Journal of Insect Science: Vol. 11 | Article 7 Brown et al.
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Tortricid moths reared from the invasive weed Mexican palo 
verde, Parkinsonia aculeata, with comments on their host 
specificity, biology, geographic distribution, and systematics
John W. Brown1a, Ricardo Segura2b, Quiyari Santiago-Jim?nez2c, Jadranka Rota3d, Tim A. 
Heard4e
1Systematic Entomology Laboratory, Plant Sciences Institute, ARS, U.S. Department of Agriculture, National 
Museum of Natural History, Washington, DC 20013-7012
2CSIRO Ecosystem Sciences, Mexican Field Station, A. Carl?n No. 5, Col. Ejido 1 de Mayo, Boca del R?o C.P. 
94297, Veracruz, Mexico
3Department of Entomology, National Museum of Natural History, Smithsonian Institution, Washington, DC 
20013-7012
4CSIRO Ecosystem Sciences, Brisbane, Australia
Abstract
As part of efforts to identify native herbivores of Mexican palo verde, Parkinsonia aculeata L. 
(Leguminosae: Caesalpinioideae), as potential biological control agents against this invasive 
weed in Australia, ten species of Tortricidae (Lepidoptera) were reared from Guatemala, Mexico, 
Nicaragua, and Venezuela: Amorbia concavana (Zeller), Platynota rostrana (Walker), Platynota
helianthes (Meyrick), Platynota stultana Walsingham (all Tortricinae: Sparganothini), Rudenia
leguminana (Busck), Cochylis sp. (both Tortricinae: Cochylini), Ofatulena duodecemstriata
(Walsingham), O. luminosa Heinrich, Ofatulena sp. (all Olethreutinae: Grapholitini), and 
Crocidosema lantana Busck (Olethreutinae: Eucosmini). Significant geographic range extensions 
are provided for O. duodecemstriata and R. leguminana. These are the first documented records 
of P. aculeata as a host plant for all but O. luminosa. The four species of Sparganothini are 
polyphagous; in contrast, the two Cochylini and three Grapholitini likely are specialists on 
Leguminosae. Ofatulena luminosa is possibly host specific on P. aculeata. Host trials with 
Rudenia leguminana also provide some evidence of specificity, in contrast to historical rearing 
records. To examine the possibility that R. leguminana is a complex of species, two data sets of 
molecular markers were examined: (1) a combined data set of two mitochondrial markers (a 781-
basepair region of cytochrome c oxidase I (COI) and a 685-basepair region of cytochrome c 
oxidase II) and one nuclear marker (a 531-basepair region of the 28S domain 2); and (2) the 650-
basepair ?barcode? region of COI. Analyses of both data sets strongly suggest that individuals 
examined in this study belong to more than one species.
Keywords: Australia, Amorbia, biological control, Cochylis, DNA sequences, host plants, Neotropics, Ofatulena, Playtnota,
Rudenia, taxonomy
Correspondence: a john.brown@ars.usda.gov, b csiromfs@prodigy.net.mx, c quiyari@hotmail.com, d
jadranka.rota@gmail.com, e tim.heard@csiro.au
Received: 19 June 2009, Accepted: 13 September 2010
Copyright : This is an open access paper. We use the Creative Commons Attribution 3.0 license that permits 
unrestricted use, provided that the paper is properly attributed.
ISSN: 1536-2442 | Vol. 11, Number 7
Cite this paper as:
Brown JW, Segura R, Santiago-Jim?nez Q, Rota J, Heard TA. 2011. Tortricid moths reared from the invasive weed 
Mexican palo verde, Parkinsonia aculeata, with comments on their host specificity, biology, geographic distribution, and 
systematics. Journal of Insect Science 11:7 available online: insectscience.org/11.7
Journal of Insect Science: Vol. 11 | Article 7 Brown et al.
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Introduction
Parkinsonia aculeata L. (Leguminosae: 
Caesalpinioideae), commonly known as 
Jerusalem thorn, Mexican palo verde, and/or 
jellybean tree, is a large shrub or small tree 
native to North and South America that shows 
marked genetic divergence among populations 
(Hawkins et al. 2007). It is used as an 
ornamental in tropical and subtropical 
climates, and it has been used to re-vegetate
desertified regions throughout the pantropics
(van Klinken et al. 2008). It has escaped
cultivation and become established in many 
places around the globe. It was introduced 
into Australia as an ornamental and shade tree 
around 1900 (Woods 1992) and by 1906 was 
considered weedy in some parts of 
Queensland (Bailey 1906). Now it is regarded 
as one of the most troublesome invasive 
weeds in northern Australia and is recognized 
as a Weed of National Significance. 
Parkinsonia aculeata currently infests over 
800,000 hectares, mainly along watercourses, 
in Western Australia, Queensland, and the 
Northern Territory. It has the potential to 
invade most of the semi-arid to subhumid 
tropical areas in Australia (van Klinken et al. 
2008).
Parkinsonia aculeata has been the target of 
biological control investigations by Australian 
researchers for nearly two decades (Woods 
1992; Heard 2006). During initial exploration 
in Texas and northeastern Mexico that
focused on the discovery of native herbivores 
of P. aculeata, members of six insect orders 
were recorded (Coleoptera, Diptera, 
Hemiptera, Lepidoptera, Orthoptera, and 
Thysanoptera) (Woods 1992). The five 
species of Lepidoptera reared during these 
efforts were Brachyacma palpigera
(Walsingham) (Gelechiidae), Anacamptodes
cf. obliquaria Grote (Geometridae), Melipotis
acontioides (Guen?e) (Noctuidae), Carmenta
sp. (Sesiidae), and Ofatulena luminosa
Heinrich (Tortricidae). More recently (1995-
2009) additional species of Lepidoptera have 
been recorded from this weed pest. The 
purpose of this paper is to present records of 
the ten species of Tortricidae that have been
reared from Parkinsonia aculeata, along with 
comments on their host range, damage, and 
geographic distribution. The results of host 
specificity trials for Platynota stultana
Walsingham and Rudenia leguminana
(Busck) are also presented. All of the material
listed in the ?Specimens Examined? sections 
was reared from Parkinsonia aculeata (unless 
stated otherwise) during the most recent 
ongoing study (1995-2009); other records of 
host-use and geographic distribution are from 
the literature and/or museum collections.
