Host Records for Tortricidae (Lepidoptera) Reared from Seeds and Fruits in a Thailand Rainforest

Home , Archipini, Archips, Chlidanotinae, Cryptaspasma, Cryptophlebia, Cryptophlebia illepida

PROC. ENTOMOL. SOC. WASH. 121(4), 2019, pp. 544–556

HOST RECORDS FOR TORTRICIDAE (LEPIDOPTERA) REARED FROM SEEDS AND FRUITS IN A THAILAND RAINFOREST

JOHN W. BROWN,YVES BASSET,MONTARIKA PANMENG,SUTIPUN PUTNAUL, AND SCOTT E. MILLER

(JWB) Department of Entomology, National Museum of Natural History, P.O. Box 37012, Washington, DC 20013-7012 (e-mail: [email protected]); (YB) ForestGEO Smithsonian Tropical Research Institute, Apartado 0843-03092, Balboa, Ancon, Panama City, Republic of Panama (e-mail: [email protected]); (MP, SP) ForestGEO Arthropod Laboratory, Khao Chong Botanical Garden, Na Yong, Thailand ([email protected]); (SEM) Department of Entomology, National Museum of Natural History, P.O. Box 37012, Washington, DC 20013-7012 (e-mail: [email protected])

Abstract.—A survey of insects reared from seeds and fruits in a rainforest in Thailand yielded 337 specimens of Tortricidae representing 16 species. Based on this material, we present host records for the following: Hilarographa muluana Razowski complex (Chlidanotini), Archips machlopis (Meyrick) (Archipini), Cryptaspasma brachyptycha (Meyrick) (Microcorsini), Collogenes squamosa (Diakonoff) (Microcorsini), Gatesclarkeana idia Diakonoff (Olethreutini), Heli- ctophanes prospera (Meyrick) (Olethreutini), Lobesia aelopa (Meyrick) (Oleth- reutini), Hoplitendemis sp. A (Olethreutini), Cryptophlebia rhynchias (Meyrick) (Grapholitini), Cryptophlebia ombrodelta (Lower), Cryptophlebia sp. (undetermined) (Grapholitini), Thaumatotibia sp. (undetermined) (Grapholitini), An- drioplecta shoreae Komai (Grapholitini), Andrioplecta subpulverula (Obraztsov) (Grapholitini), Andrioplecta (?) species (undetermined) (Grapholitini), and Cydia (?) species (undetermined) (Grapholitini). Tortricids were reared from 30 plant species representing 12 plant families, with Sapindaceae and Annonaceae sup- porting the greatest number of species, seven and six, respectively. Consistent with other surveys of seed- and fruit-feeding tortricids, the tribe Grapholitini represented 50% of the total tortricid species and 73% of the total tortricid specimens in the Thailand survey. Key Words: Andrioplecta, Annonaceae, barcodes, Cryptophlebia, Dipterocarpaceae, Fabaceae, Grapholitini, Hilarographa, Indo-Australia, Sapindaceae DOI: 10.4289/0013-8797.121.4.544

Large-scale insect rearing projects et al. 2002, Sam et al. 2017), Ecuador conducted in various tropical sites (e.g., Bodner et al. 2010, 2012), Costa throughout the world, including Kenya Rica (e.g., Janzen and Hallwachs 2009, (e.g., Copeland et al. 2002, 2009; Miller 2016), Thailand (e.g., Basset et al. et al. 2014), New Guinea (e.g., Novotny 2018), Panama (e.g., Gripenberg et al.

