PROC. ENTOMOL. SOC. WASH. 114(2), 2012, pp. 173–185

THE LIFE HISTORY AND IMMATURE STAGES OF THE MONOSTIGMA CHAMPION (COLEOPTERA: ) ON MICONIA CALVESCENS DC (MELASTOMATACEAE)

EDUARDO CHACO´ N-MADRIGAL,M.TRACY JOHNSON, AND PAUL HANSON

(ECM, PH) Escuela de Biologı´a, Universidad de Costa Rica, A.P. 2060, San Pedro de Montes de Oca, San Jose´, Costa Rica (e-mail: ECM [email protected], PH [email protected]); (MTJ) USDA Forest Service, Pacific Southwest Research Station, Institute of Pacific Islands Forestry, Volcano, Hawaii 96785, U.S.A. (e-mail: [email protected])

Abstract.—We describe and illustrate the life history and immature stages of Anthonomus monostigma Champion (Curculionidae: : ). This weevil is a fruit borer in Miconia calvescens DC (Melastomataceae), a Neotrop- ical tree that is invasive in Pacific islands. The has three instars, and development from egg to adult requires approximately two months. In Costa Rica, A. monostigma larvae were found in three Miconia , and adults fed only on Miconia species. Host relationships of the A. monostigma group suggest that this group could be related to the A. partiarius and A. albocivitensis groups (sensu Clark 1992, 1993b). The potential of A. monostigma as a biological control agent is discussed. Key Words: biological control, fruit-borer, invasive species, host , Costa Rica, Hawaii DOI: 10.4289/0013-8797.114.2.173

The velvet tree, Miconia calvescens For this reason researchers at the Univer- DC (Melastomataceae), is native to the sity of Costa Rica have been studying po- Neotropics and is invasive in Hawaii, tential biocontrol agents of M. calvescens Tahiti and other Pacific islands where it in its native habitat; examples of species poses a serious threat to natural ecosys- that have been studied include a psyllid tems because of its ability to displace na- (Hemiptera) (Burckhardt et al. 2005) and tive plants (Meyer 1996, 1998; Medeiros riodinid (Lepidoptera) (Allen 2010, Nishida et al. 1997). Classical biological control 2010). Several weevil (Curculionidae) via the introduction of natural enemies from specieshavealsobeenfoundfeedingon the native (Neotropical) habitat is probably this , including Cryptorhynchus mel- the best long-term means of managing this astomae Champion (Reichert et al. 2010) invasive plant (Badenes-Perez et al. 2008). and Copturus tricolor Champion in stems, and Pedetinus halticoides (Champion) and Anthonomus monostigma Champion in * Edited by Jens Prena; accepted by Robert R. fruits. The latter species is the focus of the Kula current investigation. 174 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