Voucher specimens are deposited in the 
collections of the National Museum of Natural
History (USNM), Smithsonian Institution, 
Washington, D.C., U.S.A.; and the Australian 
National Insect Collection (ANIC), Canberra, 
Australia. EME refers to Essig Museum of 
Entomology, University of California, 
Berkeley, U.S.A. LPL stands for Long Pocket 
Laboratories, CSIRO Entomology, Brisbane, 
Queensland, Australia; each reared specimen 
is assigned an LPL number for tracking.
Species Accounts
Amorbia concavana (Zeller) 
(Figures 1, 2)
In a systematic revision of Amorbia, Phillips-
Rodriguez and Powell (2007) recorded A.
concavana from Costa Rica, Cuba, 
Guatemala, Honduras, Mexico, and Panama 
and provided illustrations of the adult and 
genitalia. Specimens that key to Amorbia
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concavana (Phillips-Rodriguez and Powell 
2007) were reared from P. aculeata in 
Guatemala and Mexico. However, the 
specimens also resemble A. emigratella
Busck, another widespread, polyphagous 
species of Amorbia.
Biology. Phillips-Rodriguez and Powell 
(2007) listed rearing records from Mimosa
pigra L. (Leguminosae) in Mexico, Phaseolus
sp. (Leguminosae) in Cuba, and Inga vera
Willd. (Leguminosae) and Hammelia sp. 
(Rubiaceae) in Costa Rica. Although most 
recorded hosts are in Leguminosae, A.
concavana likely is a polyphagous leaf-roller
as are other members of the genus.
Specimens Examined. GUATEMALA: 
Jutiapa: Lago Guija, 14? 73.9? N, 89? 32? W, 
14 January 2007 (1
), R. Segura & M. 
Mart?nez, LPL 10955. MEXICO: San Luis 
Potos?: Laguna Ajinche, 22? 10.5? N, 98? 21? 
W, 18 May 2006 (1, 2
), R. Segura & C. 
Pascacio, LPL 10688, 10887. La Marland, 22? 
08.4? N, 98? 24.9? W, 9 November 2006 (1),
R. Segura & M. Mart?nez, LPL 10897. 
Veracruz: Canoas, 22? 10.1? N, 98? 07.7? W, 
16 August 2002 (6, 3
), M. Mart?nez & C. 
Pascacio, LPL 12047, 12056.
Platynota rostrana (Walker) complex 
(Figures 3, 4)
Platynota rostrana, which may represent a 
complex of closely related species, ranges 
throughout much of the southeastern U.S. and 
the northern Neotropics, including the 
Caribbean (based on specimens in USNM).
Males are characterized by a complex hood of 
scaling on the frons of the head and a long 
costal fold that extends about 0.7 times the 
length of the forewing.
Biology. Platynota rostrana has been reared 
from more than 100 different plant species in 
over 20 different families (Heinrich 1921; 
MacKay 1962; d'Araujo Silva et al. 1968; 
Okumura 1974; Bruner et al. 1975; Diniz and 
Morais 1995; McClay et al. 1995). Examples 
of this species were reared from P. aculeata in 
Veracruz and San Luis Potos?, Mexico.
Specimens Examined. MEXICO: San Luis 
Potos?: Playa Buda, 22? 10.8? N, 98? 22.8? W, 
10 November 2006 (1
), R. Segura & M. 
Mart?nez, LPL 10926. El Caracol, 22? 09.6? 
N, 98? 01.9? W, 10 November 2006 (1), R. 
Segura & M. Mart?nez, LPL 10912. L?zaro 
C?rdenas, 22? 15.7? N, 98? 8.73? W, 17 May 
2006 (1), M. Mart?nez & C. Pascacio, LPL 
10657. Ciudad Cuauht?moc, 22? 12.8? N, 97? 
53.8? W, 17 May 2006 (1), R. Segura & C. 
Pascacio, LPL 10664.
Platynota helianthes (Meyrick)
(Figures 5, 6)
Platynota subargentea ranges from Mexico to 
Venezuela (based on specimens in USNM). 
The male is similar to P. rostrana, with a 
complex hood on the frons and a long 
forewing costal fold; but the two species are 
easily distinguished by features of the 
genitalia, in particular those of the female.
Biology. This species has been reared from 
Jatropha gossypifolia L. (Euphorbiaceae), 
Casearia corymbosa H. B. & K. 
(Flacourtiaceae), Leucania leucocephala
(Lam.) De Wit (Leguminosae), Mimosa pigra
(Leguminosae), and Psidium guajava L. 
(Myrtaceae) in Mexico (all USNM). We 
reared P. helianthes from Parkinsonia
aculeata in Nicaragua and Mexico.
Specimens Examined. MEXICO: San Luis 
Potos?: La Marland 2, 22? 10.1? N, 98? 24.4? 
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W, 1 March 2008 (1
), R. Segura, M. 
Mart?nez & C. Pascacio, LPL 11587.
Veracruz: Canoas, 22? 10.1? N, 98? 07.7? W, 
16 August 2002 (2, 3
), M. Mart?nez & C. 
Pascacio, LPL 12054, 12058, 12062, 12064. 
NICARAGUA: Matagalpa, Las Guayabas 1, 
11 December 1999 (1
), T. Heard & R. 
Segura, LPL 8839.
Platynota stultana Walsingham 
(Figures 7, 8)
Platynota stultana is one of the smaller 
species in the genus. It ranges throughout the 
southwestern United States and northern 
Mexico; it is adventive in California (Powell 
1983), Hawaii (Miller and Hodges 1995), 
Florida (Kimball 1965), Virginia, and
Washington, D.C. (USNM).