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2019), and others, have contributed a seminatural conditions in pots lined with remarkable amount of new host plant tissue paper and covered with very ﬁne data for many groups of phytophagous netting for ventilation and to avoid sub- insects including the large, and eco- sequent colonization and/or contamina- nomically important lepidopteran family tion. The pots were checked twice a Tortricidae (e.g., Brown et al. 2014). week, and any emerging insects were During a recent three-year survey of collected, preserved, and pinned. Seeds seed- and fruit-feeding insects in Thai- and fruits were kept for three months and land (Basset et al. 2019), 1,970 sam- then dissected to determine the presence ples were collected representing 39,252 of developing larvae. Those with live seeds or fruits from 357 liana and tree larvae were retained for an additional species (and a few herbs) representing rearing period, and the remaining were 66 plant families. Among the insects discarded. reared were 337 specimens of tortricid DNA Barcoding.—Tissue samples (one moths, representing 16 species. The leg of an adult moth) were used to am- purpose of this contribution is to present plify a ;650bp region of the mitochon- host records for these tortricids in the drial gene cytochrome oxidase I (COI), context of previously recorded larval commonly referred to as the DNA bar- food plants. code, using standard Sanger sequencing procedures employed at the Biodiver- MATERIALS AND METHODS sity Institute of Ontario, University of Study site.—The survey was con- Guelph (Hebert et al. 2003, Craft et al. ducted on a 24-ha permanent vegetation 2010, Wilson 2012). Barcode Index plot at Khao Chong Botanical Garden Numbers (BINs) using the RESL algo- (KHC) and surrounding habitat, located rithm were used to help delineate spe- in lowland seasonal evergreen rainforest cies (Ratnasingham and Hebert 2013). of the Khao Ban Thad Wildlife Sanctuary Unfortunately, many of the specimens in southern Thailand. The site supports at yielded poor or incomplete sequence least 593 tree species, representing 285 data owing to their exposure to high genera and 82 families. humidity. Next-generation sequencing Sampling and rearing.—Methods of (NGS) (Hebert et al. 2018) yielded sig- seed and fruit collection and insect niﬁcantly improved results for a small rearing are detailed by Basset et al. sample of these. (2019) and can be summarized as fol- Depositories.—Vouchers of adults are lows. In 2013, seeds and fruits were deposited in the Thai Department of collected from locally abundant tree, National Parks, Wildlife and Plant Con- shrub, and liana (more rarely herb) spe- servation, Bangkok, Thailand, and the cies. During 2014 and 2015, sampling National Museum of Natural History, was restricted to the 10 most common Smithsonian Institution, Washington, D.C., plant families at KHC: Annonaceae, USA. Arecaceae, Ebenaceae, Euphorbiaceae, Fabaceae, Lauraceae, Meliaceae, Phyl- RESULTS lanthaceae, Rubiaceae and Sapindaceae. The survey yielded 337 specimens of Seeds and fruits still on plants or freshly Tortricidae representing 16 species. We fallen (without apparent decomposition) recorded tortricids from 30 plant species were collected and brought into the representing 12 plant families (Table 1). laboratory where they were kept under The reared species represent only a small

percentage of the tortricid fauna of the (K. Nishida, personal communication), site, but likely constitute a reasonable and Loganiaceae and Rubiaceae in sample of the fruit- and seed-eating Papua New Guinea (Sam et al. 2017). members of that fauna. Below we pres- This fairly wide range of host families ent host records for all specimens that provides little evidence of host speci- could be identified at least to tribe. ficity at the tribal or generic levels; Using Sanger sequencing we obtained the only feature these records have in barcodes for only three species: Hilar- common is the internal feeding habit. ographa muluana complex (n = 5), During the study in Thailand, 44 Andrioplecta shoreae (n = 7), and specimens of Hilarographa muluana Cryptophlebia rhynchias (n = 7). Using Razowski complex were reared from NGS we were able to add two more three different hosts: Pseuduvaria ru- species: Hoplitendemis sp. A (n = 2) and gosa (n = 40), Mezzettia parviflora (n = Helictophanes prospera (n = 1) (Table 3), and Artabotrys sp. (n = 1) (all An- 2). Full specimen data (including host nonaceae). plants) for specimens sequenced (including The adult and male genitalia of our those that failed) and images are avail- specimens are identical to those illus- able on BOLD (www.boldsystems.org), trated by Razowski (2009: figs. 29, 30, accessible from the dataset KHCFRUIT 80) for Hilarographa muluana, which using a DOI (dx.doi.org/10.5883/DS- he described from Sarawak. They are KHCFRUIT). Barcode data are depos- also identical to those of a species of ited in GenBank (Table 2). Hilarographa reared from fruit in Papua New Guinea, listed by Sam et al. (2017) as Hilarographa nr muluana. However, Chlidanotinae: Hilarographini barcodes show a 3.2% difference be- Hilarographa muluana Razowski tween the specimens from Thailand and complex those from Papua New Guinea; hence, it Hilarographini is a small tribe in the is uncertain which of these species, if primitive tortricid subfamily Chlidano- either, is actually H. muluana. There- tinae (Regier et al. 2012). The tribe is fore, we refer to all of them as “H. primarily pantropical in distribution muluana complex.” Specimens from (Heppner 1982), and its foodplants are Papua New Guinea were reared from the poorly known. Species of Thaumatog- fruit of Phrynium macrocephalum and rapha are reported to be internal borers P. pedunculatum (Marantaceae) (Sam in the stems and cambium of several et al. 2017). plant families, including Lauraceae (Diakonoff 1982), Myrsinaceae (Diakonoff Tortricinae: Archipini and Arita 1981), Pinaceae (Diakonoff and Archips machlopis (Meyrick) Arita 1981, Heppner 1982, Brown et al. 2008), Rubiaceae (Heppner 1982), and Archipini is one of the largest tribes in Zingiberaceae (Heppner 1982). Idiot- Tortricidae with over 2000 described hauma near africanum Walsingham was species distributed worldwide (Brown reared from the fruit of Maesa lanceo- 2005, Regier et al. 2012). The vast ma- lata (Myrsinaceae) (Brown et al. 2014) jority of Archipini are polyphagous leaf in Kenya, and undetermined species of rollers, reported from numerous plant Hilarographa have been reared from families. The occurrence of Archipini on Costus sp. (Costaceae) in Costa Rica fruit is usually “opportunistic,” as larvae

