun 2011, S. Ríos & R. Franco, 11-SRNP-1034946 (1♀), 11-SRNP-103992 (1♂), 11-
and the new species from Panama fits convincingly into one SRNP-103940 (1♂) (USNM). PANAMA: Barro Colorado Island: r.f. Guapira 
of those groups. Spinipogon is primarily Neotropical, ranging standleyana 16 May 2011 (1♂), I. Simón (USNM); 29 Mar 2011, r.f. Tabebuia 
from Central America (Costa Rica, Mexico) and the Caribbean guayacan, 27 May 2013 (1♂) (USNM); 18‒28 Apr 1964 (1♂), 10‒20 Apr 1965 
(British Virgin Islands) to South America (Argentina, Brazil); (1♀), W. D. & S. S. Duckworth (USNM).
a single Nearctic species occurs in the midwestern U.S.A. Distribution and Biology. Spinipogon triangularis sp. nov. is 
(Metzler & Sabourin, 2002). During the study in Panama, a known from Panama and Costa Rica. It has been reared from 
previously undescribed species of Spinipogon was reared from the fruit of Guapira standleyana (Nictaginaceae) and Tabebuia 
fruit, and that species is described below. guayacan (Bignoniaceae).
 
Spinipogon triangularis sp. nov. Etymology. The specific epithet refers to the triangular process 
(Figs. 3, 4, 10) of the valva in the male genitalia.
Diagnosis. Among congeners, Spinipogon triangularis is Olethreutinae: Grapholitini
superficially most similar to S. virginanus Razowski & Becker, 
but it is also similar to many species of Cochylis as well. The Ricula Heinrich, 1926
male genitalia (Fig. 4) are easily distinguished by the narrow 
valva with a conspicuous triangular process at the middle of  Ricula was proposed by Heinrich (1926) for the single North 
the venter that is present in no other species of Spinipogon. American species Lipoptycha maculana Fernald, 1901 (type 
In the female genitalia (Fig. 11), the large, truncate, cone-like locality: USA, Florida). Brown (2005) included two species in 
process from the corpus bursae confirms the assignment of the the genus, the type species and R. dubitana Kuznetsov, 1992 
new species to Spinipogon. from Vietnam. Razowski (2011) synonymized the monotypic 
Riculoides Pastrana, 1952 (type locality: Argentina), described 
Description. Adult (Fig. 3). Head: Vertex and upper frons tawny cream, paler four new species, and transferred two previously described 
on lower frons; length of labial palpus (all segments combined) ca. 1.2 times 
diameter of compound eye, mostly tawny cream, paler on inner surface and species (i.e., Carpocapsa comptana Walker, 1863 and 
ventrally, third segment extending beyond scaling of second; ocellus minute; Hemimene limenita Meyrick, 1922) to the genus. Most recently, 
antenna extending to ca. middle of forewing costa, with two rows of tawny Razowski & Becker (2011b) described an additional 31 species 
scales per flagellomere. Thorax: Nota with smooth appressed tawny cream from the Neotropics, and transferred Grapholita figurana 
scales, tegula tawny. Legs unmodified. Forewing length 5.0–6.5 mm (n = 6); 
forewing ground color cream and tawny, with weak yellowish suffusions and Zeller, 1877 to the genus, resulting in a total of 41 species. With 
brownish streaks, circular blotch in tornal region with costal 0.5 dark brown the exception of R. dubitana from Asia, the genus is restricted 
and lower 0.5 olive-brown, a dark brown, slightly arched band in subterminal to the New World tropics from southern Florida to Argentina.
