© 2010 National Research Coullcil ofCanada AlJ rights reserved. No part 01' this pubJicatíon may be reproduced in a retrieval system, or transmitted by any means, electronic, mechanical, photocopying, recording or otherwise, without the prior written permission 01' the Natíonal Research Coullcil 01' Canada, Ottawa, ON K 1A OR6, Caoada. Printed 00 acid-free paper. §l

ISB N 978-0-660-19958-0 NRC 49739

l. CIP for print publicatíon:

Library and Archives Canada Cataloguing in Publication

Manual 01' Central American Diptera. Volume 2 / edited by S.v. Brown ... [et aL].

!ssued by: Natiooa! Research Counci I Canada. Ineludes bibliographical references and indexo ISBN 978-0-660-19958-0 (bound)

l. Diptera--Central America--Classificatioo. 2. Diptera--Central America --Identification. 3. F/ies--Central America--Classification. 4. --Central America--Identi fication. 1. Brown, Brian Victor 1\. National Research Couoci]

QL535.2.C35 M36 20 IO 595.7709728 C20 l0-980031-]

NRC Monograph PubJishiog Program

Editor: PS. Cavers (University ofWestern Ootario)

Editorial Board: W.G.E. Calclwell, OC, FRSC (University 01' Western Ontario); M.E. Cannon, FCAE, FRSC (University 01' Calgary); K.G. Davey, OC, FRSC (York University); M.M. Ferguson (University of Guelph); S. Gubios (Aonual Reviews); S.K. Hall, FRSC (Dalhousie University); P. Hicklenton Ph.D, P.Ag.; W.H. Lewis (Washington University); A.W. May, OC (Memorial University 01' Newfoundland); B.P Dancik, Editor-in-Chief, NRC Research Press (University 01' AJberta)

Inquiries: Monograph Publishing Program, NRC Research Press, National Research Council 01' Canada, Otlawa, Ontario K J A OR6, Canada. Web site: http://pubs.nrc-cnrc.gc.ca

Correet citatíon for this publication: Brown, B.V. el af. 2010. Manual ojCenlraf American Diptera: Vofume 2. NRC Research Press, Ottawa, Ontario, Canada. 728 pp. A Publication of the National Research Council of Canada Monograph Publishing Program

anual of eral American D·ptera

Vo unte 2

Edited by

B.V. Brown Natural History Museum of Los Angeles County, Los Angeles, California, USA A. Borkent Royal British Columbia Museum, Victoria, British Columbia, Canada J.M. Cumming Agriculture and Agri-Food Canada, Ottawa, Ontarío, Canada D.M. Wood Agriculture and Agri-Food Canada, Ottawa, Ontario, Canada N .E. Woodley USOA Systematic Entomology Laboratory, Washington, OC, USA M.A. Zumbado Instituto Nacional de Biodiversidad, Costa Rica

,.;:,c·c,.;:,c NRC Research Press OUawa 2010 YRGOTIDAE (PYRGOTID FLIES) 69

cente Hernández-Ortiz

''hl

Fig. 69.1. Female of Pyrgola undala Wiedemann, (Nearctic, MND, fig. 65.1).

Diagnosis 1+2 elongate, usually with several long hairs on lateral sides near base. Female terminalia (Figs. 8-12) with syntergoster­ Medium to large-sized flies (body length 5-30 mm), nite 7 robust and strong basally forming conical tube turned body uSllally robust with long legs, wing pattern often spot­ ventrally, sometimes longer than rest of abdominal length. ted, banded, 01' reticulated, rarely hyaline (Fig. 1). Head Apical margin of syntergosternite 7 sometimes provided (figs. 2-7) spherical with fronslarge, antennal bases project­ with ventral hooklike sclerite, 01' two apicolateral clawlike eel anteriorly, pedicel elongate; arista setiform; ocelli usually sclerites (according to Steyskal, 1972, probably derived from lacking in American species (except in Descoleia Aczél); tergum and sternum 8); aculeus uSllally sborter than length frons weakly depressed, frontal bristles and vibrissa absent, ofsyntergosternite 7. Male terminalia similar to Tephritidae face broad, with deep antennal grooves; mouthparts of mod­ and Platystomatidae; epandrium globose, curved ventrally; erate size. Scutum mostly bare, setulose, 01' distinctly haired, lateral surstyli usuaJly short and robust basally, medial sur­ sometimes black spotted, major bristles usually shol1er than styli without prensisetae developed; aedeaglls coiled and in Tephritidae; scutellul11 short and broad with one to four eJongated; glans more 01' less developed and partially sclero­ pairs of margj nal bristles. Legs (Fjgs. 13-15) long in botb tized in some species. sexes, frequently without strong bristles, but covered by short are most similar to certain members of the setulae; bind tibia tapering to base 01' with strong constriction fami Iy Conopidae by the similar shape of the body, and par­ in basal third 01' near midlength. Wing (Figs. 16-23) with ticularly the bead shape, in addition to the absence of ocelli