Table 1. Herbivore utilization by host plant.

FAMILY Plant species Hervibvore # of specimens

ANNONACEAE Artabotrys sp. Hilarographa muluana 3 Mezzettia parviflora Hilarographa muluana 3 Polyalthia cauliflora Gatesclarkekana idia 1 Pseuduvaria rugosa Hilarographa muluana 40 Platymitra siamensis Hoplitendemis species A 1 Andrioplecta shoreae 1 unidentified Annonaceae sp. 14 Collogenes squamosa 1 Hoplitendemis species 6 Andrioplecta (?) sp. 1 ARECACEAE Arenga pinnata Cryptophlebia rhynchias 1 Calamus godefroyi Andrioplecta shoreae 1 Wodyetia bifurcata Cryptophlebia species 1 CHRYSOBALANACEAE Maranthes corymbosa Hoplitendemis species A 5 DIPTEROCARPACEAE Dipterocarpus grandiflorus Andrioplecta shoreae 4 Andrioplecta subpulverula 1 Dipterocarpus sp. Andrioplecta shoreae 1 Parashorea stellata Hoplitendemis species A 1 Andrioplecta shoreae 47 Shorea roxburghii Andrioplecta shoreae 9 Shorea hypochra Andrioplecta shoreae 3 EUPHORBIACEAE Macaranga denticulata Andrioplecta shoreae 3 Ptychopyxis sp. 1 Thaumatotibia species 1 Andrioplecta shoreae 1 FABACEAE Entada sp. 1 Cryptophlebia rhynchias 59 Millettia atropurpurea Cryptophlebia rhynchias 81 Andrioplecta shoreae 1 Parkia speciosa Cryptophlebia species 1 LAURACEAE unidentified Lauraceae sp. 1 Cryptaspasma brachyptycha 2 Collogenes squamosa 6 Cryptophlebia rhynchias 2 Andrioplecta (?) species 7 MELIACEAE Aglaia sp. 14 Andrioplecta shoreae 1 Synoum glandulosum Helictophanes prospera 1 MYRTACEAE Syzygium sp. 8 Archips machlopis 1 Syzygium sp. 3 Hoplitendemis species A 17 PHYLLANTHACEAE Baccaurea polyneura Hoplitendemis species A 1

Table 1. Continued.