region beyond circular blotch, a small subrectangular blotch at costa ca. 0.7  Heinrich (1926) indicated that Ricula is most similar 
distance from base to apex, a small brown blotch near middle of discal cell, 
encircled by a crescent-shaped patch of paler brown, an ill-defined semicircular to Talponia but “…differs chiefly in the more approximate 
patch near middle of hind margin; fringe concolorous with ground. Hindwing condition of veins 6 and 7 of the hindwing, the convexity of the 
rather uniform brownish gray throughout; frenulum with one spine in male, termen, and the absence of one signum from the corpus bursa of 
three in female. Fringe pale gray-ocherous. Abdomen: Dorsum tawny cream the female.” Razowski (2011) further refined the definition of the 
with scattered dark brown scales. Male genitalia (Figs. 4) with tegumen 
somewhat rounded dorso-posteriorly, vinculum rounded V-shaped, arms fused genus, mentioning that “The female of Ricula has a broader and 
distally; uncus absent; socius membranous, inconspicuous; transtilla well shorter ductus bursae than other genera of the Dichrorampha-
70 TROP. LEPID. RES., 29(2): 67-73, 2019 BROWN:  New Tortricidae from Panama
Figs. 1‒8. Adults and male genitalia of new species. 1. Holotype female of Histura panamana. 2. Male genitalia of H. panamana, (a) phallus to 
right, USNM slide 150,638. 3. Holotype female of Spinipogon triangularis. 4. Male genitalia of S. triangularis, (a) phallus to right, USNM slide 
144,559. 5. Holotype male of Ricula croceus. 6. Male genitalia of R. croceus, USNM slide 144,609. 7. Female paratype of Ricula lacistema. 8. 
Male genitalia of R. lacistema, USNM slide 144,595.
group, and the corpus bursae has a single signum. The male they are much shorter (or fused to the tegumen) in about half the 
genitalia of Ricula have a pair of setae at the top of the tegumen species currently placed in Ricula. Hence, although features of 
and long sender socii of variable length.” The male genitalia of the male genitalia do not convincingly separate the two genera, 
most species of Ricula and Talponia have remarkably long and the number of signa in the female genitalia is less ambiguous, 
digitate socii. Although they are somewhat variable in length, one in Ricula, two in Talponia.
BROWN:  New Tortricidae from Panama TROP. LEPID. RES., 29(2): 67-73, 2019 71
Figs. 9‒12. Female genitalia of new species. 9. Histura panamana, USNM slide 144,886. 10. 
Spinipogon triangularis, USNM slide 144,606. 11. Ricula croceus, USNM slide 144,610. 12. Ricula 
lacistema, USNM slide 144,596. 
Ricula croceus sp. nov. narrow neck; and socii that are about 0.35 times the length of the 
(Figs. 5, 6, 11) tegumen. The genitalia of R. croceus can be distinguished from 
those of the latter by the more oblique excavation of the valva, 
Diagnosis. The male genitalia of Ricula croceus are most similar to that of R. trachalea Razowski & Becker, 2011 (that 
similar to those of R. excavata Razowski & Becker, 2011, with of R. excvata is more rounded), by the triangular termination 
a relatively narrow valva, broad in the basal half, with an abrupt of the sacculus (more rounded in R. excavata), and the much 
excavation between the sacculus and the cucullus creating a shorter cucullus. The new species is assigned to Ricula on the 
72 TROP. LEPID. RES., 29(2): 67-73, 2019 BROWN:  New Tortricidae from Panama
basis of the long digitate socii in the male genitalia and a single be distinguished from those species by the more distant origin 
signum in the corpus bursae of the female genitalia. of the pair of setae on the top of the tegumen; by a deeper 
Description. Adult (Fig. 5). Head: Vertex and upper frons with shaggy, pale excavation of the venter of the valva between the sacculus and 
gray brown scales, lower frons with white appressed scales; length of labial the cucullus (shallow in R. deflexana); and a more rounded 
palpus (all segments combined) ca. 2.0 times diameter of compound eye, outer margin of the cucullus (slightly angled in R. amethystina). 