The curculionid Anthonomus MATERIALS AND METHODS Germar, with over 500 species worldwide, This study was conducted in Costa includes about 400 Neotropical species Rica from June 2005 to December 2006 (Clark 2008). It is the largest genus in the at three locations: Vereh (Cartago Prov- Anthonomini and contains many ince, Turrialba; 09°479000 N, 83°319400 species that are still undescribed. Several W; 1200 m elevation), located on the species are of economic importance as crop Caribbean slope of the Cordillera de pests (Burke and Cross 1966, Dieckmann Talamanca with annual rainfall of 3000 1968, Burke 1976), while others have been mm (unpubl. data, Centro Agrono´mico proposed as biological control agents Tropical de Investigacio´n y Ensen˜anza, (Pedrosa-Macedo et al. 2000, Medal Turrialba); La Selva Biological Station et al. 2002, Caxambu 2003). (Heredia Province, Sarapiquı´;10°259270 Because of their host specificity and N, 84°009050 W; 50 m elevation), lo- ability to damage flowers and fruits, An- cated in the Caribbean lowlands, with an thonomus have been studied as annual rainfall of ca. 4000 mm, peaking potential biological control agents for three from May to December, and a mean weeds of South American origin. These annual temperature of 25.8 °C (Sanford include Anthonomus tenebrosus Bozeman et al. 1994); and the campus of the Uni- for control of viarum Dunal versity of Costa Rica (San Jose´ Province, (Solanaceae) in the southeastern United ° 9 0 ° 9 0 States (Medal et al. 2002), Anthonomus Montes de Oca; 09 56 16 N, 84 03 00 santacruzi Hustache for control of Sola- W; 1200 m elevation), located in the Valle num mauritianum Scopoli (Solanaceae) in Central, with mean annual rainfall of ca. South Africa (Pedrosa-Macedo et al. 2000) 1800 mm, concentrated between May and and Anthonomus partiarius Boheman for November (unpubl. data 2004–2005, Uni- control of Tibouchina herbacea Cogniaux versity of Costa Rica Meteorological Sta- (Melastomataceae) in Hawaii (Pedrosa- tion) and a mean annual temperature of ca. ° Macedo et al. 2000, Caxambu 2003). 20 C (Stiles 1990). Anthonomus monostigma, described by Larvae of A. monostigma were col- Champion in 1903, has been collected lected at Vereh from dissected fruits of from Mexico to Panama (Champion 1903, M. calvescens and were preserved in O’Brien and Wibmer 1982). In Costa 70% ethanol. Additional fruits were held 3 3 Rica, adults were collected on Miconia in plastic containers (7 21 28 cm) to nervosa Triana (Melastomataceae) at the rear larvae to the adult stage. Adults La Selva Biological Station, Heredia were identified by Robert S. Anderson (Clark 1993a), but the specific nature of its (Canadian Museum of Nature). Ten third host plant associations was otherwise instar larvae were selected for descrip- previously unknown. Clark (1993a) clas- tion; these were macerated and boiled sified A. monostigma in the A. monostigma in a 10% KOH solution and washed in species group; however, the phylogenetic distilled water and 95% ethanol (May relationships between this and other spe- 1979). These larvae, their mouthparts cies groups of Anthonomus are unresolved and four pupae were mounted in euparal (Clark 1993a). Here we describe the life on glass slides for microscopic observa- history and illustrate the immature stages tion and illustration. The terminology for of A. monostigma and discuss its host re- larval and pupal descriptions follows lationships and potential as a biocontrol Burke (1968) and Ahmad and Burke agent for Miconia calvescens DC. (1972). VOLUME 114, NUMBER 2 175

To determine the number of instars, RESULTS 176 preserved larvae were digitally Larval development.—The frequency photographed at a fixed magnification distribution of head capsule widths under a microscope, and widths of head showed three separate peaks, indicating capsules were measured using Imagetool three larval instars (Fig. 1). Larvae 3.0 software (UTHSCSA 2002). The hatched from eggs within 5–9 days of duration of immature stages was de- oviposition. The first, second and third termined by rearing A. monostigma in stadia lasted approximately five, 10 and bagged infructescences of M. calves- 20 days, respectively (Table 1). After 45 cens trees cultivated on the campus of days from oviposition, when specimens the University of Costa Rica. Groups of were in the prepupal stage, we ran out of six adult A. monostigma were enclosed fruits for dissection, so it was not possible in seven mesh bags (10 3 15 cm), each bag covering a portion of a developing to determine the duration of the pupal infructescence, for a period of 5–6 days stage. Adults lived up to nine weeks in the to allow oviposition. After removal of laboratory. Ten specimens of one parasit- adults from the bags, samples of 20–30 oid species, Bracon sp. (Braconidae), fruits per infructescence were dissected emerged from approximately 600 M. every five days. Ages of dissected lar- calvescens fruits which contained Antho- vae were determined by days elapsed nomus larvae. No weevil species other than after the removal of adults. The larval A. monostigma were reared from M. cal- instar was determined based on head vescens fruits. capsule width. The longevity of adults Immature stages of A. monostigma was determined by maintaining 5–20 were monitored at Vereh during the M. adults in each of five petri dishes with calvescens fruiting period in 2005–2006. mature and immature M. calvescens First instars were observed in late November fruits given as food. Fresh food was 2005 and early January 2006 in small- and supplied weekly. medium-sized immature fruits. Second in- The host range of A. monostigma was stars were present between December 2005 evaluated by searching selected mela- and March 2006 in immature fruits. Third stome species at the La Selva Biological instars were observed between December Station and Vereh, Turrialba. A voucher and April in immature and nearly mature of each melastome species was deposited fruits. A few pupae were observed in March in the Herbarium of the University of and April in mature fruits. Costa Rica. At each locality 1–15 plants Host range.—AttheLaSelvaBi- of each potential host species were ological Station, 13 melastome species searched at 1–2 month intervals for one (Fig. 2) were surveyed for the presence year. Plant phenology was recorded of A. monostigma. Adults were found during each visit. Particular attention was feeding on fruits of Miconia affinis DC., given to flowers and fruits because the Miconia impetiolaris (Sw.) D. Don ex weevils use these structures for their DC., Miconia longifolia (Aubl.) DC. development. When fruits were present, and Miconia nervosa (Sm.) Triana, but samples of approximately 20–400 fruits larval development was detected only in per melastome species were collected fruits of M. longifolia. This host plant and taken back to the laboratory where produced fruits during almost the entire they were dissected or kept in plastic year, and adult weevils were observed containers for rearing. on M. longifolia during every visit (Fig. 2). 176 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Fig. 1. Frequency distribution of head capsule widths of A. monostigma larvae. Vertical dotted lines indicate probable limit of each instar (n = 176).