Biology. This species is known in the 
American economic literature as the 
omnivorous ?leaf-roller?. Larval hosts in 
native situations include western ragweed 
(Ambrosia psilostachya DC.; Asteraceae), 
gooseberry (Ribes sp.; Grossulariaceae), and 
youngberries (Rubus sp.; Rosaceae). 
Agricultural and ornamental hosts include a 
wide range of native and cultivated plants 
(Atkins et al. 1957; MacKay 1962; Powell 
1983; Miller and Hodges 1995). Rolled leaves 
of Parkinsonia aculeata harboring larvae of 
Platynota stultana were collected at several 
sites in a large swamp known as Laguna 


Figure 1-8. Adults of leaf-roller moths reared from Parkinsonia aculeata. 1) Amorbia concavana (Zeller) (
). 2) A. concavana
(). 3) Platynota rostrana (Walker) (
). 4) P. rostrana (). 5) Platynota subargentea Walsingham (
). 6) P. subargentea (). 7)
Platynota stultana (
). 8) Platynota stultana (). Scale bar = 5 mm. High quality figures are available online.
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 5
Ajinche near the town of Ebano, San Luis 
Potos?, Mexico, on 10 November 2006. First 
instars fed inside the pinnule and soft rachis of 
the host, eventually feeding externally, tying 
the young leaves together with silk to make a 
shelter from which they foraged on young 
leaves. In a host specificity test, oviposition,
development, and adult emergence of P.
stultana occurred on all test plant species 
(Mimosa asperata L., Leucaena leucocephala
(Lam.) de Wit, Caesalpinia pulcherrima (L.) 
Sw., Delonix regia (Bojer ex Hook.) Raf., 
Tamarindus indica L., and Parkinsonia
texana) (all Leguminosae).
Specimens Examined. MEXICO: San Luis 
Potos?: Pozo La Pez No. 1, 22? 5? N, 98? 23.4? 
W, 10 November 2006 (13, 3
), R. Segura 
& M. Mart?nez, LPL 10937, 10940, 10941, 
10943, 10945-10950. Playa Buda, 22? 10.8? 
N, 98? 22.8? W, 10 November 2006 (1, 2
),
R. Segura & M. Mart?nez, LPL 10927, 10929. 
La Marland, 22? 08.4? N, 98? 24.9? W, 9 
November 2006 (1), R. Segura & M. 
Mart?nez, LPL 10896. Veracruz: El Caracol, 
22? 9.6? N, 99? 1.9? W, 10 November 2006 
(1, 1
), R. Segura & M. Mart?nez, LPL 
10605, 10909. Laguna Chairel, 22? 13.4? N, 
97? 52.9? W, 16 May 2006 (1), R. Segura & 
C. Pascacio, LPL 10605.
Rudenia leguminana (Busck) complex 
(Figure 9)
Rudenia leguminana is widely distributed in 
North America, ranging from southern 
Connecticut to Florida, and from the 
midwestern U.S. to California. It is abundant 
in some xeric areas of the southwestern U.S. 
(i.e. Texas, New Mexico, Arizona) where 
Prosopis sp. (mesquite) and/or Acacia spp. 
(both Leguminosae) are the dominant shrubs. 
Biology. Based on specimen label data and 
one published record (i.e., Busck 1907), this 
species has been reared from Acacia
farnesiana (L.) Willd. (USNM), A. glauca
(L.) Moench (USNM), A. novernicosa Isley 
(USNM), Gleditsia japonica Micq. (Busck 
1907), Leucaena pulverulenta (Schltdl.) 
Benth. (USNM), Mimosa aculeaticarpa var. 
biuncifera (Benth.) Barneby (USNM), 
Prosopis glandulosa Torr. (USNM), and 
Senna lindheimeriana (Scheele) H. S. Irwin & 
Barneby (USNM) (all Leguminosae).
Rudenia leguminana was reared from P.
aculeata in Guatemala (Jutiapa), Mexico (San 
Luis Potos?, Oaxaca, and Veracruz),
Nicaragua (Granada, Managua, and 
Matagalpa), and Venezuela (Miranda and 
Anzo?tegui), and from Pithecellobium dulce
(Leguminosae) in Mexico (San Luis Potos?).
On P. aculeata eggs are laid on leaves (rachis 
and pinnules) and tips of other vegetative 
parts. First instars feed inside the rachis. They 
then move to the axil and make a tunnel 
within it. Larvae leave the tunnel at night to 
feed on the pinnules and rachis of leaves. 
Frass can be seen protruding from tunnels 
especially during the mid-life of larvae when 
the tunnel is being actively excavated. 
Pupation usually occurs in tunnels, and when 
adults emerge the exuvium is partly exuded 
from the tunnel. Adults of Rudenia
leguminana emerged from stems (n = 417 
individuals), flowers (n = 90), and pods (n = 
29) of the host.
Host Specificity Trials. The large number of 
individuals of R. leguminana reared from P.
aculeata in the field indicated that this 
herbivore was worthy of further consideration 
as a biocontrol agent, so additional studies 
were conducted. ?No-choice? host specificity 
trials were performed in 2006. Three females 
and 4 males were placed in bags covering tips
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 6
of 6 test plant species and the control, P.
aculeata (Table 1). Successful development 
on 5 non-target hosts resulted: Parkinsonia
praecox (Ruiz & Pav.) J. A. Hawkins, Mimosa
asperata L., Mimosa pigra, Delonix regia,
and Acacia farnesiana. However, oviposition,
development, and adult emergence of R.
leguminana on non-target hosts may have 
been an artifact of the confined conditions of 
the trials. For example, R. leguminana were 
never reared from Mimosa pigra in the field 
despite considerable research on the 
herbivores of this plant over many years in the 
same geographic region.
To better establish the field host range, an
open-field trial was conducted in a plot at La 
Aguada, Veracruz, Mexico. Eleven plant 
species were grown in a plot: P. aculeata,
Mimosa asperata, Mimosa pigra, Mimosa
pudica L., Acacia farnesiana, Caesalpinia
pulcherrima, Calliandra grandiflora (L?Her.)
