Rev. Bio!. Trop., 31(1): 41-67, 1983

Leaf Mining (Díptera) in Costa Rica

Kenneth A. Spencer Department of BiologicalSciences, University of Exeter, England. Address for correspondence: Exwell Farm, Bray Shop, Callington PL1 7, 8QJ, England

(Received for publication November 26, 1982)

Abstract: During a 3 weeks' visit to Costa Rica in January/February, 1982, 9 new species of Agromyzidae were discovered and are de scribed in the genera Me lanagromyza, Ophiomyia, Japanagromyza, Liriomy za and . Eighteen species new to Costa Rica wer e also found, inc1uding the three important pests, huidobrensis, L. sativae and L. tri[o lii. In addition, leaf mines were collected on 10 plants , 2 certainly and 6 probably representing further undcscribed species. Keys to the 13 gen era and 53 species now known in Costa Rica are provided. centralis Frost is synonymised with Phytomyza loewii Hende!.

No specialised collecting of Agromyzidae has of leaf mining pests. The opportunity was also hitherto been undertaken in Costa Rica and this taken fo r general collecting on wild plants interesting and important fauna has been between visits to nurseries of ornamentals and poorly known. Only 27 species have been vegetable growing areas, in order to obtain a recorded (Spencer, 1973a) out of a total of better knowledge of the Agromyzidae as a approximately 220 species known in the whole in Costa Rica. Neotropical Regíon (Spencer, 1963 ; 1965; Throughout my stay I was based at San José, 1966; 1973a; 1973b; 1982 ; Valladares, 1982). apart from one night spent at Limón. Visits A catalogue of 141 species was provided were made to the main chrysanthemum by Spencer (1967) and a review of species in nurseries at Fraijanes, below Volcán Poás and at the Caribbean area was gíven by Spencer and San José de la Montaña, aboye Heredia, and Stegmaier (1973). also to the carnation nursery at Dano Grande During the last few years damage caused by (Cartago). The large nursery of ornamental leaf miners to a wide range of cultivated plants, shrubs, "Matas de Costa Rica", was inspected at including trees, vegetables and ornamentals, has Siquirres. Several visits were made to the greatly increased. In nature leaf miners are important vegetab1e growing areas north of normally effectively controlled by a large Cartago and at Santa Ana and San Rafael de complex of hymenopterous parasites, mainly in Ojo de Agua, west of Alajuela. General the families Braconidae and Eulophidae. collecting was undertaken, normally near the However, this natural balance has now been roadside, between these visits, and also on seriously disrupted with the widesp�ead use of waste ground near the Hotel Irazú, San José. modern pesticides resulting in the destruction I established that the serious pest, Liriomyzll of beneficial parasites. The economic trifolii, was present at both chrysanthemum importance of the Agromyzidae was reviewed nurseries and also on onions, beans and by Spencer (1973c). tomatoes. L. sativae was found on tomatoes, My visit to Costa Rica was funded by the beans, cucumbers and pepper (chili). The third German Agency fo r Technical Cooperation most important Liri omyza pest , L. (GTZ), with local support from Sanidad huidobrensis, was found at Fraijanes and Llano Vegetal, Ministerio de Agricultura y Ganadería Grande but only on wild hosts and the garden (MAG) and the American Flower Corporation, ornamental, Tropaeolum. primarily to identify and assess the importance maculosa, a species which can damage lettuce,

41 42 REVIST A DE BIOLOGIA TROPICAL was comrnon in the San José area on weeds. No In order to assist in fu ture identification, pests were fo und on carnations. keys are provided to the 13 genera and 53 The nine new species described below were species now known in Costa Rica. These are aH obtained by sweeping but leaf mines were simplified to cover the Costa Rican fauna but also found on a number of plants c1early more comprehensive keys are available in other representing fu rther undescribed species. De­ works which may need to be consulted during tailed' records are given below of the 18 species any thorough study of the Agromyzidae. discovered as new to Costa Rica. Populations of References are gíven to all relevant papers on both pests and other species were low, as was to the Agromyzidae of the Neotropical Regíon. In be expected at the begínning of the dry season the interests of economy the number of but it is now established that the Agromyzidae illustrations has been restricted but it is hoped are weH represented in Costa Rica and many that sufficient have been inc1uded to show the new species certainly await discovery. essential characters of the Agromyzidae .

Key to genera of Costa Rican Agromyzidae l. Subcosta developed throughout its length , coalescing with vein RI before reaching costa (Subfamily Agromyzinae) ...... 2 Subcosta becoming a fo ld distally and ending at costa separately and basad of Rl (Subfamily Phytomyzinae). 5 2(1). Pre-scutellars lacking 3 Pre-scutellars present 4

3(2). Mesonotum or abdomen normal1y with sorne metallic coloration , greenish or coppery, more rarely entirely black; antennae not normally separated by raised facial keel, male never with vibrissal fasciculus ; male genitalia: aedeagus symmetrical (Figs. 2,9) ......

...... • ...... Melanagromyza Hendel Uniformly black species; antennae frequently divided by conspicuous facial kee1, male with vibrissal fasciculus (Fig. 11); male genitalia: aedeagus asymmetrical (Figs. 13.15) ...... Ophiomyia Brashnikov 4(2). Two pairs ol" dorso-centrals ..Ja panagromyza Sasaka wa At least 3 pairs of de Agromyza Fallén

5(1). Orbital sctulae erect, reclina te or absent . 6 Orbital setulae distinctly proclinate . 12

6(5). Vein R4+5 ending nearest wi ng tip Lioy Vein Ml+2 ending nearest wing tip ...... 7

7(6). Male genitalia: sperm pump with enlarged, bowl- shaped base (Fig. 29 ). Amauromyza Hendel Male geni talia: sperm pump without such enlarged base ...... 8

8(7). Third antennal segment with a spine, scutel1um with only one pair of bristles (Subgenus ); or lunnle conspicuously higher than a semicirc1e (subgenus Po emiza) . Cerodontha Rondani

Third antcnnal segment wi thout spin e, lunule broader, lower ...... 9

9(8). Fore-tibia with lateral bristle; abdomen in male yel10w . Nemorimyza Frey Only mid-tibia sometimes with bristle, fore-tibia never 10

10(9). No pre-su tural de; epandrium ""i th eonspieuous patch of spines at hind-co rner (Fig. 45)...... Ca lycomyza Hendel Pre-sutural de pr esent ; epandrium w'Í thout such spines . . . . . 11

11(10). Frons and seutellum characteristically brigh t yel10w Liriomyza Mik Frons and scutellum variable, darker Phytoliriomyza Hendel in part

12(5 ). Costa extending to vein M 1+2 Phytoliriomyza Hendel in part Costa extending only to vein R4+5 13

13(12) Male genitalia: distal section of aedeagus simple (not bifid), Iying below a lobe with supporting sc1erites on dorsal side of aedeagus; pupation 111 mme Hardy Male genitalia : distal section of aedeagus bifid, without su eh lobe a boye; pupation on ..

ground ...... Ph ytomyza Fal1én SPENCER : Agromyzidae of Costa Rica 43

Sub-family Agromyzinae Ten apparently undescribed species have been seen from San José, La Caja (Spencer and Stegmaier, 1973: 150) but it was not felt Genus Me lanagromyza Hendel desirable to describe these largely on the basis of male genitalia of specimens in poor This large genus occurs throughout condition. Rearing specimens from puparia t he wo rld but is best represented in found in stems is not difficult and after such the tropics, with 80 species now known in the collecting, with the host identified and larval or Neotropical Region. The larva e are internal puparial characters known, many fu rther fee de rs, mainly in stems but also in species can certainly be discovered in Costa flower-heads. Five species have hitherto been Rica. known in Costa Rica. Two new species are Many species in this genus damage cultivated described below, one species only known from plants by boring in stems and two species California has been tentatively identified at San feeding in this way on tomatoes in Colombia, José and additional records are given of two Ecuador and Venezuela were described by fu rther species. Steyskal (cf. Spencer, 1973b: 24).

Key RicanMelanagromyza to Costa species

1. Squamal fringe pale, whitish ...... 2 Squamal fringe dark, brown or black 4

2(1). Jowls narrow, 1/8 height of eye ; small species, wing length 1.75·2.0 mm . floris Spencer Note: Described by Spencer, 1963: 313 from Mexico, Puerto Rico and one female from Higuito, Costa Rica. Subsequen tly recorded from Venezuela (Spencer, 1973b: 18 and Chile, Spencer, 1982); also known from Argentina (Valladares). Host Bidens pilosa, larvae feeding in seed-heads .