At Vereh, 12 melastome species were and reaching antennal base. Mandibles, surveyed (Fig. 3), and A. monostigma setae and sensilla reddish. Antenna 1- adults were found feeding on fruits of M. segmented, with sensorium encircled by affinis, M. calvescens and Miconia nutans sensorial papillae on basal membrane. Donn. Sm. Larval development was de- Frontal seta 4 long; setae 1, 3, and 5 very tected only in fruits of M. affinis and M. reduced and inconspicuous; frontal seta calvescens.NeitherM. affinis nor M. 2 absent. Four pairs of dorsal epicranial calvescens fruited year-round, but fruits setae present; seta 1 slightly shorter than of one or the other of these species were others;setae2and4similarinlength; present throughout the year (Fig. 3). seta 3 absent or not evident; and seta 5 Description of last instar larva (Fig. nearly twice the length of setae 2 and 4. 4).—Body moderately convexly curved Four short and obscure posterior epicranial dorsoventrally, white, length 2.42 ± 0.03 setae present. Two lateral epicranial setae mm (mean ± SD; n = 6). present and equal in length. One pair of Head: 0.51 ± 0.02 mm wide (n = 33), stemmata apparent, each positioned ante- hypognathous, semicircular, sclerotized, rior to epicranial seta 5 and near frontal yellow orange, slightly retracted in pro- suture. Clypeus with two inconspicuous thorax. Epicranial suture nearly half of setae on each side with one sensillum head length, frontal suture forming “V” between them. Labrum with three pairs VOLUME 114, NUMBER 2 177