white scaled, with slightly shiny, gray scales at distal end of segment II, third Most other species of Ricula have a more elongate-triangular 
segment exposed; ocellus large; antenna thickened, short, ca. 0.4 times length 
of forewing costa, with extremely short sensory setae in both sexes, with pale tegumen similar to that of Talponia. The species is assigned 
brown scales. Thorax: Nota and tegula brown, venter white. Hindleg of male to Ricula on the basis of the long, digitate socii in the male 
with compact patch of pale orange-ocherous secondary scales (hairpencil) genitalia and a single signum in the corpus bursae of the female 
along tibia, originating near trochanter, fitting into receptacle along abdominal genitalia.
plueron. Forewing length 4.0‒4.5 mm (n = 5), costa gently arched throughout, 
termen mostly convexly rounded, with extremely shallow, subapical notch, Description. Adult (Fig. 7). Head: Vertex and upper frons with brown scales, 
male costal fold absent; ground color brown with irregular lines and small lower frons with pale brown appressed scales; length of labial palpus (all 
blotches of orange in distal 0.5 of wing, mostly in form of three slender oblique segments combined) ca. 2.0 times diameter of compound eye, outer surface 
fascia from costa with basal two intersecting four slender straight orange lines pale brown basally, dark gray brown distally, inner surface cream; ocellus large; 
originating near middle of hind margin, and an orange patch in speculum; antenna thickened, short, ca. 0.4 times length of forewing costa, with extremely 
middle of hind margin with broad pale patch encompassing four slender orange short sensory setae in both sexes, with pale brown scales. Thorax: Nota and 
lines extending to ca. lower margin of discal cell, innermost curving distally tegula brown, venter cream. Legs in male unmodified. Forewing length 3.5–4.0 
at discal cell; five conspicuous white costal strigulae from distal 0.5 of wing, mm (n = 4), costa gently arched throughout, termen mostly convexly rounded, 
with two fainter pairs more basal. Fringe dark gray brown. Hindwing nearly with barely discernible trace of subapical notch, male costal fold absent; ground 
uniform brown; frenulum with one spine in male, two in female. Fringe brown. color brown with faint, ill-defined, broad band of sparse individual yellow 
Abdomen: Male genitalia (Fig. 6) with tegumen triangular, a pair of long setae scales in middle portion of wing and in subapical region; faint trace of two 
near top; socius long, ca. 0.35 length of tegumen, slender, with fine setae; basal or three slender metallic bluish lines obliquely from costa; three conspicuous 
0.5 of valva broadest, with weakly convex costa, valva narrowed at short neck white costal strigulae from distal 0.5 of wing, with two fainter pairs more basal. 
by oblique excavation along venter between sacculus and oblong cucullus, Fringe dark gray brown. Hindwing nearly uniform brown; frenulum with one 
latter with dense patch of spines along lower margin; termination of sacculus spine in male, two in female. Fringe brown. Abdomen: Male genitalia (Fig. 8) 
triangular. Phallus long, curved, only slightly larger basally, with 30–40 small with tegumen somewhat pear-shaped, with a pair of long setae near top; socius 
cornuti. Female genitalia (Fig. 11) with papillae anales unmodified; apophyses ca. 0.3 length of tegumen, slender, with fine setae; basal 0.33 of valva broad, 
slender; sterigma a broadly U-shaped, lightly sclerotized plate, weakly bilobed rhombodal, followed by deep rounded concavity in costa creating narrow neck 
posteriorly, with a slender, linear sclerite from each antero-lateral corner of separating sacculus and cucullus; costal margin of cucullus weakly curved, 
sclerotized edge; ductus bursae ca. 0.7 length of corpus bursae, with sclerite ventral margin demarcated basally by abrupt notch, lower half of cucullus 
near middle; corpus bursae pear-shaped with posterior lobe; a single small, with long spines. Phallus short, curved, with 30–40 small deciduous cornuti 
thorn-like signum. (lost in illustrated genitalia). Female genitalia (Fig. 12) with papillae anales 
unmodified; apophyses slender; sterigma a somewhat broad, inverted, crescent-
DNA barcodes. There are six sequences available for this shaped plate, with strong line of sclerotization along anterior margin; antrum 
species in BOLD (BIN ABV2186), with an average distance sclerite restricted to posterior 0.15 of ductus bursae, with ductus seminalis 
of 0.15% among them, and a distance of 8.03% to its nearest originating at anterior end of sclerite; ductus bursae comparatively broad 
neighbor, an undescribed species of Talponia from Costa Rica. throughout, slightly broadened anteriorly, ca. 1.4 times as long as corpus bursae; corpus bursae round, surface somewhat dimpled but without distinct 
Types. Holotype ♂, Panama, Barro Colorado Island, 9 Mar 2011, larva on spiculae or anterior lobe; a single tiny, thorn-like signum with blunt tip.