Jowls broader, 1/5-1/4 height of eye . .. 3

3(2). Male gelÚtalia: aedeagus as in Fig. 10 neotrop íea Spencer Note: Described from Mexico and Brazil (Spencer, 1963:319). Subsequently fo und Venezuela (Spencer, 1973b: 22) and in Chile (Spencer, 1982) wh ere specimens werein reared from seed-heads of Bidens pilosa at Arica; also present at Córdoba , Argentina (Valladares). One male seen from San José, Caja, 1930 (H. Schmidt). Male genitalia : aedeagus as in Fig. 5 La . . cirsíophíla Spen cer

4(1). Mesonotum and abdomen greenish 5 Mesonotum and abdomen black 6

5(4). Eye bare in male ; female ovipositor greatly elongated mínima (Malloch) Eye wi th patch of conspicuous hairs a t level of ors ; ovipositor normal ... mayí sp .n. Note: Discussed by Spencer and Stegmaier (1973 :44) ; larva fe eding and pupating seed-heads of Asteraceae, confirmed on Wedelia and probably on Eupatorium; kno,,; n fromin Florida, Trinidad, Jamaica. One male, one fe male previously reared from San José, Caja . One female caught at José, waste ground ncar Hotel Irazú, 21-1-82 and oneLa a t Santa Ana, 3-II-82. San

6(4). Arista conspicuously pubescerit; wing length in male 2.1 mm . .. eh ita riensís sp.n. Arista virtually bare; smaller species, wing length 1.7-2.0 mm ...... 7

7 (6). Male genitalia : aedeagus as in Figs. 6, 7 . . .. . eleuth erensis Spencer . Note: Described from Eleuthera Is. (Bahamas) and two males fr om San José, La Caja (Spencer and Stegmaier, 1973 : 155). One male, one female caught at San José, waste ground near Hotel Irazú, 25 and 31-1-82. This minute black species is only reliably distinguishable from M. earibbea by the m ale genitalia, \\hich are of the same general form , but siglÚficantly different in detail. Male genitalia : aedeagus as in Figs. 1, 2 . . ea ribbea Spencer Note: Described from Florida , Dominica and one male from Higuito, near San Mateo, Costa Rica (Spencer and Stegmaier, 1973: 34). 44 REVISTA DE BIOLOGIA TROPICAL

.,.. ;;---:"'-� �'� ��� .... �? i 1

\�.-. 10

Figs. 1, 2. Melanagromyza caribbea: l. aedeagus, Melanagromyza chitariensis sp.n. side view; 2. same, ven tral view. (Figs. 3,4)

Figs. 3, 4. Melanagromyza chitariensis: 3. aedeagus, side view; 4. same, ventral view. Head: Frons broad, twice width of eye, Fig. 5. Melanagromyza e irsiophila: aedeagus, side not significantly projecting above eye in view (San José). profile ; 2 upper orbital bristles (ors), 2 lower. orbitals (ori), orbital setulae sparse, Figs. 6, 7. Melanagromyza eleutherensis: 6. 7. reclinate ; ocellar triangle extending only to aedeagus, side view; same, ventral view. level of lower ors, orbits narrow ; jowls Figs . 8, 9. Melanagromyza mayi: 8. aedeagus , side deepest in front, narrow, about 1/7 height view; 9. same , ventral view. of eye, this bare; third a ntennal segment srnall, round, arista long, little shorter than 10. Melanagromyza neotropica : Fig. aedeagus , side height of eye, conspicuously pubescent. view (ex Bidens, Argentina). SPENCER : Agromyzidae of Costa Rica 45

Mesonotum: 2 strong dc, acrostichals (acr) Sorne thistles were growing where the main numerous, in about 10 rows. series was caught but the dominant composite was Bidens pilosa and this is possibly the host, Wing: Length in male 2.1 costa with the larvae feeding and pupating internally mm; extending strongly to vein M 1 +2, last section in the stems. of M 3+4 shorter than penultimate, in ratio 15: 25, inner cross-vein at rnidpoint of discal cell. Material seen: San José, waste ground near Hotel Irazú, 5 9 9, 21-25-1-82; Llano Ó, Colour: Black; ocellar triangle and orbits Grande, aboye Cartago, 2100 m, 2 9, 26-1-82; W only weak1y shining; mesonotum moderately of Atenas, Nature Reserve, 1 9, 1-11-82. shining from rear, more mat from front; squamae dark grey, margin and fringe black; Melanagromyza mayi sp.n. halteres black. (Figs. 8,9)

Male genitalia: Aedeagus as in Figs. 3, 4; Head: Frons narrow, equal to width of eye, sperm pump with blade long and narrow; inner not projecting aboye eye in profile;2 ors, 2 ori; margin of spandrium slightly rounded, with orbits broad, with orbital setulae short but numerous short bristles; end of hypandrium numerC'lls, in 2 rows, those nearest eye margin broad, blunt. reclínate, the inner row proclinate; apex of ocellar triangle extending' to just below level of Holotype: Costa Rica, roadside E of lower ors; jowls deepest in centre, rather Chitaría (Turrialba)0, 29-1-82, in AC (author's narrow, about 1/7 height of eye, this round, collection). upright, with distinct patch of hairs in male at leve! of ors; third antennal segment smaIl, Remarks: The pubescent arista and larger round, arista appearing bare. size distinguish this species from the two other entirely black species known in Costa Rica, M. Mesonotum: 2 strong dc, acr numerous, in caribbea and M. eleutherensis. about 8 rows.

Me la nagromyza cirsiophila Spencer Wing: Length 1.9-2.0 mm in both sexes; New to Costa Rica costa extending strongly to vein M 1 +2, last (Fig. 5) section of M3+4 short, little more than 1/2 length of penultimate. (In holotype there is a Me la nagromyza cirsiophilaSpencer, 1981:41 venational aberration with MI +2 curvingup to join R4+5 at the distal third of the discal cell). This greenish species with the squamal fringe pale was reared from "thistle" in southem Colour: Head black, both ocellar triangle California. The type specimens are large, with and orbits moderately shining; mesonotum and wing length of 2.5-3.25 mm. The male genitalia scuteIlum distinctly shining green viewed from are distinctive (cf. Spencer, 1981 : Figs. 23, 24). rear, more mat seen from front, abdomen A series of 14 specimens from San José and 3 almost brilliantly shining green; squamae grey, others fr om Atenas and Uan0 Grande are margin and fringe black; halteres black but substantially smaller, with wing length of 2-2.5 paler aboye in holotype. mm. However, the male genitalía (Fig. 5) agree exactly with the holotype and 1 hesita te to Male genitalia: Aedeagus as in Figs. 8, 9, treat the Costa Rican population as distinct basal sclerite closely adjoining distiphallus me rely on the smaIler size. It wiIl be of the complex; sperm pump large, with well-defined utmost interest to findaddit ional material from 'central vein; inner margin of epandrium intermediate areas further north in Central rounded, with a row of short bristl es; America and in Mexico to ascertain the size of epandrium with greatly elongated apodeme. populations there. Further clarification of the Costa Rica species will only be possible when Holotype: Costa Rica, San José, waste additional material becomes available and in ground near Hoteló, Irazú, 31-1-82; paratypes: 2 particular when the host can be establíshed. 9, same locality, 25-1-82, all in AC. A male 46 REVISTA DE BIOLOGIA TROPICAL

Key to Rican Ophiomyia spedes Costa 1. Halteres brown below, with circular white area aboye . . .. punctohalterata (Frost) Note: Described from Guatemala by Frost (1936, Ann. Entorno!. Soc. Amer., 29: 311). Eight males and 4 females were seen from San José, La Caj a (SpenceJ and Stegmaier, 1973: 171); also known from Venezuela (Spencer, 1973b: 29). Halteres urúformly dark . . . . 2

2(1). Squamal fringe white .. buscki (Frost) Squamal fringe brown or black 3

3(2). Proboscis elongate (Fig. 16) 4 Proboscis short, normal . 5 0 4(3). Large species, wing length in female 2.9 mm ; vibrissal corner forrIÚng angle of 80 • • •• ...... magna sp.n. Smaller species, wing length 2-2.2 vibrissal corner acute, forming angle of less than 0 mm; 40 • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • valida Spencer Note: Described from Bahamas; Jamaica and 7 males from San José, La Caja (Spencer and Stegmaier, 1973: 173).

Ocellar triangle brilliantly shlning; wing with veins pale, whltish . .gen tilis Spen cer 5(2), Ocellar triangle only moderately shlning; veins dark brúwn ... costaricensis sp. n.

from Santa Ana, 3-Il-82 closely resembles the Over 150 species are known and the genus is type series but is not included as a paratype. lt weIl represented in the Neotropical Regíon, is larger, with wing length of 2.25 mm, with extending to Chile, with 7 species known in very slight differences in the male genitalia. lt Venezuela. Three species have hitherto been may possibly represent a fu rther species but known in Costa Rica. Two new species are more material wiIl be required before this can described below and one, o. buscki (Frost), is be decided. recorded as new to Costa Rica, originaIly described from Panamá. Remarks: 1 have pleasure in naming this Larvae feed predominantly as external species after Ing. Juan José May, Vice-President stem-miners, pupating in the stem with the of the Convenio Costarricense-Alemán, MAG, anterior spiracles projecting through the whose courtesy and assitance throughout my epidermis. Possibly the most serious agromyzid visit was greatly appreciated. pest in the world, o. phaseo/i (Tryon), feeds on beans throughout the Old World tropics. This M. mayi generally resembles M. erechtitidis species recently reached Hawaii but is not so far Spencer (1966a: p.l O) known from Florida, known in mainland America or in the feeding in fl ower-heads of Erechtites Neotropical Regíon. hieraciifolia. The aedeagus somewhat resembles that of M. neotropica (Fig . .10) but in this Ophiomyia buscki (Frost) species the squamal fringe is paleo New to Costa Rica

Genus Ophiomyia Braschnikov Agromyza buscki Frost, 1936, Ann. Entorno!. Soco Amer. 29: 315. Species in this genus are entirely black, as Ophiomyia buscki, Frick, 1957, Ann. Entorno!. opposed to the normaIly greenish or bluish Soco Amer. 49: 200. colour of the mesonotum or abdomen normaIly found in Melanagromyza. CharacteristicaIly, This is a distinctive species with white squamal males have a pronounced vibrissal fasciculus fringe and elongated proboscis. Spencer (1963: (Fig. 11) and there is a prominent raised facial 325) considered specimens bred from stem­ keel in both sexes. However, in a few species mines on Alternan thera philoxeroides in Argen­ these characters may be lacking and correct tina and Brazil to represent O. buscki described generic assignment may only be possible from from Panamá and the male genitalia and the male genitalia. puparial spiracles were illustrated. However, SPENCER : Agromyzidae of Costa Rica 47

12

11

-J'4��" :'�::' , ��':.- , , ' > '" . . : . , . ' "" ' �.,.-- , ,�"./��. -""-'-"', ,:�' " -,, �;, � ;;: "�;;: " ... ' . . ,,::.; .. ���, ,; ._,'�::::. .';/...... " ( . " "�'' - ". " <' 14