Table 1. Frequency of A. monostigma larvae in M. Thorax and abdomen (description of calvescens fruit exposed to adults for 5–6 days. Fruits left lateral aspect of body, Fig. 4): Thoracic were dissected at several time periods after removal spiracle bicameral, its air tubes with 6–8 of adults to estimate time of development. annuli. Pronotum with three long setae and Number of larvae per instar Days since Number of one minute seta near spiracle. Mesothorax st nd rd removal of adults 1 instar 2 instar 3 instar fruits dissected and metathorax segments with two folds. 5 - - - 150 Prodorsum of mesothorax and metathorax 10 7 - - 132 with one moderately long seta. Postdorsum 11 - - - 20 of mesothorax and metathorax without 15 12 4 - 119 evident setae. Epipleural lobe of mesotho- 16 - - - 19 rax and metathorax with one long and one 20 - 14 - 143 21 - 2 - 10 minute seta. Pedal area of each thoracic 25 - 2 12 67 segment with five setae, two long and three 30 - - 16 66 minute. One pair of sternal setae evident 35 - - 14 72 only on segment I. Abdominal segments 38 - - 3 16 with eight pairs of lateral bicameral spira- 40 - - 11 55 45 - - 15 49 cles, their air tubes with 4–9 annuli, spi- Totals 19 22 71 918 racle 8 generally with more annuli and larger than others. Segments I–VIII each with one minute prodorsal seta and five postdorsal setae, setae 1, 2 and 4 minute, of setae and one pair of sensilla. Epi- setae 3 and 5 long. Spiracular area with pharynx bearing six anterolateral and four two minute or inconspicuous setae. Ep- anteromedial setae; labral rods moder- ipleurum of segments I–VIII with one ately long, converging posteriorly; four seta as long as postdorsal long setae and epipharyngeal sensory pores arranged in one very short seta. Pleuron with one two clusters with two pores in each. long and one minute seta. Pedal area Mandibles robust and pyramidal, dark with one minute or inconspicuous seta. orange, with two apical brown teeth, two Anus subterminal. mandibular setae equal in length, and two Description of pupa (Fig. 5).— sensilla. Maxillary palpus with two seg- Length 2.27 ± 0.05 mm (n = 11). Body ments; basal segment longer and wider yellow white; eyes creamy white in color, than apical segment. Two sensilla and one brown when mature. Rostrum with one short bristle on basal segment; apical pair of short distirostral setae located on segment with at least two tiny palpal se- conical tubercle. One pair of basirostral tae. Mala with six dorsal malar setae setae, like distirostral setae in shape and equal in length, five ventral malar setae length, also located on small conical tu- and a sensillum. Four stipital setae: 1 bercles. Head with one pair of small slightly longer than or equal to seta 4, 3 frontal setae. Supraorbital setae absent. slightly shorter than 4, and 2 reduced. Prothorax with slender and slightly curved Labial palpus consisting of two articles, pronotal setae, all similar in length; each each with one sensillum; premental scler- located on conical tubercle. Two pairs ite with long posteromedial extension; of anterolateral setae. One pair of ante- premental setae equal to stipital seta 3; romedial setae slightly more widely sep- glossa bearing four minute setae; post- arated than one pair of dorsomedial setae mental setae 1 and 3 short, 2 slightly located posteriorly. One pair of posteri- longer than stipital seta 1. omedial and three pairs of posterolateral 178 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Fig. 2. Bimonthly reproductive phenology of melastome species surveyed as potential A. mono- stigma hosts between July 2005 and May 2006 in La Selva. Numbers of trees sampled in parentheses; * = host fruit utilized by A. monostigma larvae and adults; ** = fruits eaten by adults. setae, all set on tubercles, arranged in DISCUSSION strongly curved row on each side. Meso- This is the first formal description of notum with two pairs of setae, similar in immature stages for any species in the A. length and borne on tubercles. Metanotum monostigma species group. Keys for the with three pairs of setae, similar in length, immature stages of Anthonomus are only two inner setae closer to each other than to available for a few species principally outer seta. Abdomen with five pairs of belonging to the Anthonomus grandis discotergal setae on terga I–VI, three of Boheman species group (Ahmad and these slender, curved and borne on small Burke 1972). Three dorsal folds in ab- tubercles, other two small, not evident or dominal segments, the subterminal anus absent in some cases. Three pairs of dis- and the absence of frontal seta 2 in cotergal setae on terga VII and VIII, setae A. monostigma larvae are diagnostic slender, curved and borne on tubercles. characters found in all Anthonomini Laterotergal seta 1 minute and laterotergal species (Ahmad and Burke 1972). seta 2 long, slender and borne on spinelike Compared with described larvae of tubercle. Spiracles evident only on first other Anthonomus species, A. mono- five segments. Abdominal sternal setae stigma larvae differ by having frontal absent. Segment IX without setae and setae1,3and5veryshortandonlyfrontal prolonged into sclerotized median pro- seta 4 long (Burke and Gates 1974, Clark cess terminating in double hook (like and Burke 1986, Caxambu 2003). The inverted T). pupa of A. monostigma differs considerably VOLUME 114, NUMBER 2 179