Benth., Desmanthus virgatus (L.) Willd., Inga
jinicuil G. Don, Leucaena leucocephala, and 
Tamarindus indica (all Leguminosae). Two 
releases of lab-reared adults were made; the 
first consisted of 42 females and 42 males on 
24 July 2006 and the second of 52 females 
and 52 males on 9 August 2006. When 
damage to tips of the plants was observed, the 
terminal area was bagged to capture emerging 
adults. In these trials, R. leguminana
successfully developed only on P. aculeata
(20 from the first trial and 34 from the 
second). Although R. leguminana successfully 
developed on Mimosa asperata, Mimosa 
pigra, and Acacia farnesiana in the no-choice
trials, no feeding or adult emergence was 
detected on these plant species in the open-
field trials. This suggests that if not confined 
with the non-target hosts, R. leguminana may 
not oviposit on them, or at least development 
is far less likely.
Molecular Analyses. Because few native 
tortricids range from the northeastern U.S. to 
Venezuela, it is possible that more than one 
species is concealed within the material 
examined. Razowski (1985) indicated that 
specimens from Sonora, Sinaloa, and Baja 
California have genitalia very similar to ?R.
leguminana and may be conspecific with that 
species,? even though they exhibit some 
external and genital variation. Although some 
variation (in facies and genitalia) is present in 
the material reared from P. aculeata, the 
variation does not co-vary and is not 
concordant with geography. 
Hence, four genetic markers were analyzed to 
see if molecular data could differentiate 
groups within this broad geographic 
distribution. In one data set (Data Set I) two 
mitochondrial markers were examined, a 781-
basepair region of cytochrome c oxidase I 
(COI) and a 685-basepair region of 
cytochrome c oxidase II (COII), and a 531-
basepair region of the nuclear marker 28S 
domain 2 (D2). In Data Set I 5 individuals 
were sampled from Mexico, 3 from 
Nicaragua, 1 from Arizona, and 1 from 
Nebraska. In addition, another data set (Data 
Set II) was analyzed which consisted of the 
barcode region of COI. In Data Set II 29 
individuals of Rudenia leguminana were 
sampled ranging from Virginia to Venezuela, 
and 3 other tortricids as outgroups (Acleris
semipurpurana, Aethes biscana, and Eugnosta
Table 1. Adult emergence of "no choice" host specificity trials 
of Rudenia leguminana from six plant species (as recorded per 
individual plant).
Plant species
Average adult 
emergence
Parkinsonia aculeata 9.1
Parkinsonia praecox 4.7
Mimosa asperata 6.3
Mimosa pigra 1.3
Delonix regia 3.3
Acacia farnesiana 1
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busckana). Evolutionary pairwise distances in 
PAUP* 4.0b10 (Swofford 2002) were 
calculated using a maximum likelihood model 
with parameter estimates from a maximum 
likelihood analysis carried out in Garli 0.951 
(Zwickl 2006) with the GTR+I+G model, 
selected as best by Modeltest 3.7 (Posada and 
Crandall 1998) based on the AIC criterion. A 
tree based on Data Set II was inferred using 
MrBayes 3.1.2 (Huelsenbeck and Ronquist 
2001) with a data set partitioned by codon 
position into 3 subsets, with uninformative 
priors except for the branch length prior, 5 
million generations (burnin=2 million), 4 
chains (1 cold, 3 heated), and 2 concurrent 
runs. For establishing branch length prior, the 
empirical Bayes method was employed, 
calculating mean branch length for the 
maximum likelihood tree from 4 separate 
Garli analyses and using the resulting value 
for creating an exponential distribution for 
this prior. Whether the MCMC chains 
converged was assessed by examining the 
potential scale reduction factors and log-
likelihood plots over time.
Evolutionary distances between individuals in 
both Data Set I and Data Set II strongly 
suggest that these individuals belong to more 
than one species. In Data Set I, evolutionary
distances for individuals from the same 
geographic area are at the level expected for 
conspecifics (below 1% for both 
mitochondrial markers) (Hebert et al. 2003), 
whereas distances for individuals from more 
distant locations show a divergence an order 
of magnitude greater (Tables 2 and 3). There 
is very little variation in D2. Based on the 
analysis of the Data Set II (COI barcode) it 
appears that there are at least three different 
species (Figure 15), with distances within 
species ranging 0.0-2.3% and between species 
4.9-13% (Table 4). Divergence levels of 
greater than 3% frequently are interpreted to 
represent separate species (Hebert et al. 2003;
Hebert et al. 2004). It is odd that one of the 
individuals from Venezuela (VE 1) is about 
5% distant from the three other Venezuelan 
specimens (however, it is possible that this is 
the result of contamination). Also, the origin 
of the four specimens from Mexico (MX 1-4)
that cluster with the Venezuelan outlier is 
uncertain because these specimens are larvae 
that were intercepted at U.S. ports-of-entry
from ?Mexico? on Pithecellobium dulce (i.e. 
the exact point of origin is unknown, although 
it is likely to be northern Mexico).
Although geographic sampling is limited, 
these results strongly suggest that more than 
one species is present in the material 
examined: species A, including individuals 
from the eastern U.S. (e.g. Kansas, Louisiana, 
Mississippi, Texas, and Virginia); species B, 
Table 2. Evolutionary distances among individuals from Data Set I for the COI marker. Distances between individuals from 
the same locality are in bold.
1 2 3 4 5 6 7 8 9 10
1 Nicaragua 1
2 Nicaragua 2 0.10%
3 Nicaragua 3 0.15% 0.05%
4 Mexico 1 2.35% 2.23% 2.29%
5 Mexico 2 2.17% 2.06% 2.11% 0.15%
6 Mexico 3 2.17% 2.06% 2.11% 0.35% 0.20%
7 Mexico 4 1.55% 1.36% 1.36% 0.08% 0.08% 0.17%
8 Mexico 5 1.55% 1.36% 1.36% 0.08% 0.08% 0.17% 0.00%
9 Arizona 2 2.37% 2.25% 2.31% 2.44% 2.26% 2.26% 1.47% 0.01%
10 Nebraska 1 3.16% 2.96% 2.96% 2.65% 2.65% 2.75% 2.56% 2.56% 0.03%
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including individuals from Mexico (Oaxaca 
and Veracruz) and Venezuela; and species C, 
including individuals from southwestern U.S. 