Figs. 11, 13. Ophiomyia costaricensis: 11. head; identical and a further rather similar species, O. 12. aedeagus, side view; 13. same, ventral view. haydeni, was described from the Bahamas and southern Florida (Spencer and Stegmaier, Figs. 14, 15. Ophiomyia gentilis: 14. aedeagus, side view; 15. same, ventral view (San José). 1973: 57). A single female from Moín, near Limón, Fig. 16. Ophiomyia magna: head. 29-1-82, is tentatively identifiedas O. buscki, as this habitat certainly closely resembles that of later sorne doubt was expressed as to whether the type locality, Corozal, Panamá but this the populations in Argentina and Brazil on the group can only be further clarifiedas additional one hand and Panamá on the other were really material becomes available. 48 REVISTA DE BIOLOGIA TROPICAL

Ophiomyia costaricensis sp.n. keel narrowe r, less raised; vibrissa not curving (Figs. 11-13) uniformly, with distinct bend at midpoint (cf. Spencer and Stegmaier 1973 : fig. 126) ; last Head (Fig. 11): Frons twice width of eye, section of ve in M3+4 sligktly shorter, in ratio not projecting aboye eye in profile; 2 ors, 2 ori, 15: 20 with penultimate, inner cross-vein at orbital setulae sparse, reclinate; ocellar triangle midpoint of discal cell; veins unusually pale; elongated, narrowly extending to below lower male genitalia: aedeagus as in Figs. 14, 15. The ori ; jowls fo aning angle of 800, approx. 1/10 two distal tubules more distinct, in ventral view height of eye; antennal bases narrowly divided distiphallus less obviously divided; inner comer by low fa cial keel which widens in lower halL or surstyli with only 2 teeth. vibrissal fasciculus finely tapering. Ophiomyia magna sp.n. (Fig. 16) Mesonotum: 2+0 de, acr in about 6 rows. Head (Fig. 16): Frons almost twice width of Wing: Length in male 1.9 mm; last section of eye, narrowly projecting aboye eye in profile ; 2 vein M3+4 relatively long, in ratio 16: 19 with ors, 2 ori ; orbital setulae long, sparse, reclinate; penultimate; inne r cross-vein well beyond ocellar triangle extending to lower ors; jowls midpoint of discal cell. broad, 1/3 height of eye , fo rrning angle of 0 about 80 , extending to the lateral margin of Colour: Black; ocellar triangle moderately the fa ce on which the vibrissa is located; shining; mesonotum and ab domen only slightly proboscis greatly elongated : antennae widely shining ; squamae grey, margín and fr inge black; separated by a broad, raised fa cial kee!. wing with veins brown, normal . Mesonotum: 2+0 de, acr in 10 rows. Male genitalia: Aedeagus highly asymmetrical, as in Figs. 12, 13, distiphallus Wing: Length in fe male 2.9 mm ; costa distinctly divided in ventral view ; inner comer extending st rongly to vein M 1+2, last section of surstyli with 4 teeth. ofM 3+4 in ratio 20:34 with penultimate, inner cross-vein at rnidpoint of discal cel!. Holotype: Costa Rica, San José, waste ó, ground near Hotel Irazú, 31-1:82, in AC. Colour: Black; ocellar triangle moderately shining; mesonotum weakly shining, abdomen Remarks: This species is c1early c10sely more so; squamae pale grey, margín and fringe related to O. gentilis Spencer. The mal e black; halteres black. genitalia are of the same distinctive, asymmetrical foan but differ conside rably in Holotype: 9, Costa Rica, below Volcán Poás, detai!. The distinctness of the two species is 1950 m, at bridge 5 km aboye two cafes near also apparent from a number of morphologícal Fraijanes, 6-11-82, in AC. diffe rences. Remarks: This is the largest species known Ophiomyiage ntilis Spencer in the Neotropical Regíon. It is normally (Figs. 14, 15) undesirable to describe species in this genus on the basis of females; however, the elongate Ophiomyia gentilis Spencer, in Spencer and proboscis, large size, broad jowls and broad, Stegmaier, 1973: 57. high facial keel are distinctive characters and This species was described from specimens there should be no difficulty in associating from southem Florida, Dominica and one mal e males with the female holotype. It can be [rom San José, La Caja. A further male was assumed wi th certainty, that the male will be obtained from San José, waste ground near typical of the genus in having a developed Hotel Irazú, 25-1-82. vibrissal fa sciculus. o. gentilis closely resembles O. costaricensis Genus Japanagro my za Sasaka wa with the following points of difference: frons narrower, slightly less than 1.5 times wid th of As its name implies, this genus was originally eye; ocellar triangle brilliantly shining; fa cial described from Japan but it was later found to SPENCER : Agromyzidae of Costa Rica 49

occur widely in the Oriental Region and also in Spencer and Stegmaier, 1973 : 140). Dr. von southern Florida and the Caribbean area. Two Tschirnhaus has recently informed me that he species are now known in Costa Rica, of which has seen yellow species of Japanagromyza one, occurring as a minor pest on Phaseolus, is during a visit to Perú in 1981 and it therefore formally described below. seems probable that Geratomyza will prove to All species hitherto known have been dark, be synonymous with Japanagromyza. However, black or gr eenish. Two species with this synonymy will not be fo rmally proposed, conspicjJous yellow coloration from Jamaica, until the two species of Geratomyza can be Grand Cayman and the Bahamas were placed in re-examined. a new genus Geratomyza by Spencer (in

Key to Costa Ri Japanagromyza species can

1. Large speeies , wing 2.5-2.75 mm; male genitalia: hypandrium with short but distinct ap odeme (Fig. 19); host Phaseolus ...... phaseoli sp. n. Smaller species, wing length 1.9-2.4 mm; male genitalia : hypandrium rounded at end, without apodeme (ef. Speneer and Stegmaier, 1973: Fig. 36B) ...... perpetuaSpeneer Note: Described from Florida, Bahamas, Puerto . and. 7 males from. San José,. La Caja (Speneer and Stegmaier, 1973: 28b). There is no indication of theRico host.

Japanagromyza phaseoli sp.n. with several strong bristles (Fig. 20A), cerci (Figs. 17-22) large, with numerous strong bristles (Fig. 20B).

Head (Fig. 17): Frons narrow , equal to Host: Phgseolus spp., also reported in Perú width of eye, not projecting aboye eye in on Brassica oleracea larva forming irregular profile ; 2 equal ors, 2 ori, the upper only blotch mine , sometimes several in a leaf (Fig. slightly weaker than the ors and reclinate, the 22), pupating externally ; puparium lower weaker, more incurved; orbital setulae brownish-orange, posterior spiracles each with 3 sparse, reclinate; jowls narrow , approx. 1/10 flat processes again subdivided into 3 small, height of eye; third antennal segment large, elongate pores (Fig. 21). arista long. Holotype: 0, Venezuela, Rancho Grande, Mesonotum: 2 strong dc, acr in 8 rows in Aragua, eme rge d 5-V -72 ex mine on Phaseolus fron t; pre-scutellars well-developed. 18-IV-72; El Valle , DF, 1 9, 24-11-50 ex "caraotas" (R. Hernández); Perú, Carie te, 1 0, 3 Legs: Fore-tibia with 1 lateral bristle, 9,20-VIA1 (EJ. Hambleton) ; La Molina, 20, mid-tibia with 2. 59, also 4 pupal cases, ex Phaseolus vulga ris, 21-IX-62 (Olave 89-62); Huara, 6 49, 2-VI-67 o, Wing: Length from 2.5 mm in male to 2.75 (K. Raven) ; Lamb ayeque, 10, 19, "on mm in fe male; costa extending st rongly to vein cabbage", 16-V-66 (H . Torres). Leaf mines with Ml+2, last section of MH4 2/3 penultimate. larvae found in Costa Rica at Fraijanes, 1600 m, 22-1-82 , and near Chitaría de Turrialba, Co lou r: Frons slight1y variable, dark 29-1-82; also empty mines at Llano Grande, brownish to black, orbits shining black ; 2100 m, aboye Cartago, 26-1-82. Holotype and mesonotum pre dominantly mat greyish, paratypes in BM, fu rther paratypes in USNM, sometimes with faint greenish tinge, at most collection of U:niversidad Agraria, Lima and weakIy shining; pleura black, with greyish tinge; Facultad de Agronomía, Maracay . legs black ; abdomen shining greenish; squamae, including margin and ftinge, silvery; halteres Rema rks: This species was originally white. diagnosed by Spencer (1963:304) as Japanagromyza sp. (Perú) but was not formally Male genitalia: Aedeagus with 2 pale coiled described . It was later discovered again in Perú tubules (Fig. 18); hypandrium with distinct but by C.A. Korytkowski and I found it in short apodeme (Fig. 19) ; surstyli elongate, Venezuela (Spencer, 1973 b: 34; 1973c: 83 and 50 REVISTA DE BIOLOGIA TROPICAL

17

19

A

····-0·· " '-""\' 1 -, ....-.. �

21

Figs. 17, 22. Ja panagromyzo phaseoli: 17. head; now be published, as he has laken up other 18. aedeagus (h oloty pe); 19, hypandrium ;20. work and in view of the economic importance epandrium, with A. surstylus, B. cercus; 21. of Ihe species, Ihe description is published here. posterior spiracles of puparium (Chitaría) ; 22. leaf The male genitalia of \he specimen from mines on Phaseolus (Chitaría), Cariete (slide 439, USNM) have been re.examined and it can be confirmed that the 373). 1 had been proposing lo describe Ihe illustration of the aedeagus (Spencer, 1963: species al Ihal lime bul agreed Ihal \he foonal Fig. 13d) is inaccurate, owing to the loss ofthe descriplion should be prepared by coiled end of the distiphallus during KO!ylkowski. However, hls description will nol preparation (see Fig. 18). 51 SPENCER : Agromyzidae of Costa Rica