Fig. 3. Monthly reproductive phenology of melastome species surveyed as potential A. monostigma hosts between July 2005 and May 2006 in Vereh, Turrialba. Numbers of trees sampled in parentheses; * = host fruit utilized by A. monostigma larvae and adults; ** = fruits eaten by adults. from pupae of other Anthonomus described complete their development in less than by Burke (1968) in that A. monostigma has a month (Burke and Woodruff 1980). a single posterior process on segment IX, However, several temperate species feed similar to validus Dietz over a period of several months during (Burke 1968). Other Anthonomus species the winter (e.g., Anthonomus piri Kollar and Pseudanthonomus krameriae Pierce in the buds of pear and, rarely, apple trees have a paired process on segment IX (Dieckmann 1968)). The period of larval (Burke 1968, 1972). development time is likely tied to time for Three is the most common number of fruit development and maturation in hosts larval instars in the Anthonomini (Ahmad of A. monostigma. As in other anthono- and Burke 1972, Burke 1976), and A. mine species, pupation occurs within the monostigma shares this trait with A. same plant part where the larva develops grandis (Parrott et al. 1970), Anthonomus (Burke 1976). eugenii Cano (Patrock and Schuster The phylogenetic relationships between 1992) and A. partiarius (Caxambu 2003). the A. monostigma group and other groups Although this character may seem of little are unresolved (Clark 1993a). Host plant importance, it is highly conserved (Burke associations in species of Anthonomus 1976). The development from egg to seem to be useful for establishing phylo- adult required approximately two months genetic relationships between and within in A. monostigma, substantially longer groups (Clark and Burke 1996, W. Clark than some Anthonomus species that can pers. comm.). The only other Anthonomus 180 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Fig. 4. Last instar larva of Anthonomus monostigma. A, Entire larva, lateral view: as = abdominal spiracle, epl = epipleurum, pds = prodorsal seta, ptds = postdorsal setae, vpls = ventropleural seta; B, Head capsule: des = dorsoepicranial seta, ecs = epicranial suture, fs = frontal seta, fsu = frontal suture, les = lateral epicranial setae, pes = posteroepicranial setae, sen = sensilla; C, Antenna: sens = sensorium; D, Clypeus and labrum: clps = clypeal setae, labs = labral setae; E, Epipharynx: als = anterolateral setae, ams = anteromedial setae, sp = sensorial pores; F, Mandible: at = apical tooth, ms = mandibular seta; G, Maxilla and labium: dms = dorsal malar setae, labp = labial palpi, prms = premental seta, ptms = postmental setae, sen = sensilla, sts = stipital seta, vms = ventral malar setae. VOLUME 114, NUMBER 2 181