FAMILY Plant species Hervibvore # of specimens

RHAMNACEAE Ziziphus angustifolius Hoplitendemis species A 1 SAPINDACEAE Lepisanthes rubinigosa Lobesia aelopa 5 Cryptophlebia ombrodelta 5 Cryptophlebia species 6 Nephelium lappaceum Hoplitendemis species A 1 Xerospermum noronhianum Gatesclarkeana idia 1 Andrioplecta shoreae 1 Thaumatotibia species 1 UNKNOWN FAMILY Unknown host Cryptophlebia rhynchias 6

feeding on leaves may also feed on ad- Convolvulaceae, Fabaceae, Lauraceae, jacent fruit. Liliaceae, Malvaceae, Marantaceae, In the Thailand study a single speci- Oleaceae, Oxalidaceae, Rubiaceae, men of the widespread Archips machlopis Rutaceae, Sapindaceae, Theaceae, and (Meyrick) (Tuck 1990) was reared from few others. Other authors (e.g., Tuck Syzygium species 8 (Myrtaceae). Yu- 1990, Bhumannavar et al. 1991, Kuroko nus and Ho (1980) reported this moth and Lewvanich 1993) reported many of species from Apiaceae, Bombaceae, the same host families.

Table 2. Specimens successfully sequenced using Sanger and NGS (*) sequencing.

Species sequence code # of bp BIN GenBank #

Hilarographa muluana complex KHCSP062-16 603 [0n] ADA7980 MN036536 Hilarographa muluana compex KHCSP060-16 668 [0n] ADA7980 MN036538 Hilarographa muluana complex KHCSP061-16 668 [0n] ADA7980 MN036542 Hilarographa muluana complex KHCSP068-16 624 [0n] ADA7980 MN036540 Hilarographa muluana complex KHCSP063-16 651 [0n] ADA7980 MN036548 *Hoplitendemis n. sp. LNAUW4057-18 658 [200n] MN036551 *Hoplitendemis n. sp. LNAUW4058-18 658 [380n] MN036546 *Helictophanes prospera LNAUW4055-18 540 [200n] MN036549 Andrioplecta shoreae KHCSP101-16 622 [0n] ACG1185 MN036543 Andrioplecta shoreae KHCSP070-16 617 [0n] ACG1185 MN036550 Andrioplecta shoreae KHCSP079-16 675 [0n] ACG1185 MN036541 Andrioplecta shoreae KHCSP071-16 675 [0n] ACG1185 MN036535 Andrioplecta shoreae KHCSP085-16 678 [0n] ACG1185 MN036547 Andrioplecta shoreae KHCSP087-16 680 [0n] ACG1185 MN036539 Andrioplecta shoreae KHCSP082-16 683 [0n] ACG1185 MN036555 Cryptophlebia rhynchias KHCSP077-16 674 [0n] AAV7917 MN036553 Cryptophlebia rhynchias KHCSP074-16 675 [0n] AAV7917 MN036552 Cryptophlebia rhynchias KHCSP076-16 676 [0n] AAV7917 MN036537 Cryptophlebia rhynchias KHCSP073-16 677 [0n] AAV7917 MN036554 Cryptophlebia rhynchias KHCSP098-16 677 [0n] AAV7917 MN036544 Cryptophlebia rhynchias KHCSP078-16 677 [0n] AAV7917 MN036556 Cryptophlebia rhynchias KHCSP099-16 678 [0n] AAV7917 MN036544