:r. �: Y' :>::" " \.,¡;� ·':"�Y, ::".':':":"

: 1 . �. , . .. . ��� -o ":" ;: /�'�'-' --:- '''':'. _--. :-.: .""�-� ' o;::;��""� �� ��2c�'�; .... , 25 ,.-: . 2 6 ::'>; ':. '

>,sm:�:';'f'\'5:.� 28 �;�?:: 27

. , ..... �:�;-

Figs. 23, 24 . Agromyza venezaana:23. hea d; 24. where Ph aseolus was examined. The larva forros wing. a relatively large blotch mine and several mines

Figs. 25, 26. Phy tobia dorsocentralis: 25. may be present in a single leaf. Plan ts attacked aedeagus, side view; 26. distiphallus, ventral view. in this way must be considerably weakened, inevitably leading to Sorne loss in yield. Y oung Fig. 27, 28. Phytobia picta: 27. aedeagus , side plants were seen to be completely destroyed 28. view ; distiphallus, ventral view. Venezuela (Spencer, 1973c: 373).in Unfortunately no adults were reared from 1. phaseoli appears to be widespread on the puparia obtained in Costa Rica but the Phaseolus in the Neotropical Region and was distinctive arrangement of the puparial present at the three localities in Costa Rica spiracles (Fig. 21) identical to that of is 52 REVISTA DE BIOLOGIA TROPICAL

reared specimens from Venezuela and Perú. Sub-family Phytomyzinae The occurrence of trus species on Brassica oleracea (cabbage) in Perú is puzzling and it Genus Phytobia Lioy remains to be established whether trus was an isolated occurrence Or whether this may serve This genus is now restricted to the mainly as a regular alternate host. large species wh ose larvae bore internally the cambium of trees. Five species are knownin in Costa Rica (Spencer and Stegmaier, 1973) but two have not been formally described in the Genus Agromyza F allén absence of males. Two species have been described from This large genus occurs predominantly in Ecuador and Perú (Spencer, 1977) and fo ur are temperate áreas of the northern hemisphere and known from Brazil (Spencer, 1966b). However, over 1 50 species are known through ou t the the genus is apparently very large in the world. Only 6 species have been recorded in the Neotropical Regíon. Dr. von Tschirnhaus Neotropical Regíon. Two species are known in recently informed me that he discovered 20 Costa Rica, both certainly grass-feeders. A new undescribed species in a rain forest in Perú at a record ofA. animata is given below. single locality.

Key to Costa Rican Agromyza species

1. Squamal fringe pale, whitsh ...... venezolana Spencer Note: Described from the Botanical Gardens, Caracas (Spencer, 1963) and later identified from two males from San José, La Caja (Spencer and Stegmaier, 1973: 139). The head and wing are shown in Figs. 23,24 .

Squamal fringe dark, brownish-black ...... animata Spencer Note: Described from a single male from Costa Rica , Higuito de San Mateo (Spencer, 1973a: 152). A female caught at Lim ón, 29-1-82 is accepted as representing tlús species.

Key to Costa Rican Phyt obia species

1. Mesonotum and scutellum entirely black ...... sp.l (Costa Rica) Note : One female seen from "Podrogoso" (as Pedregosa), Spencer and Stegmaier (1973: 179). Scutellum yellow; mesonotum partially yellow . . 2

2(1) Wings pictured ...... rabelloi Spencer Note: Described from Brazil (Spencer, 1966b: 145). A single female seen from Suiza (near Turrialba). In view of the numerous species no w known to exist Lain the Neotropical Region having marked win gs, it is possible or even probable that this represents a misidentification and tha! tlús species is distinct and undescribed. Wings clear ...... 3

3(2) Dark area of mesonotum solid, only semicircle adjoining scutel1um yel1ow. sp. 2 (Costa Rica) Note : One female seen fr om La Suiza de Turrialba (Spencer and Stegmaier, 1973: 180) Dark area of mesonotum extended into 2 bands at sides ...... 4

4(3) Very large species, wing length 3.25-4 mm ; male genitalia: aedeagus as in Figs . 25, 26 ...... dors ocent ralis (Frost) Note : Described from Panamá by Frost (1936, Ann. Entorno!. Soco Amer. 29: 307); three males, one female seen from Suiza, one male from Podrogoso (Spencer and Stegmaier, 1973 : 175). La Smaller species, wing length 2.4-3.1 mm; male genitalia: aedeagus as in Figs. 27, 28 ...... picta (Coquillet) Note: Described from Mexico by Coquillett (1902, J. New York Entorno!. Soco 10: 188). Three females seen from San Mateo, one from San José, Caja (Spencer and Stegmaier, 1973: 176). La SPENCER : Agrornyzidae of Costa Rica 53

The larvae can cause considerable damage to predominantly Nearctic and Palaearctic but wood used for the manufac ture of fumiture small populations are present in Florida and and other products. However, no trees have one female has been seen from Costa Rica, San been . identified as hosts in Costa Rica and no José, La Caja. economic damage has yet been reported. Hosts are exclusively in the Asteraceae, the main genera attacked being Aster, Baccharis Genus Amauromyza Hendel and So lidago. The adults are large, with wing (Fig. 29) length of 2.7-3.3 mm, distinctive characters T hi s genus is poorly represented in being the silvery lunule, a strong bristle on the the Neotropical Region, with only 4 fore-tibia, silvery-white squamal fringe and the species known out of the world total yellowish abdomen of the maleo The isolated of 40. Of these, one was discovered in position of this species is indicateo by the male 1967 feeding on Co mmelina in Cuba genitalia (Spencer and Stegmaier, 1973 ; Fig. and the formal description was to have 24 1). been prepared by Dr. E. Rohden do rf, Empty mines (Fig. 30) on an unidentified Prague, but this has never been done . woody species of Asteraceae at the roadside (Spencer and Stegmaie r, 1973: 186). West of San Ramón, l-II-82, represent N Twenty species are known in the United States. posticata. A distinctive charac ter of all species in this Genus Cerodontha Rondani somewhat diverse genus is the large, New to Costa Rica bowl-shaped base of the sperm pump, shown in A. maculosa in Fig. 29. In other species this is This large genus of world-wide distribution even more st rongly developed. has been split into 7 subgenera, of which two Only the single species A. maculosa are now recorded below. The genus reaches (Malloch) is known in Costa Rica and was Chile where 7 species have no w been recorded recorded by Spencer (l973a). This large, black (Spencer, 1982) but 43 are known in U .S.A. species is distinctive in having the halteres The genus has been studied in detail in Europe, largely white but partially black below. Leaf with 75 species now described. mines were not uncommon in San José in Hosts are exc lusively in the January, 1982 on the common weed Co nyza. Monocotyledoneae and the two Costa Rican The larvae of A. maculosa fo rm large blotch species are both well-known grass-feeders. mines and the species has been recorded as a Subgenus Ce rodontha Rorrlani pest on lettuce in Trinidad and Venezuela. No large populations are known to have occurred C. dorsalis (Loew) is a distinctive species on Chrysanthemum but the possibility of with a conspicuous spine on the third antennal damage being caused cannot be excluded. Hosts segment. It was described originally fr om are exclusively the Asteraceae and many Washington, D.C. but is now known to occur in genera are attacked. This is the most virtually throughout the United States. Its widespread of all Neotropical species, occurring range extends in South America to Brazil, from Argentina and Chile to New York. Bolivia and Chile (Spencer, 1982) and its occurrence in Costa Rica was to be expected. Material seen: San José, Hotel Irazú and The larvae fe ed on numerous genera of roadside nearby, many mines on Conyza Poaceae, including all the main cereal crops. As apurensis HBK , mostly empty. No adults many as eight generations ayear have been in obtained but two parasites, Opius sp. ; emerged observed in southem California and the species 7 and ll-II-82; Fraijanes, below Volcán Poás, is of sorne economic impo rtance. Young plants 1600 m, empty mines on Emilia fos bergi, can be damaged'by numerous fe eding punctures 21-1-82. in the leaves but it has not been confirmed whether there is any actual reduction in yield Genus Nemorimyza Frey (Spencer, 1973c: 293 ; 1981: 174). (Fig. 30) Material seen: San José, waste ground near Only a single species, posticata (Meigen), is Hotel Irazú, 2 9, 25-1-82 ; below Volcán Barva, known in this genus. Distribu tion is 2250 m, 19, 27 -1-82. Id, 54 REVIST A DE BIOLOGIA TROPICAL

Key te Costa Liriomyza species . Rican 1. Mesonotum yellow before seutellum . . . . . 2 Mesonotum entirely dark, blaek or grey, before seutellum 5

2(1). Third antennal segment brownish, greatly enlarged in male commelinae (Frost) Third antennal segment bright yellow, not enlarged in male ...... 3

3(2). AH bristles yellow; mesonotum largely yellow, dark areas at most faintiy reddish-brown...... micr oglossae Speneer All bristles blaek; mesonotum wi th dark areas at least partially blaek . . 4

4(3). 2+1 de; dark areas divided into distinet bands; in female ovipositor normal...... marginalis (Malloeh)

3+1 de; dark area largely solid, outer edges slightiy divided toward seutellum; in female oviposi tor greatiy elongated . . insignis Speneer

5 (1). Third antennal segmen t blaek or at least darkened , brownish 6 Third antennal segment yellow 9

6(5). . Third antennal segment blaek 7 Third antennal segment brownish-yellow 8

7(6). Frons bright yellow zui sp .n. Frons brownish-orange . schmid..tiana ira Speneer Note: Deseribed from San José, La Caja by Speneer (1973a), with illustrations of head and male genitalia.