Heisteria concinna, em: 2 Apr 2011, ID 1348, I. Simón (USNM).
Paratypes (5♂, 2♀). PANAMA: Barro Colorado Island: 5 Feb 2011, em: DNA barcodes. There are 10 sequences available for this 
28 Feb 2011 (1♂); 21 Mar 2011 (1♂); 1 Feb 2011, em: 28 Feb 2011 (1♀); species in BOLD (BIN AAY4670), with an average distance 
28 Mar 2001, em: 13 Apr 2011 (2♂), larva on H. accuminata; 16 Feb 2011, of 0.81% among them, and a distance of 9.95% to its nearest 
em: 8 Mar 2011 (1♀); 13 Jul 2011 (1♂), larva on H. accuminata, all I. Simón 
(USNM). neighbor.
 
Distribution and Biology. Ricula croceus sp. nov. is known Types. Holotype ♂, Panama, Barro Colorado Island, 3 Jun 2011, r.f. Lacistema 
only from Panama, where it has been reared from the fruit of aggregatum, I. Simón (USNM).Paratypes (3♀). Same locality data as holotype, 24 May 2012 (1♀), S. 
Heisteria concinna (Olacaceae) and H. acuminata. Gripenberg; 31 May 2012 (1♀), S. Gripenberg; 13 Jun 2011 (1♀), S. Gripenberg 
(all USNM).
Etymology. The specific epithet refers to the golden color of 
the secondary scales (hairpencil) on the hindleg of the male. Distribution and Biology. Ricula lacistema sp. nov. is known 
from a series of specimens reared from the fruit of Lacistema 
Ricula lacistema sp. nov. aggregatum (Lacistemataceae) (n = 10) on Barro Colorado 
(Figs. 7, 8, 12) Island, Panama.
Diagnosis. Based on the male genitalia, Ricula lacistema is Remarks. The generic assignment of Ricula lacistema, R. 
most closely related to R. deflexa Razowski & Becker, 2011, deflexa, R. amethysina and related species with a somewhat 
and R. amethysina Razowski & Becker, 2011. The three species pear-shaped tegumen (rather than dorso-posteriorly attenuated 
are characterized by a somewhat pear-shaped tegumen bearing tegumen) is somewhat uncertain. Although the presence of 
a pair of long setae from the top; a broad, rhomboidal basal long socii and a single signum in the corpus bursae argue for 
half of the valva followed by a deep concavity of the costa their assignment to Ricula, the relatively broad, rounded dorso-
immediately before a very slender neck separating the sacculus posterior portion of the tegumen is more similar to that of 
and the cucullus; and long, slender socii that are 0.3–0.4 the Riculorampha Rota & Brown, which also has a single signum 
length of the tegumen. The male genitalia of R. lacistema can in the corpus bursae. In addition, barcodes place R. lacistema 
BROWN:  New Tortricidae from Panama TROP. LEPID. RES., 29(2): 67-73, 2019 73
closer to Riculopampha than to Ricula+Talponia. In the northwestern Costa Rica.  PLoS One 6(8): e18123. doi:10.1371/journal.