,; (subfamily Conopinae). However, tbe Pyrgotidae possess 01' without subcostal break, C weakening after vein R4T Sc complete to C or disappearing c10se to C; R, dorsally setu­ a normally developed, sponging proboscis, and veins R4>5 + and M + are not convergent, ending well separated at the lose; R2 3 sometimes with stUl11p vein at apical third; cel! cup I 2 wing margino In contrast, Conopidae possess a long (usually closed by CuA2 usually forming acute angle in 10wer api­ geniculate) proboscis, and veins R and M + are strongly cal comer, sometimes short. Preabdomen elongate, robust, Hj I 2 more al' less petiolate; lateral membrane broad; syntergite convergent apically, and sometimes fused distally.

955 956 MANUAL OF CENTRAL AMERICAN DIPTERA ­ VOLUME 2

Biology their crepuscular or nocturnal habits that are similar to those ofmost oftheir hosts. Sorne other species, such as theAustra­ Little information is available on the biology 01' the Pyr­ lian beetle Anop/ognalhus olivieri (Dalman) and its pyrgotid parasite Maenomenus ensifer Bezzi (with ocelli developed). gotidae, although it is welJ known that larvae are endopara­ have diurnal habits (Aczél, 1956a). Probably nocturnal or sitic on adult scarabaeid beetles (Coleoptera: Scarabaeidae). diurnal activíty explains the presence or absence of ocelli The highly specialized morphology ofthe female termínalia throughout the fami Iy. suggests that all pyrgotid species are parasites. For CentraJ and South America, there is no bioJogical in­ Forbes (1908) described the oviposition behavior of formatíon on this family. Most life history data are for some Pyrgola undala Wiedemann temales. He observed the flies North American species, such as Pyrgola undala. waiting on the body 01' Phy//ophaga adults until the beetles Sorne work has be done on the val ue 01' pyrgotids as con­ opened their elytra before flying. The female deposited an trol agents 01' pest scarabaeids in the New World. Pyrgo!a egg (or larva?) into the abdomen ofthe host while both were undala was introduced in Puerto Rico as a potential conlrol of sorne injurious Phy//ophaga species (Wolcott, 1923): in l1ight. The life cycle ofP undala was described by Davis Pe/lodasiafiaviseta (Aldrich), a pyrgotid native to the Ori­ (19 J9), who found that female beetles are more frequentJy ental Region, was introduced into the USA for the control used as hosts. Parasitized beetles live 10-14 days, with only 01' the Japanese beetle, Popi/lia japonica (Gardner & Park­ one larva developing in each host. The pupates within the er, 1942). In both cases results were not satisfactory. More host body, and developmental time from oviposition to pupa­ recently, Crocker et al. (1996) conducted an evaluation 01' tion is about three weeks. the fI ight period and egg production of P undala, wilh re­ Although both males and females are attracted to Iight, spect to seasonal occurrence 01' pest Phy/lophaga species aduJt Pyrgotidae are uncommon in collections, because of in Texas.

2 Leplopyrgola d' 3 Leptopyrgola º 4 ldiopyrgola º

5 Pyrgola º 7 Slenopyrgola 9

6 Anapyrgola d'

Figs. 69.2-7. Heads: lateral view of (2) Leplopyrgola undulara Becker; anterior vjew (antennae omiued) of (3) L. albilarsis AC/él. (Soulh America, Aczél, 1956b, fig. 29); and (4) ldiopYlgola seliven!ris Aczél, (South America, Aczél, I956c, fig. 78); lateral view of(S) PY/'¡¡Ola lugens Wulp; (6) AnaPYl'gola sp.; and (7) S!enopyrgo!a sp. Figures 2 and 5-7 illustrated by Y. Hemández-Ortiz. PYRGOTlDAE (PYRGOTID FLlES) 69 957

Classification sternum and tergum 6 in males, synsternite 7 + 8 absent, phallus extremely short, female sternllm I largely desclero­ ~ormerly this family was classified in three subfami­ tized, aculeus silort and blunt, and two rudimentary cerci lies: Lochmostyliinae, Pyrgotinae, and Toxurinae (Aczél, bearing setae. 1956a; 1958; McAlpine, 1978; Steyskal, 1987), a11 of rdistinguish two subfamilies in Pyrgotidae: Teretrurinae them represented in the New World. In a recent treatment, (formerly Teretrurini), cilaracterized by sternites 1 and 2 hOll'evcr, McAlpine (1989) proposed a different subfamily fu11y developed and free, and Pyrgotinae (including Tox­ arrangement. The Ctenostylinae (=Lochmostyliinae) was mini) with sternites 1 and 2 fused into one synsternite. The removed from the Pyrgotidae as the family Ctenostylidae, subfamily Teretrurinae has an exclusively sOllthern Neo­ )uggesting that further studies cou Id support its exclusion tropical distribution with the genera Pyrgotosoma Malloch fllJm lhe . This was based on the we11-developed and Teretrura Bigot recorded from Argentina and Chile.