8(6). Femora bright yellow, at most with slight blaekish striations . . . lupini Spencer Femora more generally blaclásh, though yellowish below . . hu idobrensis (Blanehard)

9(5). Third antennal segment with a fr inge of y,hite pubeseenee y,h ich is longer than basal width of arista ...... 10 Third antennal segment with at most short pubeseenee .. 11

10(9). Mesonotal yellow patches large , inner post-alar on yellow ground; seutellum entirely yellow. . . . costaricana sp .n. Mesonotal patches smaller, inner post-alar on black ; scutellum with small late@1 black trian­ gies ...... archboldi Frost

11(9). Mesopleura largely black .. sabaziae Speneer Mesopleura at most black along lower margin ...... 12

12(1 ). Mesonotum shining blaek sa tivae Blanehard Mesonotum mat, greyish-blaek . . trifolii (Burgess)

Subgenus Poemyza Hendel presumably has a continuous distribution in the mountains from southern California (Spencer, This subgenus is characterized by the high, 1981: 181) through México to Costa Rica. narrow lunule. In Europe 24 species have been recorded and in U.S.A. 10. The subgenus has Genus Liriomyza Mik not previously been known in the Neotropical Region. A single male of Po emyza (Mg.) was Only two species have hitherto been known caught at San José, waste ground near Hotel in Costa Rica (Spencer, 1973a: 153) but the Irazú, 25-1-82. The species is distinctive, with genus is well represented throughout the the orbits partially yellow and the fe mora Neotropical Region. Two new species are yellow on the distal third. The larvae fe ed on described below and 9 species new to Costa many genera of Poaceae. Rica are recorded, inc1uding the three P. muscina was described from Germany but leaf-mining pests L. huidobrensis, sativae and is also widespread in Canada and U .S.A. It trifo lii. It was of considerable interest to SPENCER : Agrornyzidae of Costa Rica 55

discover the host 0f L. insignis, which was reclinate; jowls at rear 1/4 height of eye , this previously only kno�n from caught specimens, only slightly slanting; third antennal segment with the larvae feeding in the flower-heads of srnall, round, with fringe of conspicuous white Bidens pilosa. pubescence with a longer than basal width of arista, this weakly pubescent. Liriomyza archboldi Frost New to Costa Rica Mesonotum: 3+1 strong de, acr in 4 rows.

Liriomyza archboldi Frost, 1962, Entomol. Wing: Length in male 1.6 mm, discal cell News 73: 51-3; Spencer and Stegmaier, 1973: relatively large, last section of vein M3+4 97. almost twice length of penultimate. An uncommon species with frons and antennae yellow, mesonotum shining black, Colour: Frons, orbits, jowls, face, palps and yellow patches a t hind-comers small, inner all antennal segments bright yellow, post-alar on black ; legs with fe mora hind-margin of eye black, with both vertical conspicuously yellow; a distinctive character is bristles on black ground ; mesonotum deep the long pubescence of the third antennal black, shining, yellow hind-co mers large, with segmento The larva has once been recorded as inner post-alar clearIy on yellow ground; forming a blotch mine on Bidens pilosa. scutellum entirely yellow; side of thorax largely The only previous record is fr om Florida. yellow, mesopleura with black triangle in lower Two males were caught on rough ground near front comer, upper margin of sternopleura Hotel Irazú, San José, 31-II-82, on which broadly yellow; legs: coxae and femora bright Bidens was growing. yellow, tibiae and tarsi da rk brownish; abdomen with front tergites yellow laterally; Liriomyza commelinae (Frost) squamae yellowish-grey, margin and fringe N ew to Costa Rica black; halteres bright yellow.

Agromyza commelinae Frost, 1931, Entomol. Male genitalia: Aedeagus as in Figs. 31, 32; News, 42: 72-5. sperm pump with narrow blade but strongly Liriomyza commelinae Spencer, 1973b: 56; pigmented; surstyli with 2 spines. Spencer and Stegmaier, 1973: 101. This is a most distinctive species, with the Holotype: Costa Rica, Nature Reserve, 15 mesonotum yellow before the scutellum and km W. of Atenas,ó, 1-11-82, in AC. the third antennal segment greatly enlarged in the maleo The normal host is Commelina but Remarks: This species gene rally resembles L. larvae have also been found on Tradescantia archboldi but is paler, with the yellow (both Commelinaceae). A long linear mine is mesonotal patches smaller, the dark area ofthe formed, with the black puparium remaining in mesopleura larger and the scutellum having the leaf, lying upside down with the anterior small lateral black triangles in archboldi. The spiracles projecting through the lower aedeagus of the two species is entirely distinct. epidermis. Distribution extends from southem Florida Liriomyza huido brensis (Blanchard) to Argentina, with records from Cuba, Jamaica, New to Costa Rica St. Vincent and Venezuela. A single mine was (Figs. 33-36) found at Santa Ana, 3-11-82, on Co mmelina diffu sa Burm. from which a mal e emerged Agromyza huidobrensis Blanchard, 1926, Rev. 2-1I1-82. Soc. Entomol. Argentina, 1: 10-11. Liriomyza huidobrensis, Blanchard, 1938, An. Liriomyza costaricana sp.n. Soco Cient. Argentina, 126: 356; Spencer, (Figs. 31, 32) 1973b: 57; 1973c: 215; 1981: 24 1. Liriomyza langei Frick, 1951, Pan-Pacif. Head: Frons 1.5 times width of eye, not Entomol., 21: 81. projecting aboye eye in profile ; 2 strong, equal Liriomyza dianthi Frick, 1958, Proc. Entomol. ors, 2 weaker ori; orbital setulae sparse, Soco Wash., 60: 1-5. 56 REVISTA DE BIOLOGIA TROPICAL

This is an important pest spec�es in aboye eye in profile ; orbital bristles variable, California and Venezuela, particularly on the normally 2 strong equal ors, 1 ori but vegetables Beta, Pisum and Vicia fa ba an d the sometimes a weak lower ori is present, less omamentals Aster and Dianthus. There is also a frequently only 1 strong ors and 1 strong ori, in single record on chrysanthemums in Brazil. It 1 specimen seen this arrangement present on is darker and slightly larger than the two more one side only; jowls angular, extended at rear, important pests, L. sativae and L. trifolii. The about 1/3 height of eye, this distinctly slanting; third antennal segment small, round, only finely pubescent.

Mesonotum: 3+1 st rong dc, presutural dc equal to 3rd; acr in 4 irregular rows; no intra-alar.

Wing: Length from 2.5-3.1 in male and up to 3.6 mm in female; discalmm cell large , last section of M3+4 at most 1.5 times length of penuItimate.

Colour: Frons and jowl s yellow,. orbits 29 entirely or at least in upper half black, wi th no yellow patches at hind-corners, moderately shining, scutellum la rgely bright yellow, narrowly black at sides; rear of humerus and notopleura bright yellow, pleura otherwise deep Fig. 29. Amauro my za maculosa: spenn pump. black; legs black apart from bright yellow tips of femora; abdomen largely black but Fig. 30. Nemorimyza posticata: leaf mine on hind-margin of tergites narrowly yellow; unidentified bush (San Ramón). squamae yellowish-grey , margin and fringe black; halteres yellow. third antennal segment is normally brownish, not bright yellow, and both mesopleura and all Male genitalia: Aedeagus (Fig. 37) with femora can be largely black. In the wing the distinctive curvature characteristic of discal cell is relatively large (Fig. 33). The male grass-feeders; surstyli discrete, small with blunt genitalia are shown in Figs. 34, 35. area of sclerotization at end ; sperm pump with The larva forms a characteristic leaf mine, large rounded blade and extended base. normally following the rnidrib and the larger lateral veins. In both the larva and puparium Holotype: 0, Costa Rica, 28-1-82, below the posterior spiracles each have an ellipse of 8 Volcán Irazú near road marker 29, Ruta "km pores (Fig. 36), which permits the species to be 6", 3120 m; paratypes: 1 0, 269, same data. readily distinguished from L. sativae and L. Holotype in AC, paratypes in BM, USNM and trifolú, which both have 3 pores. Departamento de Entomología, F acuItad de No infestation was found on cultivated Agronomía, Universidad de Costa Rica. plants but mines were present on Galinsoga caracasana at Fraijanes, 21-1-82 and on Dahlia Remarks: This species most closely imperialis and Tr opaeolum at Llano Grande, resembles L. schwabei Spencer, 1963 which 26-1-82. A single female was caught on the known only in Chile; this is very similar inis latter host. colour but substantially smaller, with wing length of 2 mm. In the male genitalia of L. Liriomyza irazui sp.n. schwabei the distiphallus is greatly enlarged (Fig. 37) (Spencer, 1982: Fig. 55), confirm: 'lg that the two species are distinct. No comparable Head: Frons about twice width of eye grass-feeders are known either in Venezuela or (slightly shrunk in holotype), not projecting in the United States. SPENCER : Agrornyzidae of Costa 57 Rica

� 35 .. . \. 36

41

37

��::­ ") 42

Figs. 31, 32. Liriomyza costaricana: 31. aedeagus, Fig. 38. Liriomyza microglossae: rnesonoturn. side view; 32. sarne, ventral view. sabaz iae-:- Figs. 39, 40. Lirio myza 39. aedeagus, side view; 40. sarne, ventral view. Figs . 33, 36. Liriomyza huidobrensis: 33. wing; 34. aedeagus, side vie w; 35. distiphallus, ventral Fig. 41. Liriomyza sativae: posterior spirac1es of view : 36. posterior spiracles of pupariurn. puparium. Figs. 42, 43 Liriomyza trifo lii: 42. aedeagus, side Fig. 37. Lirio my za irazui: aedeagus, side view. view; 43. Sarne, ven tral view. 58 REVISTA DE BIOLOGIA TROPICAL