absence of more conclusive evidence, these species are retained pone.0018123 
provisionally in Ricula. Janzen, D. H., Hallwachs, W. 2018. Dynamic Database for an Inventory of the 
Macrocaterpillar Fauna, and its Food Plants and Parasitoids, of Area de 
Etymology. The specific epithet refers to the genus of the host Conservacion Guanacaste (ACG), Northwestern Costa Rica (nn-SRNP-
plant. nnnnn voucher codes). http://janzen.bio.upenn.edu/caterpillars/database.lasso. [accessed 11 September 2018]
ACKNOWLEDGMENTS Metzler, E., Sabourin, M. 2002. A new species of Spinipogon Razowski, 1967 from two remnant prairies in Ohio (Lepidoptera: Tortricidae, Cochylini). 
Fabreries 27: 69-76.
I thank Sofia Gripenberg and Yves Bassett for allowing 
me to examine specimens of Tortricidae reared from fruit in Powell, J. A., Razowski, J., Brown, J. W. 1995a. Tortricidae: Tortricinae, Chlidanotinae, pp. 138‒151. In: Heppner, J. B. (Ed.), Atlas of Neotropical 
Panama. I thank Daniel Janzen and Winnie Hallwachs for Lepidoptera, Checklist Part II: Hyblaeoidea - Pyraloidea - Tortricoidea. 
providing access to specimens and their barcodes from Área Gainesville, Association for Tropical Lepidoptera, Scientific Publishers.
de Conservación Guanacaste, Costa Rica. Paul Hebert, Jeremy Powell, J. A., J. Razowski, Brown, R. L. 1995b. Tortricidae: Olethreutinae, 
deWaard, Allison Brown, and others at the Biodiversity pp. 151–157. In Heppner, J. B. (Ed.), Atlas of Neotropical Lepidoptera, 
Institute of Ontario, University of Guelph, provided access to Checklist Part II: Hyblaeoidea - Pyraloidea - Tortricoidea. Gainesville, Association for Tropical Lepidoptera, Scientific Publishers.
sequence data and produced neighbor-joining trees. Mark Metz, 
USDA, ARS, assisted with the capture of images, and the final Ratnasingham, S., Hebert, P. D. N. 2013. A DNA-based registry for all animal species: The barcode index number (BIN) system. PLoS One: doi.
plates were prepared by David Adamski, USNM. I thank Todd org/10.1371/journal.pone.0066213.
Gilligan and two anonymous reviewers for helpful comments Razowski, J. 1967. South American Cochylidae (Lepidoptera) from the 
that enhanced the quality and clarity of the manuscript. collection of the British Museum (Natural History). Acta Zoologica 
Cracoviensia 12: 163–210.
LITERATURE CITED Razowski, J. 1981. Notes on the system of Polyorthini (Lepidoptera, 
Tortricidae). Acta Zoologica Cracoviensia 25: 309–318.
Brown, J. W., Hoddle, M. C. 2010. A new species of Histura (Lepidoptera: Razowski, J. 1984. Polyorthini (Lepidoptera, Tortricidae) in the collection 
Tortricidae: Polyorthini) from Guatemala attacking avocados (Persea of the Smithsonian Institution, Washington, D.C. Acta Zoologica 
americana) (Lauraceae). Proceedings of the Entomological Society of Cracoviensia 27: 211–234.
Washington 112: 10–21.
Razowski, J. 1986. New and little known Neotropical Cochylidii (Lepidoptera, 
Brown, J. W., Powell, J. A. 1991. Systematics of the Chrysoxena group of Tortricidae). Acta Zoologica Cracoviensia 29: 373–396.
genera (Lepidoptera: Tortricidae: Euliini). University of California 
Publications in Entomology 111: 87 pp. + figs. Razowski, J. 1994. Synopsis of the Neotropical Cochylini (Lepidoptera: 
Tortricidae). Acta Zoologica Cracoviensia 37: 121–320.