9 Sphecomyie/la Q

'.-­

lO Idiopyrgota Q 8 Sphecomyie/la Q li Undescribed Genus A

", 1,' ,1 /1 11 ':

12 Undescribed Genus A Q 13 Pyrgola cJ' i4 Stenopyrgota cJ' 15 Anapyrgola cJ'

H~ . 69.11-15. Female abdomen, terminaJia, and legs: lateral view 01' abdomen, ovipositor extended, with offset of hooked apex of ,,11il~ 7 in ventral view (8), and dorsal view of aculeus, or sclerolized apical portion of ovipositor (9) of Sphecol11yie/la p(JIida (Harris), . D. figs. 65.8-9); lateral view of apex of abdomen and syntergosternite 7 of (10) {diopyrgo/a sp.; ventral (1 \) and lateral (12) view of .orsyntcrgosternite 7 of Undescribed Genus A; posterior view of left hind tibia of (13) p¡wgola {¿¡gens Wulp; (14) SlenOp)'Igo/a sp.; and '1.llIlI{'.l'rgo/a sp. Figures 10-15 j lIustrated by V. Hernández-Ortiz. hhrcviations: ovpung, ovipositunguis; sg, segment; tg, tergite. 958 MANUAL OF CENTRAL AMERICAN DIPTERA - VOLUME2

The subfamily Pyrgotinae includes two tribes: Toxurini Differences between Slirolhrinax EnderJein and Sphcco­ with a mainJy Auslralian distribution, but also represented myiella could not be established based on currently kno\\ll in the Americas by lhe genus Descoleia, with one species, species, as weH as additional studied specilllens that repre· D. lerelrllra Aczél from Argentina; and Pyrgotini, which sent undescribed species. For this reason, Slirolhl"inux lVill is well represented in lhe NeotropicaJ Region. In my eventually be considered a synonym of Sphecolllyieffa, as sludy of lhe fauna of Central America, 1found the genera also noted by Kondratieff & Fitzgerald (I 993). Much lVork Anapyrgola, Boreolhrinux, Idiopy¡gota, Leplopyrgola, is necessary to gain a better understanding of the rclalioll­ Neopyrgola, Pyrgola, Sphecomyieffa, and Stenopyrgota ships among pyrgotid genera. to be present, in addition to three undescribed genera. Recently, Bernardi (1990) proposed the synonymy of Anapyrgola with Carrerapyrgola Aczél mainly based on the Identification absence ofthe subscuteJlum. However, his proposal does not discuss characters supporting the segregation of Carrerapy­The most comprehensive works for the recognilion 01' rgola by Aczél (1956), which are missing in Anapyrgola. NeotropicaJ taxa were pubJished by Aczél (1956a, I956b. Such characters include the costal vein ending almost at the J956c). He described, illustrated, and keyed most ofthc spe· level of vein R",; and lhe last section of vein M being barely eviclent after vein dm-cu, disappearing before the wing mar­ cjes, except those in lhe genera Sphecomyiella and Borco­ gin. In addition, the discal cell is shorter and its lower Ihrinax, which are treated by Steyskal (1978, J987). apical corner ends distant from lhe wing margin, and there Some genera cannot be presently recognized in the maje are other outstanding djfferences in wing pattem. Therefore, sex (see couplel 7). Further sludy of male terminalia is Ilec­ in the present study this synonymy is not accepted. essary to remedy lhis situation.