Liriomyza lupini Spencer yellow or the darker bands (Fig. 38) faintly New to Costa Rica reddish-brown. The type specimens were reared from Solidago microglossa at Sao Paulo, Brazil Liriomyza lu pini Spencer, 1981: 247. and the species has also recently been reared from Baccharis sp., near Córdoba, Argentina by This species has hitherto onIy been known G. Valladares. from California where it is not uncommon at A single male was caught at Santa Ana, high altitudes in the Sierra Nevada and known 3-11-82. to feed on four species of Lupinus. The larva forms an irregular, upper surface linear mine. Leaf mines were cornmon on Lupinus ashenbonnii at 2820 m near the restaurant Liriomyza sabaziae Spencer below Volcán Irazú, 28-1-82 but were already New to Costa Rica almost empty. A single female was reared, (Figs. 39, 40) emerging on 19-11-82 and in addition 2 and 1 9 were caught on the pl::nts. ° Liriomyza sabaziae Spencer, 1963 : 366; 1973b: This record represents an important 60; 1981: 262. extension of the range of the species. It is doubtless present elsewhere in Central America ri gi nally described from Brazil and and Mexico at comparable altitudes. Five Venezuelao from Sabazia, later found on Lupinus species are known in Venezuela Galinsoga in Venezuela and more recently (Vareschi, 1970) and the genus extends fu rther discovered to be widespread in California on a south along the Andes. It will be interesting to number of composite hosts, including discover in due course how fa r south L. lupini Baccharis, Carduus, Cirsium, Dahlia, has been able to follow its host plant . Gnaphalium and Silybum. In Costa Rica the two main hosts are Bidens Liriomyza marginalis (Malloch) pilosa and Galinsoga caracasana but mines were New to Costa Rica also found on Dahlia imperialis and Piqueria trinervia. Empty mines on Alomia microcarp a, Agromyza melampyga var. marginalis Malloch, Elvira biblora and Melanthera aspera are also 1913, Ann. Entorno!. Soco Amer., 6: 283. possibly referable to this species. Liriomyza marginalis, Spencer, 1963: 370; Important characters of this species are the 1973b: 58; Spencer and Stegmaier, 1973: 102. largely black mesopleura, combined with the A largely yellow species, with variable dark entirely yellow antennae and the brigh t yellow bands on the mesonotum ranging from shining femora which are narrowly black basally; wing black to reddish brow n; the mesonotum is length is 1.7-2. 1 mm; the aedeagus is shown in broadly yellow ad joining the scutellum. Hosts Figs. 39, 40. are exclusively in the Poaceae, with records from Digitaria, Panicum, Paspalum and Zea Material obtained: San José, waste ground mays. The puparium remains in the mine, with near Hotel Irazú, 2 2 9, sweeping, 21-31-1-82, o, the enlarge d anterior spiracles projecting host certainly Bidens pilosa, on which rnany through the upper epidermis. empty mines present ; Llano Grande, 2000 m, Widespread in the Neotropical Regíon and aboye Cartago, 1 1 9, 26-1-82, empty mines southern Gulf states and Florida. A single male also seen on Dahliao, imperialis and Piqueria was caught on waste ground near Hotel Irazú, trinervia; F raijanes, beside chrysan themum San José, 21-1-82. nursery, below Volcán Poás, 1600 m, 2 3 9, emerged 5-11-82 fro m mines on Galinsogo, a Liriomyza microglossae Spencer caracasana 21-1-82; empty mines seen on G. New to Costa Rica quadriradiata aboye Fraijanes, 6-11-82. (Fig. 38) Mines possibly refe rable he re : Alajuela, garage near airport, Melanthera aspera, 3-11-82; Liriomyza microglossae Spencer, 1963: 363. catarata de Caracha near Alajuela, Elvira A conspicuously pale species, with the biflora, 21-1-82; San José de la Montaña, mesonotum appearing eithe r almost completely Alomia microcarpa, 27-1-82. SPENCER : Agromyzidae of Costa Rica 59

Liriomyzq sativae Blanchard pest in Florida, from where it has spread to New to Costa Rica California, East Africa, most countries in (Fig. 41) Europe, and also Israel. Very large populations are present on chrysanthemums in Colombia Liriomyza sativae BÍancha rd, 1938, An. Soco Cient. Argentina, 126: 354; Spencer, 1973b: and it also causes damage to garlic in 61; 1973c: 219; 1981 : 264. Venezuela. I fo und that there is a large population of L. Liriomyza munda Frick, 1957, Pan-Pacif. trifolii at the two main chrysanthemum Entomol., 33 :61; Spencer and Stegmaier, 1973 : nurseries in Costa Rica at Fraiianes and San 104 (synonymy with sativae establishe d by José de la Montaña. lt is also present on onions Spencer, 1973c). at Santa Ana and on both beans and tomatoes L. sativae is one of the most important leaf at San Rafael de Ojo de Agua . Specimens were mining pests in Latin America, with large populations in Argentina, Brazil, Chile and almost ce rtainly introduced to Costa Rica with Venezuela, and it is widespread in the southern shipments of new varieties of chrysanthemums states of U.S.A., causing serious damage to supplied from U.S.A. in recent years but it is many vegetables, particularly in California and also probable that the species occurs naturally Florida. The species is highly polyphagous but in Costa Rica in small numbers on wild hosts. L. trifolii occurs most fre quent1y on genera of the is readily recognisable by its families Cucurbitaceae, Leguminosae and colour, with the mesonotum being dull Solanaceae. It is a major pest of tomatoes in greyish-black and the hind-margin of the eye Venezuela, Florida and California. largely yellow; the male genitalia, with distinctive curvature at the distal end of the The adult has the mesonotum shining black aedeagus, are shown in Figs. 42, 43. and the hind-margin of the eye largely black, these two characters distinguishing the species Material seen:Frai janes, 1600 m, below from the other widespread Liriomyza pest, L. Volcán Poás, sorne larvae and numerous adults trifolii. The larva fo rms an irregular linear mine, on chrysanthemum, mainly variety Flamenco, with the posterior spirac1es each having 3 pores 21-1-82; San José de la Montaña, 1800 m, aboye (Fig. 41). Heredia, small populations on chrysanthemum, This species is probably widespread in Costa 27-1-82; Santa Ana, large populations wíth both Rica and was noted at 4 localities. larvae and adults on onion, 3-11-82; San Rafael Material seen: San Rafael de Oj o de Agua, de Ojo de Agua, W of Alajuela, small Phaseolus vu lgaris numerous larvae on beans and young tomatoes, populations on and young 3-11-82, producing adults by 28-11-82; at same tomatoes, 3-11-82. locality old mines seen on Cu cumis sativus ' Santa Ana, 3-II-82, empty mines on Capsicur:z Genus Hendel annuum; Alajuela and San Juan de Poás, empty mines on tomato, 25 and 31-1-82 ' San José Over 50 species are known in this genus, waste ground near Hotel lrazú, mines most1; mainly in the Nearctic and Neotropical empty but sorne with larvae on Ricinus Regions. This is the largest genus known in communis, 25-1-82. Venezuela, with 21 species (Spencer, 1973b); only 10 species are known in California (Burgess) (Spencer, 1981). Three species have been known N ew to Costa Rica in Costa Rica. One new species is described (Figs. 42, 43) below and in addition four species are recorded as new to Costa Rica. Oscinis trifolii Burgess, 1880, in Rep. U .S. Identification of species in this genus is Dept. Agríe. fo r 1879 , p.200. difficult but many are host-specific on different Liriomyza trifolii, Frick, 1952, Univ. Calif. plants and the blotch mines fo rmed by the Pubis. Entomol., 8:405 ; Spencer, 1973b: 64; larvae are conspicuous. Fortuna tely the male 1973c: 226; 1981 : 281; Spencer and Stegmaier, gen italia are highly diffe rentiated (cf. Figs. 44, 197 3: 109. 48) and an important generic character is the During the past 5 years L. trifolii has presence of a patch of bristles at the developed in to the most important leaf-mining hind-corner of the epandrium (Fig. 46). 60 REVIST A DE BIOLOGIA TROPICAL

Key to Calycamyza Costa Rican species 1. Third an tennal segment yellow ...... meridiana (HendeD Note : Described fr om AJajueJa as Agromyza meridianalis Malloch, 1914 but with meridianalis preoccupied, renamed meridiana by Hendel (1923, Konowia, 2: 123) and transferred to Calycamyza by Frick (1952, Univ. Calif. Pubis. EntomoJ., 8: 433). The species has since been found in British Guiana, México and Venezuela (Spencer , 1973b) and al so two further locaJities in Costa Rica-San José, La Caja and Surubres. Third antennal segment black ...... 2

2(1). Squamal fringe entirely or partially pale, whitish-yellow .." . verbenivora Spencer Squamal fringe dark, brown or black 3

3(2). Frons brownish 4 Frons yellow . . 5

4(3). Frons 1.5 times wi dth of eye (If yellow, see couplet 7) .. devia Spencer Frons narrower, equal to width of eye ...... obscwa Spencer Note: Described by Spencer Spencer and Stegmaier, 1973: 87 fr om Florida, Jamaica and San José, La Caja. This species cJoselyin resembles the dark fo rm of dev ia (couplet 3) but the male gen italia of the two species are quite distinct. e

5(3). Third antennal segment with fringe of distinct pubescence, at lea

6(5). Orbits conspicuously black a t least to lower ors 7 Orbits dark only to upper ors 8

7(6). Femora entirely black ...... devia Spencer Femora bright yellow at apex . .. addita sp.n.

8(6). Orbits black to upper ors .. sidae Spencer Orbits entirely yellow or at most slightly brownish to upper ors hyptidis Spen cer

Calycomyza addita sp.n. yellow apically; squamae yellowish white, (Figs. 44-47) margin and fringe black; halteres white.