(Arizona and New Mexico) (Figure 15). 
Owing to weak support for the rest of the tree, 
we refrain from drawing conclusions 
regarding the assignment to species of the 
remaining individuals.
Summary of Collecting Localities.
GUATEMALA: Jutiapa: Asunci?n Mita, 
Finca El Platanar, 14? 17.29? N, 89? 33.05? 
W, LPL 10952. El Guayabo, Lago G?ija, 14?
13.9? N, 89? 32? W LPL 10958-967. El
Platanar, El Chamizal, 14? 18.22? N, 89? 
36.96? W LPL 10973, 10977, 10975, 10979-
10980, 10981, 10982, 10983, 10984, 10986. 
El Platanar, Lago G?ija, Laguna del Muerto, 
14? 17.54? N, 89? 34.33? W, LPL 10998-
11000, 100002-11004. MEXICO: Oaxaca: 
Ciudad Ixtepec, Base A?rea Militar No. 2, 16? 
26.55? N, 95? 4.65? W, LPL 10524, 10829. 
Tehuantepec: Alvaro Obreg?n, 16? 20.9? N, 
95? 4.5? W, LPL 9802-9803. Huazantl?n del 
R?o, 16? 12.7? N, 95? 6? W, LPL 9634, 9859, 
10026-10033, 10081-10084, 10159-161,
10241-251, 10277-286, 10498, 10561, 10808, 
11278-281. El Jordan, 16? 22.1? N, 95? 12.8? 
W, LPL 12227-12230, 12441-12444. San Luis 
Potos?: Lago Ajinche, La Marland, 22? 8.46? 
N, 98? 24.98? W, LPL 10861, 10865, 12446. 
Laguna Ajinche, 22? 10.5? N, 98? 21.0? W, 
LPL 12396, 12412, 12415, 12440. Laguna 
Chica, 22? 11.1? N, 98? 23.0? W, LPL 12399, 
12411, 12416. Laguna Chica, Playa Buda, 22? 
10.85? N, 98? 22.88? W, LPL 11129, 11169. 
La Muralla, 22? 11.8? N, 98? 20? W, LPL 
12413, 12419, 12420, 12428, 12450. La 
Muralla, 22? 11.8? N, 98? 20? W, LPL 12418, 
12408, r.f. Pithecellobium dulce. Pozo La Pez 
No. 1, 22? 10.5? N, 98? 23.4? W, LPL 12398, 
12447. Veracruz: Cacalilao, Lazaro C?rdenas, 
22? 15.79? N, 98? 8.73? W, LPL 10656. 
Veracruz: Canoas, 22? 10.1? N, 98? 07.7? W, 
LPL 12207, 12414, 12426, 12451, 12454. 
Tamos, El Caracol, 22? 9.63? N, 98? 1.92? W, 
LPL 11154-157, 11193, 12208, 12224, 12397, 
12409, 12417, 12422, 12457, 12458, 12459. 
Tamos, La Cortadura, 22? 10.98? N, 98? 1.36? 
W, LPL 10849, 10853, 10905, 12410, 12420, 
12427, 12461. NICARAGUA: Granada: La 
Playuela, 12? 2.8? N, 85? 55.3? W, LPL 9816, 
10041-47, 10095. Laguna La Playuela 1, 12? 
1.8? N, 85? 55.3? W, LPL 9734, 9737-9738,
9740, 9749. Laguna La Playuela 2, 12? 1.3? N, 
85? 55.2? W, LPL 9748-9750, 9760. 
Managua: road to San Francisco Libre, km 74, 
12? 22.6? N, 86? 6.7? W, LPL 9861.
Matagalpa: Tecomapa, 12? 37.6? N, 86? 2.2? 
W, LPL 9869-70.  La Playuela, 12? 2? N, 85? 
56.3? W, LPL 9871-81.  Las Guayabas, 12? 
37.7? N, 86? 17? W, LPL 8812, 8814, 8816-
8817, 8819, 8841, 8842. VENEZUELA: 
Table 3. Evolutionary distances between individuals from the 
same area and between groups from more distant areas in Data 
Set I.
Data Set I COI COII D2
Mexico
0.4-
0.8%
0-
0.3% 0%
Nicaragua 0-0.1%
0-
0.3% 0%
Arizona-Mexico
3.6-
4.3%
2.4-
2.6% 0.20%
Arizona-
Nicaragua
3.2-
3.3%
2.9-
3.3% 0.40%
Arizona-
Nebraska N/A 5.90% 0.95%
Mexico-Nebraska N/A
4.2-
4.5% 0.76%
Mexico-Nicaragua
3.32-
4.0%
2.2-
2.7% 0.20%
Nebraska-
Nicaragua N/A
4.8-
5.2% 0.95%
Table 4. Evolutionary distances within and among species A, B, 
C (see Fig. 15) in Data Set II.
Species or Species 
Pair
Evolutionary 
Distance
A 0.0-1.3%
B 0.5-2.3%
C 0.0-0.5%
A-B 10.50%
A-C 13.00%
B-C 4.90%
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 9
Anzoategui, hwy Barcelona-Caracas, km 18, 
10? 3.34? N, 64? 48.31? W, LPL 11079, 
11081, hwy Barcelona-Caracas, km 11, 10? 
2.93? N, 64? 44.56? W, LPL 11083. Miranda, 
La Guira, Mare Abajo, 10? 36.82? N, 67? 
1.23? W, LPL 11045.
Cochylis sp. 
(Figure 10)
Two females of an undetermined species of 
Cochylis were reared from Veracruz, Mexico. 