Olethreutinae: Microcorsini suggest a preference for Lauraceae Cryptaspasma and Collogenes (Laurales) and Myrtaceae (Myrtales). Although in different plant orders, trees Microcorsini includes two genera: the in these two families are often superfi- pantropical Cryptaspasma Walsingham cially similar. (37 species) and the Indo-Australian We reared seven specimens of Collo- Collogenes Meyrick (seven species). The genes squamosa (Diakonoff) from two few species of Cryptaspasma for which different hosts: unidentified Lauraceae larval hosts have been recorded feed on sp. 1 (n = 6) and unidentified Annona- the hard kernels or large seeds of a variety ceae sp. 14 (n = 1). of different plant families (Brown and Brown 2004, Dugdale et al. 2005, Horak 2006, Gilligan et al. 2011), suggesting Olethreutinae: Olethreutini that seed morphology rather than plant Gatesclarkeana idia Diakonoff chemistry determines host choice. At Gatesclarkeana includes ten described least two species have apparently ex- species restricted to the Indo-Australian panded their host range from native to region, with host plants recorded for four of cultivated plants and are now considered the ten. Gatesclarkeana domestica has been pests: C. querula (Meyrick) on Syzygium reared from Fabaceae, Euphorbiaceae, (Myrtaceae) and stone fruits (Rosaceae) Lauraceae, Theaceae, and Urticaceae in New Zealand (Dugdale et al. 2005) and (Diakonoff 1973); G. erotias from C. perseana Gilligan and Brown on av- Anacardiaceae, Boraginaceae, Fabaceae, ocado (Persea americana;Lauraceae)in Loranthaceae, Oxalidaceae, Sterculiaceae, Central America (Gilligan et al. 2011). In Sapindaceae, and Verbenaceae (Fletcher Kenya, four species of Cryptaspasma 1921, 1932, Simon Thomas 1962, Dia- were reared from fruit of native Apoc- konoff 1971, 1982); G. idia Diakonoff ynaceae, Clusiaceae, Connaraceae, Eu- from Euphorbiaceae, Lamiaceae, phorbiaceae, Myrtaceae, Podocarpaceae, Malvaceae, Myrtaceae, Theaceae, and and Rosaceae, with little evidence of host Verbenaceae (Diakonoff 1973, Kawabe fidelity within species (Brown et al. 1989); and G. senior from Euphorbia- 2014). Horak (2006) reported Collogenes ceae, Fabaceae, Theaceae, and Verbe- loricata from Endiandra sieberi (Laur- naceae (Diakonoff 1968, 1973). In aceae) in Australia. Thailand, G. idia (n = 2) was reared from Two specimens of a species of Crypt- Polyalthia cauliflora (Annonaceae) (n = aspasma that appears to be C. brachyp- 1) and Xerospermum noronhianum (Sa- tycha (Meyrick) were reared from pindaceae) (n = 1). Species in this genus unidentified Lauraceae species 1. The show little or no host specificity at the adults are in very poor condition, but the plant family level. male and female genitalia are very similar to those illustrated by Diakonoff (1959) and Horak (2006) for this species. Helictophanes prospera (Meyrick) Cryptaspasma brachyptycha was re- Helictophanes includes nine described ported by Sam et al. (2017) from the fruit species ranging from India to Thailand, of Cryptocarya depressa (Lauraceae), south to Australia and east to New Ca- Syzygium amplum (Myrtaceae), and Sy- ledonia, Guam, and Fiji (Brown 2005). zygium trivene (Myrtaceae) in Papua Sam et al. (2017) provided numerous New Guinea. These combined records records for six undetermined species of

Helictophanes from Papua New Guinea, host, Lepisanthes rubinigosa (Sapinda- where the vast majority were reared ceae). from the fruit of Meliaceae, with many fewer records from Malvaceae, Lam- Hoplitendemis species A iaceae, and Euphorbiaceae. Citing records from Turner (1946), Horak (2006) Hoplitendemis is a small genus (four mentioned that three species have been described species) that is restricted to reared in Australia, once each from Southeast Asia, ranging from Vietnam Apocynaceae, Euphorbiaceae, and Me- and Thailand to Java and the Celebes liaceae. Most of these plant families are (Brown 2005). The only published life recognized for their considerable phar- history information was presented by macological properties. During the Diakonoff (1973), who described H. Thailand survey, one specimen of H. erebodes from a small series of adults prospera was reared from Synoum reared from psyllid galls on leaves of glandulosum (Meliaceae). Syzygium (probably S. polyanthum) Although male genitalia are virtu- (Myrtaceae). ally indistinguishable among specimens During the survey in Thailand, an identified as H. prospera, barcodes form undescribed species of Hoplitendemis (n two distinct clusters that almost certainly = 33) was reared from the fruit of eight represent two species (>4% difference). different hosts: Syzygium species 3 (n = Our Thailand specimen clusters with one 17) (Myrtaceae), unidentified Annona- individual from Vietnam and one from ceae species 14 (n = 6), Maranthes cor- Australia. The second cluster includes ymbosa (Chrysobalanaceae) (n = 5), two specimens from Papua New Guinea Baccaurea polyneura (Phyllanthaceae) and two from Australia. We are uncertain (n = 1), Nephelium lappaceum (Sa- which cluster is conspecific with the ho- pindaceae) (n = 1), Parashorea stellata lotype of prospera, that is from India. (Dipterocarpaceae) (n = 1), Platymitra siamensis (Annonaceae) (n =1), and Ziziphus angustifolius (Rhamnaceae) Lobesia aelopa (Meyrick) (n = 1). The male genitalia of our adults Lobesia is nearly worldwide in dis- match perfectly those illustrated by tribution, occurring on all continents Pinkaew (2006) for “Hoplitendemis sp. except Antarctica (Brown 2005). The A” from Thailand. Like Andrioplecta majority of the species for which host (discussed below), it appears that the plants are known appear to be polyph- larvae of Hoplitendemis may be pre- agous (e.g., Nasu, 1993, Royals et al. daceous or phytophagous, potentially a 2018). For example, Lobesia aelopa species-specific life history feature. has been recorded from Actinidiaceae, Apocynaceae, Aquifoliaceae. Asteraceae, Olethreutinae: Grapholitini Ebenaceae, Euphorbiaceae, Fabaceae, Cryptophlebia species Lamiaceae, Malvaceae, Myrtaceae, Rosaceae, Rubiaceae, Rutaceae, Sa- Cryptophlebia is a large genus (55 pindaceae, Sterculiaceae, Theaceae, described species) with a pantropical Ulmaceae, and Vitaceae (e.g., Diakonoff distribution, occurring in Asia, Africa, 1982, Kodama 1988, Bae and Komai Australia, and South America, and on 1991, Nasu 1993). In Thailand, Lobesia many Pacific Islands (Komai 1999). The aelopa (n = 5) was reared from a single genus includes several notorious pests of