Head: Frons narrow, equal to width of Male genitalia: Aedeagus (Figs. 44, 45) e ye , narrowly projecting aboye eye in with distiphallus enlarged, not divided; profile ; 2 ors, 2 ori, orbital setulae sparse, mesophallus narrowing adjoining distiphallus; reclinate; jowls l/S height of eye ; third anterior ventral lo bes weakly pigmented, antennal segment small, round . with short, indistinct, posterior lobes asyrnmetrical, with normal pubescence; arista long, equal to a narrow, strongly pigmented sclerite vertical height of eye, distinctly pubescent. between; hind-corner of epandrium (Fig. 46) with characteristic patch of strong bristles. Mesonotum: 3+0 dc, third small; acr in about 8 rows. Host: Unco nfirmed, possibly Melampodium divaricatum (Fig. 47). Wing: Length in male 1.9 mm ; discal cell relatively large, last section of vein M Holotype: Costa Rica, San José, waste ó, 3 + 4 just over twice �ength of penultimate. ground near Hotel Irazú, 31-1-82, in AC. A female caught at the same locality, 21 -1-82, Colour: Frons, jowls and face bright possibly represents the same species but yellow; orbits black to lower ors; all differs in the less dark orbits, the largely antennal segments and palps black; black fo re-femora and the less numerous mesonotum and scutellum moderately acrostichals and is not treated as a shining black; notop leura and upper paratype. hind-corner of mesopleura bright yellow; Whitish blotch mines · (Fig.47) were not legs black but femora on fore-legs narrowly uncommon on Me lampodium divaricatum SPENCER : Agrornyzidae of Costa Rica 61 growing where this species was caught. Spencer and Stegmaier, 1973: 74; 1973b: Sorne larvae .were obtained but none 42. produced adults and the association of e. The larvae of this species form a addita with these mines seems probable but characteristic blotch mine with irregular requires confrrmation from reared adults. o ffshoots and the frass deposited in the centre of the mine (Fig. 51). One mine Calycomyza caguensis Spencer, 1973b was found on Hy ptis capitata at the Nature New to Costa Rica Reserve west of Atenas, l-II-82. (Fig. 48) Although the adults c10sely resemble e. malvae, the male genitalia are of different This species has hitherto only be en form, with the distiphallus ending in long known from a single male from Venezuela. paired tubules (cf. e. caguensis, Fig. 48) in A further male was caught at the roadside C. hyptidis, while in e. malvae the near Chitaría, east of Turrialba, 29-1-82. aedeagus gene rally resembles that of e. The aedeagus agrees exactly with that of devia (Figs. 49, 50) and e. sidae (Figs. 52, the holotype (Fig. 48). 53). A number of species are known with rather similar elongated tubules of the Ca lycomyza sidae Spencer distiphallus (e. ecliptae Spencer, 1963, host New to Costa Rica Eclipta; e. hyptidis Spencer, 1966a, host (Figs. 52, 53) Hyp tis; e. verbenivora Spencer, 1963, host Verbena; e. verbesinae Spencer, 1963, host Calycomyza sidae Spencer and Stegmaier, Verbesina). However, small differences in 1973: 88; Spencer, 1973b: 48. t he e x t e rnal characters of the adults, This species is widespread in the associated with the different hosts and Caribbean area, is common in sou them differences in the leaf mines confirm that Florida and has also been recorded in the species are distinct. Venezuela and Brazil. The specimen fr om Río de Janeiro was originally misidentified Calycomy za devia Spencer, 1973b as e. malvae by Spencer, 1963 and the (Figs. 49, 50) illustrations of the genitalia (Figs. 52, 53) were later used in the description of e. This species was described from two sidae. localities in Venezuela and it was found, Leaf mines on Sida sp. found at the following examination of male genitalia, roadside near Chitaría, east of Turrialba, that one of a series of 20 specimens from 29-1-82 are referred to e. sidae. The mines San José, La Caja, which were identified as are generally shorter than those of e. C. obscura (Spencer and Stegmaier, 1973: malvae (sometimes even blotch-like). It is 86) in fact represented e. devia. The two hoped that confi rmation of this species have in common the brownish frons identification can in due course be obtained and darker than normal notopleural area from adults fr om this locality. but are distinguishable by the broader frons in C. devia. Calycomyza verbenivora Spencer A fu rther male was obtained at the New to Costa Rica Nature Reserve west of Atenas, l-II-82. In this specimen the frons appears almost Phy to bia (Calycomyza) verbenivora Spencer, yellow but the orbits are conspicuously 1963: 350. black. The male genitalia agree exactlv with Ca lyeomyza verbenivora Spencer, 1973b: 49; those of the holotype (Figs. 49,50). Valladares, 1982. This species was described from Caracas, Calycomyza hyptidis Spencer Venezuela from leaf mines on Ve rbena New to Costa Rica litoralis. It has also recently been found on (Fig. 51 ) this host at Córdoba, Argentina and also on V. bonariensis and Glandularia hybrida by Calycomyza hyp tidis Spencer, 1966a: 16; Valladares, (1982). REVIST A DE BIOLOGIA TROPICAL 62

45

Figs. 4447. Ca lycomyza addita: 44. aedeagus, side Fig. 49, 50 Calycomyza devia: 49. aedeagus, side view ; 45. distiphallus,ventral view ; 46. epandriurn and view; 50. same, ventral view. sl\rstylus; leaf mine on Meiampodium 47. Fig. Calycomyza hyptidis : leaf min e on divaricatum. 51. Hyp tis pectinata (Florida)

Fig. Calycomyza caguensis: aedeagus, side view 48. Figs. 52, 53. Calycomyza sidae: 52. aedeagus , side (Venezuela). view; 53. same, ventral view (Brazil). SPENCER : Agromyzidae of Costa Rica 63

Key Costa Rican Phytoliriomyza species to l. Outer cross-vein lacking ; minute species , wing length 1.1-1.25 mm in maJe . conjunctimontis (Frick) Note: Described by Frick (1952, Univ. Calif. Pubis. Entorno!., 8 :41 3) fr om California and sUbsequentIy recorded from Dominica and Costa Rica, San José, La Caja (Spencer , 1973a). A further male has now been seen fr om San José, waste ground near Hotel 1razú, 21-1-82. The species is distinctive both in the lack of the outer cross-vein and in its small size , the specimen now seen having wing length of 1.1 mm. Outer cross-vein presen t; sligh tly larger species ...... 2

2(1). Tlúrd antennaI segment with a fringe of long pubescence 3 Third antennaJ segment with normal short pubescence .. 4

. pilosella Spencer 3(2) Third antennal segment yellowish ...... Note : Described by Spencer in Spencer and Stegmaier , 1973 : 116 fr om Puerto Rico, Florida and San José, La Caja (6 169). Although cJosely resembling P. jurgensi sp.n., the aedeagus with the long coiled distiphalld, us (cf. Spencer and Stegmaier, Figs . 297) show that the two species are not cJosely related (contrast Fig. 57 below)...... jurgensi sp.n. Tlúrd antennal segment largely black '"

. . costaricen sis (Spencer) 4(2). All antennaI segmen black ...... Note : Described fr om15 3 males from San José, La Caja by Spencer in Spencer and Stegmaier (1973: 188). The distinctive aedeagus and epandrium are sho\Vn in Figs. 54, 55. This species was described in the genus Lemurimyza, which \Vas synonymised with Phytoliriomyza by von Tschirnhaus (1971, Beitr. Entorno!., 21 : 563). Third antennaJ segment first and second more yellowish .. imperfecta (Malloch) brownish-yellow,

Leaf mines, mostly empty, were not This species has previously been known uncommon on Verbena litoralis near Hotel from Chile, Florida and California. Two Irazú, San José, 24-1-82. The larva fo rms a specirnens have now been seen from Costa linear mine in the fr rst instar but later Rica: San José, waste ground near Hotel produces a blotch, which quickly turns 1razú, one male, 31-1-82; Paso Llano, aboye brown. Unfortunately no adult was obtained Heredia, 1900 m, one female, 27-1-82. from the single larva found. The elongated, paired black tubules of the distiphal!us (Fig. 56) are distinctive . Genus Phyto liriomyza Hendel Phy toliriomyza jurgensi sp.n. Two species have hitherto been known (Figs. 57, 58) in Costa Rica. One new species fr om San Head: Frons broad, twice width of eye; José is now described below and a fu rther 2 ors, 1 ori (missing one side in holotype); species, P. imperfec ta (Mall.) which is orbital setulae virtually lacking, reduced to wi despread from Chile to Florida and 1 or 2 below ori; jowls narrow, 1/5 height California, represents a new record. of eye, this large, only slightly slanting; Throughout the Neotropical Region 15 third antennal segment small, round, wi th species are known, 6 of which were f ringe of conspicuous pu bescence ; arista recently described from Chile by Spencer long, dropping, distinctly pubescent, equal (1982). Only three species are known in in length to height of eye. Venezuela (Spencer, 1973b). Mesonotum: 3+1 de, acr sparse, in 2 Phytoliriomyza imperfecta (Malloch) rows. New to Costa Rica (Fig. 56) Wing: Length in male 1.4 discal mm; cel! relatively large, last section of M3+4 Agromyza imperfecta Malloch, 1934: 475, slight ly less than twice length of Agromyzidae in Diptera of Patagonia and penultirnate. South Chile, British Museum. Phy toliriomyza imperfecta, Spencer and Colour: Head yellow, apart from black Stegmaier, 1973: 115; Spencer, 1981 : 324. third antennal segment and palps; 64 REVISTA DE BIOLOGIA TROPICAL mesonotum predominantly pale grey but Genus Phytomyza Fallén central band between line of dc brownish; scutellum greyish, with faint yellow This large, predominantly north undertone; side of thorax largely yellow, temperate genus has hitherto be en with only central area of humerus, a small represented in Costa Rica by the single bar at upper margin of mesopleura and species P. loewii Hd. Five species only are lower half of sternopleura grey; legs: coxae known in Chile (Spencer, 1982) and one and femora bright yellow, tibiae and tarsi known only fro m leaf mines on Daucus brownish-black; squamae yellow, margin and (Umbelliferae) in Venezuela (Spencer, fr inge dark; halteres yellowish below, knob 1973b: 75). Throughout the world over distinctly darkened aboye. 450 species are known and in the United Male genitalia: Aedeagus divide d distally, States 49 . cup-shaped at end (Fig. 57); sperm pump Empty leaf mines (Fig. 59) were found mi n u te, b la de weakly pigmen ted, base on My n-hidendron donnell-smithii, a tall , slightly extended on one side : no apparent distinctive species of Umbelliferae over 2 m sursty1i but inner margin of epandrium in height , growing beside the road from the bearing about 15 short, stout spines (Fig. rim of Volcán Poás to the parking area, 58). 22-1-82. Many species occur on Umbelliferae Holotype: Costa Rica, San José, waste but this is certainly undescribed and it is ground near óHotel Irazú, 31-1-82; paratype to be hoped that adults can be obtained in same locality, 3-II-82, both in AC. due course. ó Key to Costa Rican Phytomyza species

1. Host: Clematis (Ranunculaceae); entirelY black species ...... loewii (Hende!) Note: The holotype of Phy tomyza centralis Frost fr om San José (1963, Ann Entorno!. Soco Amer., 29: 317) has been examined and this species is now synonymised with P. lo ewii, syn. nov. Known also from one male fram San José, Caja (Spencer and Stegmaier, 1973: 192). The larva forms a short linear mine on Clematis spp.La Recorded from Quebec, D.C. and California in U.S.A., also Cuba. Host: Myrrhidendron (Umbelliferae) ; adult unknown sp ...... (My rrhidendron).