Figure 15. Bayesian tree based on Data Set II. Numbers below branches are posterior probabilities. Circled clades (A, B, and 
C) are postulated to be different species. Abbreviations are as follows: USPS abbreviation for the U.S. states, Guatemala (GT), 
Mexico (MX), Mexico Oaxaca (MX, OA), Mexico Veracruz (MX, VE), and VE (Venezuela). High quality figures are available 
online.
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 10
The taxonomic disarray of the group and the 
large number of undescribed taxa combine to 
inhibit accurate species-level identification.
Biology. Although most Cochylini worldwide 
feed on Asteraceae (Razowski 1970), there are 
numerous deviations from this pattern, and 
Leguminosae is not an unusual host plant 
family for members of this tribe.
Specimens Examined. MEXICO: Veracruz: 
La Cortadura, 22? 10.9? N, 98? 1.36? W, 14 
April 2007 (1
), R. Segura, T. Heard, M. 
Mart?nez, C. Pascacio, LPL 11152. San Luis 
Potos?: Playa Buda, 22? 9.63? N, 98? 1.92? W, 
14 April 2007 (1
), R. Segura, T. Heard, M. 
Mart?nez, C. Pascacio, LPL 11168.
Ofatulena Heinrich, 1926
As defined by Heinrich (1926), Ofatulena,
included two species: O. duodecemstriata
(Walsingham) and O. luminosa Heinrich. The 
two share a whitish gray, finely striated 
forewing with a distinct ocellar patch in the 
tornal region; as in most Grapholitini, males 
of Ofatulena lack a costal fold. The male 
genitalia (illustrated by Heinrich 1926) have a 
distally swollen cucullus with a row of long, 
stout, flattened, marginal spiniform setae 
along the outer portion of the valva. 
?Enarmonia? jamaicana Walsingham was 
transferred by Razowski to Ofatulena without 
comment in the checklist of Neotropical 
Lepidoptera (Powell et al. 1995). However, 
because it lacks the typical forewing pattern 
and shape, and the male genitalia deviate 
considerably from those of O.
duodecemstriata (Walsingham) and O.
luminosa Heinrich, we remove it from the 
genus and transfer it provisionally to Cydia
(new combination), with which the facies and 
genitalia are more similar. Ofatulena
duodecemstriata usually can be distinguished 
superficially from O. luminosa by its slightly 
greater forewing length and absence of peach 
or orange scaling in the forewing ocellar 
patch. The male genitalia of O.
duodecemstriata have a linear patch of larger 
spiniform setae along the lower edge of the 
valva subapically and an extra, stout spiniform 
seta near the middle of the apical region. In O.
luminosa the setae are more numerous, but 
much finer and thinner, and usually are easily 
dislodged in slide-mounted preparations. 
Specimens reared from Parkinsonia praecox
in Oaxaca, Mexico apparently represent a 
third and undescribed species of Ofatulena.
Based on collection records (USNM) and 
published literature (Heinrich 1926; MacKay 
1959; Woods 1992), Ofatulena has been 
reared only from Leguminosae.
Ofatulena duodecemstriata (Walsingham)
(Figure 11)
Ofatulena duodecemstriata ranges across the 
western United States from California to 
Texas, north to Utah and south into Mexico. 
In addition, a series of this species (n = 6) was 
discovered in the USNM with the following
data: Venezuela, Lara, Puente Torres, 24 km 
E Carora, 10 March 1978, thorn forest, 
blacklight, J. B. Heppner. The latter represent 
a considerable increase in the previously 
documented geographic range of the species. 
Although the Venezuelan specimens are 
considerably smaller than those from North 
America, the genitalia are identical to North 
American specimens.
Biology. The primary larval host in North 
America is mesquite (?mesquite beans?), 
Prosopis sp. (Leguminosae) (Heinrich 1926; 
MacKay 1959). Although there is a specimen 
in the USNM reared from Verbascum thapsus
L. (Scrophulariaceae), this is almost certainly 
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 11
an error. One specimen from P. aculeata was 
reared in Mexico and two in Venezuela.
Specimens Examined. MEXICO: Oaxaca: 
Tehuantepec, 16? 17.5? N, 95? 13.8? W, 5 
February 2002 (1), M. Mart?nez & M. 
Ju?rez, LPL 9622. VENEZUELA: Nueva 
Esparta: Bah?a de Plata, 11? 6.47? N, 63? 
56.8? W, 12 March 2007 (1, 1
), R. Segura 
& C. Pascacio, LPL 11060.
Ofatulena luminosa Heinrich 
(Figure 12)
This species is recorded from California 
(EME), Arizona (Woods 1992), and Texas 
(USNM, ANIC), USA, and Sonora, Sinaloa, 
and Nuevo Leon, Mexico (ANIC). Larvae 
have been reported previously only from P.
aculeata (Woods 1992). According to Woods 
(1992), ?Up to 5 larvae have been dissected 
from the distal 25 cm of a stem [of P.
aculeata]. A single larva may also develop in 
a green seed, eating out the seed and killing it. 
Heavily infested plants appear to grow at a 
slower rate than uninfested ones.? Woods 
(1992) also mentions that the larvae of O.
luminosa are heavily parasitized by 
Hymenoptera.
Biology. Specimens of O. luminosa were
reared from P. aculeata in Oaxaca, San Luis 
Potos?, and Veracruz, Mexico and from P.
texana in San Luis Potos?. The abundance of 
individuals in the field and the damage it 
causes indicated that this herbivore could have 
potential as a biocontrol agent, so additional 
observations and studies were conducted.
Larvae bore in leaf and stem tips, mature 
green stems, and green seeds. The stem may 
appear swollen as a result of the presence of 
the larvae. Most damage is concentrated in the 
pith in the center of the stems. When mature, 
the larva prepares an exit hole distinguished
by a frass and silk structure. From 218 bagged 
stems which showed evidence of damage, 96 
adults emerged. Two to 3 damaged stems 
were observed per plant on 2 transects 
showing that this insect is consistently and 
abundantly available (Table 5).  Overall, 
adults of Ofatulena luminosa emerged from 
stems (n = 164 individuals), pods (n = 55), 
and flowers (n = 2) of the host. Adults live for 
a mean of 6 days. 