fruit and nuts including the koa seed- (Fabaceae) (n = 1). Our records agree worm (Cryptophlebia illepida (Butler)) with previously documented findings - and the macadamia or litchi nutborer species of Cryptophlebia have a strong (Cryptophlebia ombrodelta (Lower)). preference for Fabaceae but also feed on Host plants encompass numerous species the fruit of a wide range of plant families. in about 15 fruit- and nut-bearing families, with a preponderance of Fabaceae. Thaumatotibia species During the survey in Thailand, Cryp- tophlebia rhynchias (Meyrick) (n = 149) Thaumatotibia includes about 25 de- was reared from five different hosts: scribed species occurring primarily in Millettia atropurpurea (Fabaceae) (n = the tropical regions of Asia, Australia, 81), Entada species 1 (Fabaceae) (n = and Africa (Komai 1999). The genus 59), Arenga pinnata (Arecaceae) (n = 1), includes a number of fruit pests, undetermined Lauraceae species 1 (n = most notably the false codling moth, 2), and an unknown host (n = 6). Cryp- Thaumatotibia leucotreta (Meyrick). tophlebia rhynchias was the most com- Whereas most species of Thaumatotibia monly reared tortricid during the survey. appear to be fairly host specific, at least It previously was reported from Faba- at the family level, T. leucotreta has a ceae (Meyrick 1912, Clarke 1976, host range that encompasses 23 different Ghesquiere 1940) with a single record plant families in Kenya alone (Brown on Rosaceae from the rearing files of the et al. 2014). late I. F. B. Common (Australian Na- During the study in Thailand, we tional Insect Collection, Canberra). reared two specimens of an unidentified In Thailand, Cryptophlebia ombrodelta species of Thaumatotibia from Ptycho- (n = 5) was reared from a single host, pyxis species 1 (Euphorbiaceae) (n = 1) Lepisanthes rubinigosa (Sapindaceae). and Xerospermum noronhianum (Sa- This species has been reared numerous pindaceae) (n = 1). The genitalia of our times from many different genera of dissected male are very similar but not Fabaceae, with a few scattered records of identical to those illustrated by Pinkaew Oxalidaceae (Ho 1985), Polygonaceae (2006) for “Thaumatotibia sp. C” from (Clarke 1976, Zimmerman 1978), Pro- Thailand. teaceae (Zimmerman 1978, Jones 1994, Hung et al. 1998, Horak 2006), Rutaceae Andrioplecta species (Bradley 1953, Diakonoff 1968, Clarke 1976), and Sapindaceae (Clarke 1976, Andrioplecta includes ten species re- Zimmerman 1978, Horak 2006). Accord- stricted to Southeast Asia (Komai 1992, ing to CABI (1976), this species is widely Brown 2005). According to Komai distributed throughout southeastern Asia (1999), the larvae of three related spe- (Japan, India, Sri Lanka, Thailand, Cam- cies (i.e., A. subpulverula (Obraztsov) A. bodia, Vietnam, Philippines, Indonesia, shoreae Komai, and A. dierli Komai) New Guinea) and northern Australia. bore into the seed or seedlings of Ani- During the study in Thailand, an un- soptera, Dipterocarpus, Parashorea, and determined species of Cryptophlebia Shorea (all Dipterocarpaceae). Similar (n = 8) was reared from three different findings were reported by Komai (1992) hosts: Lepisanthes rubinigosa (Sapinda- and Nakagawa et al. (2003). Komai ceae) (n = 6), Wodyetia bifurcata (Are- (1999) also reported that three other re- caceae) (n = 1), and Parkia speciosa lated congeners are entomophagous, with