Genus Chromatomyia Hardy 73). In U.S.A. 27 species are known . New to Costa Rica Leaf mines with puparia (Fig. 60) were found on the grass Brachiaria mu tica beside This is also an essentially north the airport near the coast at Limón, temperate genus with approx. 100 species 29-1-82. Unfortunately both puparia were now known. On external characters the parasitized and no adults were obtained. adults cannot be separated from Phyto myza The species is certainly undescribed. but the distinctive method of pupation, with the puparium lying upside down and Leaf Mines of unidentified species the anterior spiracles projecting ventrally through the epidermis and particularly the 1. Bomariea (Amaryllidaceae), aboye form of the male genitalia, have be en Dano GrandeSP . in gu11y on E side of treate d as of generic significance in road, 2280 m, 26-1-82. Many mines revIslonary stu dies of this group by found, a11 with dead larvae, mine white, Griffi ths (1974, Quaest. Entomol., 10: narrow, normally closely fo11owing a 35-69 and 1980, Entomol. Scand. Suppl. vein (Fig. 61). 13: 1-61). Almost certainlv a Liriomyza sp.n., Only 3 species have hitherto been known related to L. smilacinae Spencer, known in the Neotropical Region (e. platensis from western Canada and California, Brethes feeding on Labiatae in Argentina feeding on Smilacina spp. (Liliaceae). ( and Chile, cf. Spencer, 1982 and two 2. Browallia demissa L. = americana L.) species known in Venezuela originally (Solanaceae), San José, near Hotel placed in Phy to myza, cf. Spencer, 1973b: Irazú, 24-1-82 and 15 km W of San SPENCER : Agrornyzidae of Costa Rica 65

Ramón, l-I1-82. Irregular white linear 7. Stachytarpheta sp. (Verbenaceae), 15 mines (Fig. 62), all emptv. Probably a km E of A tenas, Nature Reserve, Liriomyza sp.n. 1-II-82. Narrow, irregular, white linear 3. Conyza canadensis (L.) Cronquist mines, all empty. Probably Liriomyza (Asteraceae), below Volcán Poás, 1950 sp.n. m, 6-Il-82. Short, white linear mine, 8. Ve monia canescens HBK, (Asteraceae), possibly Liriomyza sp.n. 15 km W of Atenas, Nature Reserve, ( 4. E lvi ra martyni Cass. = biflora DC) 1-II-82. Blotch mine , empty. Probablv (Asteraceae), Catarata de Caracha, N of Calycomyza sp. Alajuela, 21-1-82 . White linear mines, in 9. Vi guiera sylvatica Klatt (Asteraceae), 15 part following veins, aH empty, possibly km W of San Ramón, 1-II-82. Both Liriomyza huidobrensis. blotch mines and linear mines, empty, 5. Sa lvia sp . (Labiatae), aboye Llano fo rmer probably Calycomyza sp., latter Grande, 2280 m., 26-1-82. Linear mine, Liriomyza sp. with the channels short and adjoining, 10. Genus, sp. indet.. large-leaved species, fo rming a secondary blotch. Probab1y (Asteraceae), aboye Llano Grande, Liriomyza sp.n. Cartago, in gully on E si de of road, 6. Solanum nigrum L. (Solanaceae), 2280 m, 26-1-82. Long, white linear Fraijanes, 1600 m, 21-1-82. Irregular mines, sorne with larvae (Fig. 63). linear mines, all empty. Possiblv Probably Liriomyza sp.n. Liriomyza sativae.

� ""\." .>.

'5j(l l ��-�7 t•�.. -�> 55 J58

ACKNOWLEDGMENTS

Figs. 54, 55. Phytoliriomyza costaricensis: 54. I wish to take this opportunity of a edeagu s, view; 55. epandriurn. ventral formally thanking the GTZ for sponsoring my visit to Costa Rica at short notice and Fig. 56. Phytoliriomyza imperfecta: aedeagus (San José). the American Flower Corporation for their generosity in covering my living expenses Fig. 57, 58. Phytoliriomyza jurgensi: 57. aedeagus, throughout my stay. side view; 58. epandriurn. Ing. Juan José May, Dr. G. J ürgens and Dr. G. Vaagt of the Convenio Costarricense- 66 REVISTA DE BIOLOGIA TROPICAL

Alemán provided constant help with many San José and Sylvia Gould at the matters and the hospitality received was Department of Botany, British Museum, greatIy appreciated. My special thanks are Natural History, proved invaluable and is due to Dr. Jürgens whose initiative was gratefully acknowledged. instrumental in arranging my visit. 1 must also thank Sres. C. Sanabria and Assistance provided by Sanidad Vegetal, L. Zamora fo r their willing assistance on MAG, with transport, and in particular the our frequent field trips. loan of a microscope by Sr. Alberto Freer, Finally, must thank Sr. Jorge Ramírez and the identification of plants by Luis for kindly 1agreeing at short notice to act Diego Gómez at the Museum HerbariumDr. in as interpreter during my lecture.

!;:

62 63 60

Figs. 59, 63. Leaf mines : 59. Phytomyza sp. on descu biertas 9 especies nuevas de My rrhidendron donneIl.smithii, Volcán Poás ; 60. Chromatomyia sp. on Brachiaria mutica, Limón; Agromyzidae que fueron descritas en los 61. Liriomyza sp. on Bomariea sp., abo ve llano géneros Me la nagromyza, Ophiomyia, Grande; 62. Liriomyza sp. (?) on BrowaIlia Japanagromyza, Liriomyza y Phytoliriomyza. demissa, San José; 63. Liriomyza sp. on gen., También se encontró 18 especies nuevas (? ) sp. indet. (Asteraceae) above Uano Grande. para Costa Rica incluyendo 3 que son plagas de importancia económica: Liriomyza RESUMEN huidobrensis, L. sativae y L. trifolii. Se En un estudio de 3 semanas en Costa r.olectó llÚnas en hojas de 10 plantas: 2 Rica entre enero y febrero de 1982, fueron representan especies hasta ahora no descritas SPENCER : Agromyzidae of Costa Rica 67

6 probablemente especies no descritas. Se Spencer , K.A. 1973b. The Agromyzídae (Díptera) ypr esenta claves para 13 géneros 53 espe­ of Venezuela. Rev. Fac. Agron., Maracay. cies hasta ahora conocidas en Costay Rica. 7 :5-108. Phy tomyza centralis Frost se coloca en si­ Spencer, K.A. 197 3c. Agromyzidae (Diptera) of nonimia con Phy tomyza loewii Hendel. economic importance. Series Entorno!. 9:1-418. DI. W. Junk, The Hague.

Spencer , K.A. 1977. Notes on world Agromyzidae, with the description of 16 new species. Beitr. Entorno!., Berlin, 27 :233-253. UTERATU CITED RE Spencer, K.A. 1981. A revisionary study of the Spencer, K.A. 1963. A synopsis of the leaf-mining (Agromyzidae) of California. Neotropical Agromyzidae. Trans. Roy. Entorno!. Special Pub!. 3273: 1-489. Div. Agric. Sciences, Soc., London, 115:291-389. University of California.

Spencer, K.A. 1966a. New and interesting Spencer, K.A. 1982. Agromyzidae (Diptera) in Agromyzldae fr om Florida. Stuttgart Beitr. Chile. Stuttgart BeitI. Naturk. Staatlich. Mus., Naturk. Staatlich. Mus., 158: 1-20. 357:1055.

Spencer, K.A. 1966b. Notes on the Neotropica1 Spencer , K.A. & C.E. StegmaieI. 1973. The Agromyzidae of Florida, with a supplement on A�omyzidae . Papeis Avulsos Dept. Zoo!. Sao Paulo 19:141. 1-150. species fr om the Caribbean. of Florida and neighboring land areas 7: 1-205. Spencer, K.A. 1967. A catalogue of the diptera FIa. Dept. Agric., Gainesville. of the Americas South of the United States. Valladares, G. 1982. Contribución al conocimiento 83. Family Agromyzidae pP. 1-23. Dept. Zoo!. las especies de Hendel (Diptera,de Sao Paulo. Ca Iycomyza Agromyzidae), minadores de hojas en la Argentina. Rev. Soco Entorno:. Argent., 40 :221-229. Spencer, K.A. 197 3a. Agromyzidae (Diptera) in Costa Rica. Beitr. Entorno!., Berlin, 23: 151-156. Vareschi, V. 1970. Flora de los páramos. University Press, Mérida, Venezuela , pp. 1-429.