Clarke, J. F. G. 1958. Catalogue of the Ttype Specimens of Microlepidoptera 
in the British Museum (Natural History) Described by Edward Meyrick, Razowski, J., Becker, V. O. 1981. Brazilian Polyorthini (Lepidoptera: 
Volume 3. London, Trustees of the British Museum. 600 pp. Tortricidae). Acta Zoologica Cracoviensia 25: 389–404.
Craft, K. J., Pauls, S. U., Darrow, K., Miller, S. E., Hebert, P. D. N., Razowski, J., Becker, V. O. 1983. Brazilian Cochylidii (Lepidoptera: 
Helgen, L. E., Novotny, V., Weiblen, G. D. 2010. Population genetics Tortricidae). Acta Zoologica Cracoviensia 26: 421–464.
of ecological communities with DNA barcodes: An example from New Razowski, J., Becker, V. O. 1986. Cochylidii (Lepidoptera: Tortricidae) 
Guinea Lepidoptera. Proceedings of the National Academy of Sciences, collected in Central America and Mexico. Acta Zoologica Cracoviensia 
U.S.A. 107: 5041–5046. 29: 441–500.
Gripenberg, S., Basset, Y., Lewis, O. T., Terry, J. T. D., Wright, S. J., Razowski, J., Becker, V. O. 2002. Systematic and faunistic data on Neotropical 
Simón, I., Fernandez, D. C., Cedeño, M., Rivera, M., Barrios, H., Cochylini (Lepidoptera: Tortricidae), with description of new species. 
Brown, J. W., Calderón, O., Cognato, A. I., Miller, S., Morse, G. Part 1. Acta Zoologica Cracoviensia 45: 287–316.
E., Pinzon-Navarro, S., Quicke, D. L. J., Robbins, R. K., Salminen, 
J.-P., Vesterinen, E. 2019. A highly-resolved food web for insect seed Razowski, J., Becker, V. O. 2011a. Systematic and faunistic data on Neotropical 
predators in a species-rich tropical forest. Ecology Letters 2019: 1–12. Tortricidae: Phricanthini, Tortricini, Atteriini, Polyorthini, Chlidanotini 
doi: 10.1111/ele.13359. (Lepidoptera: Tortricidae). SHILAP Revista de Lepidopterología 39: 
161–181.
Hebert, P. D. N., deWaard, J. R., Zakharov, E. V., Prosser, S. W. J., Sones, J. 
E., McKeown, J. T. A., Mantle, B., La Salle, J. 2013. A DNA ‘Barcode Razowski, J., Becker, V. O. 2011b. Systematics and faunistics of Neotropical 
Blitz’: Rapid digitization and sequencing of a natural history collection. Grapholitini, 1: Ricula Heinrich and its allies (Lepidoptera: Tortricidae). 
PLoS One 8 (7): e68535. Acta Zoologia Cracoviensia 54B(1-2): 129–168.
Heinrich, C. 1926. Revision of the North American moths of the subfamilies Wilson, J. J. 2012. DNA Barcodes for Insects, pp. 17–46. In: Kress, W. J., 
Laspeyresiinae and Olethreutinae. United States National Museum Erickson, D. L. (Eds.), DNA barcodes: Methods and Protocols. New 
Bulletin 132: 1–216. York, Springer.
Hoddle, M., Brown, J. W. 2011. Lepidoptera associated with avocado fruit in 
Guatemala. Florida Entomologist 93: 649–650.
Horak, M. 1998. Tortricoidea, pp. 199–215. In: Kristensen, N. (Ed.), 
Lepidoptera, Moths and Butterflies. Volume 1: Evolution, Systematics, 
and Biogeography. Handbook of Zoology 4 (35), Arthropoda: Insecta. 
Berlin & New York, Walter de Gruyter.
Horak, M. 2006. Olethreutine moths of Australia (Lepidoptera: Tortricidae). 
Monographs on Australian Lepidoptera 10. 522 pp.
Janzen, D. H., Hallwachs, W. 2011.  Joining inventory by parataxonomists 
with DNA barcoding of a large complex tropical conserved wildland in