Surveys of Natural Host Plant Use. Twenty-
three legume species growing in the same 
habitat as P. aculeata were recognized (5 
species of Caesalpinioideae, 11 species of 
Mimosoideae, and 7 species of Faboideae) 
(Table 6). Stems with evidence of damage 
were bagged for adult emergence. In October 
2008 when the site was threatened with 
flooding, all stems were collected and taken to 
the lab where they were held in plastic bottles 
until insects emerged, or were dissected if the 
stem showed signs of drying out. All insects 
that emerged were pinned, labeled, and 
identified. Ofatulena luminosa emerged only
from Parkinsonia aculeata and P. texana (A. 
Gray) S. Watson var. macra (I.M. Johnst.) 
Isely.
Table 5. Density of O. luminosa larval damage in the field at 
La Muralla and El Caracol.
Sep-08 Jan-09
La Muralla branches/trees 6.5 6.5
La Muralla
Ofatulena % 
damage 45.7 22.1
El Caracol branches/trees 8.6 8.6
El Caracol
Ofatulena % 
damage 26.5 18.8
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 12
Specimens Examined. MEXICO: Oaxaca: 
Comitancillo, 16? 28.8? N, 95? 5.5? W, 18 
August 2009 (1
), M. Mart?nez & C. 
Pascacio, LPL 12537. Huazantlan del Rio, 16?
12.7? N, 95? 6? W, 19 August 2009 (6, 8
),
M. Mart?nez & C. Pascacio, LPL 12546. San 
Jos? del Palmar, 16? 13.84? N, 95? 10.77? W, 
19 August 2009 (1), M. Mart?nez & C. 
Pascacio, LPL 12550. San Luis Potos?:
Laguna Ajinch?, 22? 10.5? N, 98? 21.0? W, 18 
May 2006 (12, 5
), R. Segura & C. 
Pascacio, LPL 10681, 10690, 10691, 23 April 
2009 (1, 1
), M. Mart?nez & C. Pascacio, 
LPL 12404, 12464. Ajinch?, 22? 11.8? N, 98?
21.1? W, 16 May 2006 (1, 1
), R. Segura & 
C. Pascacio, LPL 10644. Laguna Chica, 22?
11.1? N, 98? 23.0? W, 19 May 2008 (1, 1
),
M. Mart?nez, LPL 11725, 11742, 21 April 
2009 (1), R. Segura, M. Mart?nez, C. 
Pascacio, T. Heard, LPL 12424. La Marland, 
22? 11.8? N, 98? 24.9? W, 8 November 2006 
(1
), Segura & Mart?nez, LPL 10866, 13 
April 2007 (3, 2
), R. Segura, M. Mart?nez,
T. Heard, C. Pascacio, LPL 11147, 11148, 21 
April 2009 (1
), Segura, Mart?nez, Pascacio 
& Heard, LPL 12423. La Marland 2, 22? 08.4?
N, 98? 24.9? W, 13 January 2009 (1), M. 
Mart?nez & C. Pascacio, LPL 12219. Pozo La 
Pez No. 1, 22? 10.5? N, 98? 23.4? W, 13 April 
2007 (1, 1
), Segura, Mart?nez, Pascacio & 
Heard, LPL 11118, 21 April 2009 (1
),
Segura, Mart?nez, Pascacio & Heard, LPL 
12403. La Muralla, 22? 11.8? N, 98? 20? W, 
18 May 2008 (5 specimens), LPL 11683, 
11684, 11685, 11721, 11722, 21 May 2008 (5 
specimens), LPL 11687, 11688, 11689, 
11690, 11744, 13 January 2009 (1
), M. 
Mart?nez & C. Pascacio, LPL 12212, 22 April 
2009 (1), M. Mart?nez & C. Pascacio, LPL 
12462, 21 April 2009 (1, 1
), Segura, 
Mart?nez, Pascacio & Heard, LPL 12402, 
12449. Veracruz: Canoas, 22? 10.1? N, 98? 
07.7? W, 10 November 2006 (1
), Segura & 
Mart?nez, LPL 10678, 10 November 2006 
(1
), R. Segura & C. Pascacio, LPL 10919, 
16 August 2008 (1), Mart?nez, LPL 12191,
22 April 2009 (2, 1
), Segura, Mart?nez, 
Pascacio & Heard, LPL 12401, 12452, 12455, 
14 September 2008 (1
), C. Pascacio, LPL 
12197, 17 September 2008 (1, 1
), Segura 
& Pascacio, LPL 12200. Ciudad Cuauhtemoc, 
22? 12.8? N, 97? 53.8? W, 17 May 2006 (2),
R. Segura & C. Pascacio, LPL 10663. El 
Caracol, 22? 09.6? N, 98? 01.9? W, 10 
November 2006 (3, 3
), R. Segura & M. 
Mart?nez, LPL 10913, 10910, 10911, 22 April 
2009 (2, 1
), Segura, Mart?nez, Pascacio & 
Heard, LPL 12456, 12460. El Caracol, 22? 
09.3? N, 98? 02.4? W, 16 August 2008 (1),
M. Mart?nez & C. Pascacio, LPL 12192. La 
Cortadura, 22? 10.9? N, 98? 01.3? W, 8 
Table 6. Related legumes growing in the same habitat as 
Parkinsonia.