Fig. 5. Pupa of Anthonomus monostigma. A, Dorsal view: dts = discotergal setae, lts = laterotergal setae, ps = pronotal setae; B, Frontal view of head and rostrum: brs = basirostral setae, drs = distirostral setae, fs = frontal setae. species previously known to utilize host group because both groups have scler- plants in the Melastomataceae are otized enlargements of the ejaculatory Anthonomus coactus Clark, A. partiarius duct. Recently Anthonomus stellatus and Anthonomus opis Clark, all in the A. Clark, belonging to the A. albocivitensis partiarius group (Clark 1992, Caxambu group, was reared from fruits of Con- 2003). According to Clark (1993b), the ostegia oerstediana O. Berg ex Triana A. partiarius group could be related to (Melastomataceae) (E. Chaco´n unpubl.), the Anthonomus albocivitensis Clark supporting the link with the A. partiarius 182 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON group suggested by Clark (1993b). Given for only part of the year (Fig. 3). Preference that the known host plants of A. mono- for M. longifolia over M. affinis for re- stigma are Melastomataceae, the A. mon- production was notable at La Selva, sug- ostigma species group also appears to be gesting that the use of M. affinis at Vereh related to the A. partiarius and A. alboci- may have resulted from seasonal absence vitensis groups. However, in addition to of M. calvescens fruit. Elevated levels this host range, morphological or mo- of feeding by weevils on the few M. lecular studies are needed to de- calvescens fruiting structures that ma- finitively discern the phylogenetic re- tured out of season at Vereh provide ad- lationships among these species groups ditional circumstantial evidence that A. (W. Clark pers. comm). monostigma is highly host specific and Anthonomus species are associated has a strong preference for M. calvescens with a broad range of flowering plants, fruit at this site (Chaco´n-Madrigal 2007). with larvae occurring in 22 plant families To adapt to seasonal scarcity of host across 13 orders. However, individual fruit, A. monostigma may need to survive species appear to be narrowly specialized for long periods as adults. Adults lived without exception. Each of the 113 spe- for nine weeks in the laboratory, but we cies with recorded larval host plants has have no knowledge of how long or where been reported from just a single plant this weevil might remain concealed away family and may be restricted to a few from host plants in the field. Some An- closely related plant species within the thonomus species are known to overwinter same genus (Anderson 1993, Jones 2001). for almost a year as adults, waiting for fruit Anthonomus larvae commonly develop in in which to lay eggs (Burke 1976). floral buds or fruits, but some species in- Classical biological control has been duce galls or are inquilines in galls in- identified as an essential tool for long duced by other (Burke 1976). term management of M. calvescens in There is little information about adult diet, Pacific islands (Smith 2002). The po- but Anthonomus adults may be less spe- tential for A. monostigma as a biological cific than their larvae. For example, adults control agent appears high given its host of A. grandis are known to feed on pollen specificity and possible impacts on seed of five different plant families (Cuadrado production (E. Chaco´nunpubl.).Restric- 2002). tion to the genus Miconia would be a suit- Although we did not search for A. able level of specificity in Tahiti where monostigma weevils in plant families there are only a few native melastome other than Melastomataceae, finding this species, and family-level specificity species only in a few Miconia species likely would be suitable in Hawaii, where and no other genera suggests it is very there are no native Melastomataceae narrowly host specific. Physiological (Wagner et al. 1990). Although the re- and ecological factors governing host productive phenology of M. calvescens use by A. monostigma will require ad- may limit its suitability as a year-round ditional study. in just one host for A. monostigma in Costa Rica host at La Selva versus two hosts at (Chaco´n-Madrigal 2007), this does not Vereh may have been a function of fruit appear to be a barrier in Pacific islands, production at each site given that M. where there are multiple reproductive longifolia fruits were available year-round events per year, and fruits can be present at La Selva (Fig. 2), whereas each of the all year (Meyer 1998). Because the suc- two larval hosts at Vereh produced fruits cess of M. calvescens as an invasive plant VOLUME 114, NUMBER 2 183 is largely due to its prolific production of R.Sforza,M.C.Bon,H.C.Evans,P.E. bird-dispersed seeds (Medeiros et al. Hatcher, H. L. Hinz, and B. G. Recto, eds. 1997, Meyer 1998), fruit-feeding special- Proceedings of the XII International Sympo- ists such as A. monostigma merit consid- sium on Biological Control of Weeds. CAB International, Wallingford, UK., Montpellier, eration for their potential to slow the France. spread of M. calvescens into new areas Burckhardt, D., P. Hanson, and L. Madrigal. 2005. (Badenes-Perez et al. 2008). Diclidophlebia lucens n. sp. (Hemiptera: Psy- llidae) from Costa Rica, a potential control ACKNOWLEDGMENTS agent of Miconia calvescens (Melastomataceae) in Hawaii. Proceedings of the Entomological The authors extend their thanks to Society of Washington 107: 741–749. Edgar Rojas, Luis Madrigal, Alexander Burke, H. R. 1968. Pupae of the weevil tribe An- Castillo, Pablo Allen, Manuel Alfaro, thonomini (Coleoptera, Curculionidae). Texas Ronald Sanchez and Kenji Nishida for Agricultural Experiment Station Technical assistance in the field; and to Kenji Monograph 5: 1–92. Nishida, Edgar Rojas and two anonymous Burke, H. R. 1972. Observations on the life his- reviewers for their comments on the tory of Pseudanthonomus krameriae, with descriptions of larva and pupa (Coleoptera: manuscript. We were generously assisted Curculionidae). Annals of the Entomological in the identification of Anthonomus spe- Society of America 65: 619–622. cies by Robert Anderson. The Proyecto Burke, H. R. 1976. Bionomics of the Anthono- Miconia has been funded by the state of mine weevils. Annual Review of Entomology Hawaii, the National Park Service and the 21: 283–303. doi:10.1146/annurev.en.21. USGS Biological Resources Division, via 010176.001435 the University of Hawaii Pacific Co- Burke, H. R. and W. H. Cross. 1966. A new species of Anthonomus attacking cotton in operative Studies Unit, and by USDA Colombia with a review of the of Forest Service International Programs. Anthonomus vestitus (Coleoptera: - All work was possible thanks to the nidae). 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