observations of A. leucodora (Meyrick) tinct, dense, rounded patch of short black feeding on scale insects on Pithecellobium scales from the posterior margin of the (Fabaceae) (Diakonoff 1968, Komai dorsum of abdominal segment III that is 1992); A. pulverula (Meyrick) feeding absent in other species of Andrioplecta, on cynipid gall wasp larvae on Castanea but similar to some species of Pammene and Quercus (Fagaceae) (Park 1983, Hu¨bner [1825]. Specimens were reared Abe 1990, 1995, Komai 1992, Abe and from unidentified Lauraceae species 1 Sanari 1992); and an undescribed spe- (n = 7) and unidentified Annonaceae cies from Malaysia feeding on imma- species 14 (n = 1). The strong preference tures of Beesoniidae in galls on a species for Dipterocarpaceae shown by three of Dipterocarpaceae (Komai 1999). described species of Andrioplecta sug- During the Thailand survey, An- gests that this undetermined species may drioplecta shoreae Komai (n = 73) was not belong to that genus. reared from 12 different hosts: Parashorea = stellata (Dipterocarpaceae) (n 47), Cydia species Shorea roxburghii (Dipterocarpaceae) (n = 9), Dipterocarpus grandiflorus Cydia includes over 100 species dis- (Dipterocarpaceae) (n = 4), Macaranga tributed worldwide (Brown 2005); denticulata (Euphorbiaceae) (n = 3), however, it is likely that the genus rep- Shorea hypochra (Dipterocarpaceae) (n = resents a polyphyletic assemblage of 3), Aglaia species 14 (Meliaceae) (n = 1), species with similarly plesiomorphic Calamus godefroyi (Arecaceae) (n = 1), genitalia. We reared a single individual Dipterocarpus species (Dipterocarpaceae) of an undetermined species that may (n = 1), Millettia atropurpurea (Fabaceae) represent a species Cydia. The larva was (n = 1), Platymitra siamensis (Annona- reared from the fruit of Millettia atro- ceae) (n = 1), Ptychopyxis species 1 (Eu- purpurea (Fabaceae). phorbiaceae) (n = 1), and Xerospermum noronhianum (Sapindaceae) (n = 1). DISCUSSION Although several plant families were We recorded tortricids from 30 plant documented as hosts, there is a strong species representing 12 plant families preference for Dipterocarpaceae, with (Table 1). Among the families, Sa- 88% of the specimens reared from this pindaceae was utilized by the greatest family. number of tortricid species (i.e., seven), One specimen of Andrioplecta sub- but over half of these (i.e., four) were pulverula (Obraztsov) was reared from represented by single individuals. The Dipterocarpus grandifloras (Dipter- next most utilized family was Annona- ocarpaceae) in Thailand. ceae, with six tortricid herbivores, but The survey also resulted in eight again, with four of these represented by specimens of an undetermined species single specimens. that may represent another species of Fabaceae produced the greatest num- Andrioplecta based on genital morphol- ber of specimens (n = 142): 140 of ogy, but the generic assignment is un- Cryptophlebia rhynchias, one of Cryp- certain. Although the adult and male tophlebia species, and one of An- genitalia are similar to those of A. pul- drioplecta shoreae. The next highest verula, the tiny signa of the female total was from Dipterocarpaceae, with genitalia are dissimilar to those of all 66 individuals, all but one of which is a Andrioplecta. Also, the male has a dis- species of Andrioplecta. Hence, plant