Plant species Family Site
Acacia cornigera Mimosaceae La Muralla
Acacia cornigera Mimosaceae Canoas
Acacia farnesiana Mimosaceae La Muralla
Enterolobium 
cyclocarpum Mimosaceae La Muralla
Leucaena glauca Mimosaceae La Muralla
Mimosa asperata Mimosaceae El Caracol
Mimosa asperata Mimosaceae Canoas
Pithecellobium dulce Mimosaceae El Caracol
Pithecellobium ebano Mimosaceae La Muralla
Prosopis laevigata Mimosaceae La Muralla
Prosopis laevigata Mimosaceae Canoas
Albizia lebbeck Faboideae Ebano (urban)
Delonix regia Faboideae La Muralla
Desmanthus virgatus Faboideae Canoas
Erythrina herbacea Faboideae La Muralla
Piscidia piscipula Faboideae La Muralla
Sesbania sp. Faboideae La Muralla
Vigna speciosa Faboideae La Muralla
Bauhinia purpurea Caesalpiniaceae Ebano (urban)
Caesalpinia mexicana Caesalpiniaceae La Muralla
Caesalpinia sp. Caesalpiniaceae La Muralla
Parkinsonia texana Caesalpiniaceae La Muralla
Tamarindus indica Caesalpiniaceae La Muralla
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 13
November 2006 (1), R. Segura & M. 
Mart?nez, LPL 10855, 22 April 2009 (1
),
Segura, Mart?nez, Pascacio & Heard, LPL 
12425. Cardenas, 22? 15.7? N, 98? 8.73? W, 
17 May 2006 (1
), R. Segura & M. Mart?nez, 
LPL 10650.
Ofatulena sp. 
(Figure 13)
Two specimens of Ofatulena were reared 
from Parkinsonia praecox in Oaxaca, Mexico. 
The adults have a forewing length 
conspicuously less than that of either O.
duodecemstriata or O. luminosa, the forewing 
has slightly darker scaling, and the third 
segment of the labial palpus is brown (white 
in the other two species). In the male genitalia 
(n = 1), the swollen portion of the valva is less 
inflated, and the flattened scales in a row 
along the perimeter of the valva are somewhat 
uniformly spaced and easily dislodged. The
two specimens likely represent an undescribed 
species.
Biology. Apart from the host, nothing is 
known of the biology of this apparently 
undescribed species.
Specimens Examined. MEXICO: Oaxaca: El 
Jordan, 16? 24.3? N, 98? 11.6? W, 21 January 
2009 (1, 1
), M. Mart?nez & C. Pascacio, 
LPL 12435, 12231.
Crocidosema lantana Busck 
(Figure 14)
Crocidosema lantana is indigenous to 
Mexico; it is recorded from Florida to Costa 
Rica. It was described from Hawaii (Busck 
1910) based on specimens imported from 
Mexico for biological control of weedy 


Figure 9-14. Adults of leaf-roller moths reared from Parkinsonia aculeata. 9) Rudenia leguminana (Busck). 10) Cochylis sp. 11) 
Ofatulena duodecemstriata (Walsingham). 12) Ofatulena luminosa Heinrich. 13) Ofatulena n. sp. 14) Crocidosema lantana Busck. 
Scale bar = 5 mm. High quality figures are available online.
Journal of Insect Science: Vol. 11 | Article 7 Brown et al.
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 14
lantana (Lanana camara L.: Verbenaceae) 
(Perkins and Sweezy 1924; Zimmerman 
1978). It subsequently was introduced into 
Australia (Tyron 1914; Common 1957) and 
Miocronesia (Zimmerman 1978) for 
biological control.
Biology. This species has been reared from 
Tabebuia chrysantha (Jacq.) G. Nicholson 
(Bignoniaceae), Tecoma stans (L.) Juss. ex 
Kunth (Bignoniaceae), Lantana camara L. 
(Verbenaceae), and Litchi chinensis Sonn. 
(Sapindaceae) (Busck 1910; Tyron 1914; 
Perkins & Swezey 1924; Common 1957; 
MacKay 1959; Kimball 1965; Clarke 1976; 
Zimmerman 1978, Ibrahim & Zakaria 1988; 
Muniappan 1990). A single female of C.
lantana was reared from Parkinsonina
aculeata in San Luis Potos?, Mexico.
Specimen Examined. MEXICO: San Luis 
Potos?: La Muralla, 22? 11.8? N, 98? 20? W, 
13 January 2009 (1
), M. Martinez & C. 
Pascacio, LPL 12210.
Concluding Remarks
The ten species of Tortricidae reared from P.
aculeata in this study show a range of feeding 
modes from leaf rollers to borers in stems and 
reproductive parts. Their systematic positions 
vary from clearly defined species to possible 
species complexes, and include members of 
both of the largest subfamilies ? Tortricinae
and Olethreutinae. Their host specificity also 
varies widely from extreme generalists to 
possible specialists on the genus Parkinsonia.
Although 7 of the 10 species are clearly not 
suitable as biocontrol agents, 2 species show 
some potential. Rudenia leguminana is 
probably a complex of species, one or more of 
which may be a specialist. Further work is 
needed to understand the systematics of this 
group. Ofatulena leguminosa is possibly a 
specialist on the genus Parkinsonia. This level 
of specificity may be suitable for the 
importation of this species into Australia as no 
other species of Parkinsonia occur there. 
Further work is ongoing to determine its host 
specificity and its impact on the target plant.
Acknowledgements
We thank Moises Mart?nez and Carlos 
Pascacio (both CSIRO) for assistance with 
field work in Mexico. Diane Hartley (CSIRO) 
provided sequence data. The following 
provided helpful reviews of the manuscript: 
Natalia Vandenberg and Thomas Henry, 
Systematic Entomology Laboratory, U.S. 
Department of Agricultural, National Museum 
of Natural History, Washington, D.C.; and 
John Heppner, Florida State Collection of 
Arthropods, Gainesville, Florida. The
Australian Government Department of 
Agriculture, Fisheries and Forestry provided 
financial support.
Editor?s note: Paper copies of this article 
will be deposited in the following libraries. 
The date of publication is given in ?About the 
Journal? on the JIS website.
Universitaetsbibliothek Johann Christian
Senckenberg, Frankfurt Germany; National 
Museum of Natural History, Paris, France; 
Field Museum of Natural History, Chicago, 
Illinois USA; University of Wisconsin, 
Madison, USA; University of Arizona, 
Tucson, Arizona USA; Smithsonian 
Institution Libraries, Washington D.C. USA; 
The Linnean Society, London, England.
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