families that produced the greatest coding Opportunity FY013 and FY014 number of individuals hosted very lim- (to YB) and in-kind assistance from the ited herbivore diversity. Canadian Centre for DNA Barcoding From a taxonomic perspective, Hilar- and Southern China DNA Barcoding ographini was represented by a single Center facilitated the sequencing of species that was fairly common (n = 44) specimens. Nicholas Silverson provided and found only on Annonaceae. Archi- curatorial assistance at the National pini, likewise, was represent by a single Museum of Natural History, Wash- species, but only a single individual. ington, DC. We thank Todd Gilligan Members of this tribe are primarily leaf- (USDA, APHIS, Fort Collins, Colorado) rollers and are infrequently encountered and Richard Brown (Mississippi State on fruit. Two species of Microcorsini University) for helpful reviews of the were recorded, and as mentioned pre- manuscript. viously, this tribe appears to be restricted to seeds and fruits with little regard to Literature Cited plant taxonomy/chemistry. Four species Abe, Y. 1990. Notes on moths attacking cynipid of Olethreutini were reared, along with galls. Akitu (N.S.) 118: 6. eight species of Grapholitini. Larvae of Abe, Y. 1995. Relationships between the gall the last tribe are well known for their wasp, Trichagalma serratae (Ashmead) fruit-feeding habits, and the tribe in- (Hymenoptera: Cynipidae), and two moth cludes the most important tortricid fruit species, Andrioplecta pulverula (Meyrick) (Lepidoptera: Tortricidae) and Characoma pests throughout the world, including ruficirra (Hampson) (Lepidoptera: Noctui- codling moth (Cydia pomonella (Lin- dae). Applied Entomology and Zoology 30: naeus)), false codling moth (Thamatoti- 83–89. bia leucotreata), Oriental fruit moth Abe, Y., and T. Sanari. 1992. Larval feeding (Grapholita molesta (Busck)), macad- habits of two species attacking cynipid galls. amia nutborer (Cryptophlebia om- Proceedings of the XIX International Con- gress of Entomology (Abstract): 363. brodelta), and many others. Hence, it is Bae, Y.-S. and F. Komai. 1991. A revision of the not surprising that in the Thailand sur- Japanese species of the genus Lobesia vey, this tribe represented 50% of the Gueneé (Lepidoptera, Tortricidae), with de- total species and 73% of the total spec- scription of a new subgenus. Tyo to Ga 42: imens. These findings are similar to 115–141. those reported from surveys of seed- and Basset, Y., R. Ctvrtecka, C. Dahl, S. E. Miller, D. L. J. Quicke, S. T. Segar, H. Barrios, R. A. fruit-feeding tortricids in Kenya (Brown Beaver, J. W. Brown, S. Bunyavejchewin, S. et al. 2014) and Panama (Gripenberg Gripenberg, M. Knizek, P. Kongnoo, O. T. et al. 2019). Lewis, N. Pongpattananurak, P. Pramual, W. Sakchoowong, and M. Schutze. 2019. Insect assemblages attacking seeds and fruits in a ACKNOWLEDGMENTS rainforest in Thailand. Entomological Sci- ence: https://doi.org/10.1111/ens.12346. We thank ForestGEO and the Khao Basset, Y., C. Dahl, R. Ctvrtecka, S. Gripenberg, Chong Botanical Garden, Thailand, for O. T. Lewis, S. T. Segar, P. Klimes, H. logistical support. Collecting permits Barrios, J. W. Brown, S. Bunyavejchewin, B. were obtained from the National Re- A. Butcher, A. I. Cognato, S. Davies, O. search Council of Thailand. This study Kaman, P. Klimes, M. Knizek, S. E. Miller, G. E. Morse, V. Novotny, N. Pongpattana- was supported in part by the Czech nurak, P. Pramual, D. L. J. Quicke, R. K. Science Foundation (16-20825S). Grants Robbins, W. Sakchoowong, M. Schutze, E. J. from the Smithsonian Institution Bar- Vesterinen, W.-Z. Wang, Y.-Y. Wang, G.

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