Key to the genera of Pyrgotidae of Central America

l. Face without median keel, 01' antennal grooves complete!y joined (Fig. 3); ifpresent, median keel weak 01' poorly dilTerentiated; alula indistinct to well developed. . 2 Face with antennal grooves separated by strongly developed median keel (Fig. 4); alula well developed...... , . . . .. 6

2. Alula well developecl (Figs. 16, 18-21); several pairs of scutellar bristles usually present .... J Alula greatly reduced 01' absent (Figs. 17,22); scutellar bristles usually absent, 01' sometimes with only short pair of sJender apical bristles present ...... 5

3. Subscutellum indistinct 01' absent; wings with brownish bands well dilTerentiated by hya!ine areas; legs normally clevelopecl; female ovipositor sheath with neither apicolateral ovipositungues R nor apicoventral unguis; Stulllp vein in 21l present (Fig. 21) Anapyrgola Steyska! Subscutellum well developed; wings mostly hyaline, with dilTuse pattern, 01' hyaline spotted; legs various; remale ovipositor sheath with two apicolateral ungues 01' one apicoventral unguis; stump

+ vein in R2 i usually absent ...... 4

4. Wing broadly hyaline 01' witll slightly infuscate pattern (Fig. 18); legs e10ngated and slender with hind tibia constricted near basal half; female ovipositor sheath with two c1aw-shaped apicolateral ungues (Figs. 11-!2) ...... Undescribed GenusA Wing brownish, hyaline spotted; legs normally developed with hind tibia tapering gradually to base; female usualJy with hook-shaped apicoventral unguis (as in Fig. 8) ... Neopyrgola Hende!

5. Vein R ,) straight throughout length, becoming paralJeI with respect to .,; alula completely 2 R1 absent; wing pattern irrorate, with small hyaline spots throughout surface (Fig. 22); pedicel usual/y sllorter thanAagellum; head strongly depressed anteriorly, triangular shaped in lateral view (Fig. 7) Slenopyrgola Malloch

' R~_,; Vein R2 1 strongly sinuate on apical one-third, becoming convergent with alula greatly reduced; wing mostly hyaline usually with large anteroapical dark spot (Fig. 17); pedice! elongate, usually lOllger than fiagellum; head not as aboye (Fig. 2) .... Leplopyrgola Hendel PYRGOTIDAE (PYRGOTID FLlES) 69 959

6. Hind tibia tapering to base (as in Fig. 13)...... 7 Hind tibia strongly constricted near basal one-third (as in Fig. 15) . . 10

7. Abdominal sternites 3 and 4 in femaJes each with comb-shaped row of strong long bristles; syntergosternite 7 lacking apicoventral unguis (Fig. 10); ocellar and postocellar bristles usually well differentiated (Fig. 4)...... Tdiopyrgolo Aczél Abdominal sternites in fe11lales without any distinct strong bristles; other characters various .. 8

8. Syntergosternite 7 without apicoventral hook-shaped unguis; body dark brown to black; wing pattern brown with extensive irregular hyaline spots Undescribed Genus B Syntergosternite 7 with strong apicoventral hook-shaped unguis (as in Fig. 8); body yellow, reddish or brownish, rarely black; wing pattern uniformly darkelled, 01' pale brown and hyalille spotted on most of surface ...... 9

16 Sphecomyiella 9 17 LeploPJJigolo cJ'

18 Undescribed Genus A 9 19 PJlrgola 9

20 Tdiopyrgola d' 21 Anapyrgolo d'

22 SlenopYlgola 9 23 ndescribed Genus e cJ'

Fi~s. 69.16-23. Wings: dorsal view of (16) Sphecomyie//a valida (Harris), (M O, fig. 65.2); (17) Leplopyrgola undu/ala Becker; f IX) Undescribed Genus A; (J 9) Pyrgo/a /¡¡gens Wul p; (20) Idiopyrgola sp.; (2l) Anapyrgola sp.; (22) Slenopyrgola sp.; and 1231 Undcscribed Genus C. Figures 17-23 illustrated by V. Hernández-Ortiz. . 960 MANUAL OF CENTRAL AMERICAN DIPTERA ­ VOLUME 2

9. Robust fiies; wing pattern with extensive large brownish dark areas, not mottled; vein R + 2 J usually with stump vein in apical one-third (Fig. 19); paraf,!cial and genal groove broad and usually wrinkled (Fig. 5) Pyrgola Wiedemann

+ Slender f1ies; wing pattern pale brown and hyaJine spotted throughout surface; vein R2 1 without stump veins (Fig. 16); parafacial and genal groove short and smooth (including Slirothrinax Enderlein) Sphecomyiella Hendel

10. Female ovipositor sheath without apicoventral unguis; wing pattell1 with numerous hyaline spots; legs 01' norma I length Boreothril1CLI. Steyskal Female ovipositor sheatb with apicoventral hook-shaped unguis well developed; wing pattem mostJy pale brown (Fig. 23); legs long and slender Undescribed Genus C

Synopsis of the fauna but the present study added three undescribed species from Costa Rica and Mexico. The pyrgotid fauna south 01' the United States includes around 59 known species (Aczél 1956a, 1956b, 1956c; Stey­ Neopyrgola Hendel. Two species ol'this genus are known skal 1967, 1978; Kondratieff & Fitzgerald 1993; Bernard i to occur in Costa Rica, N. majar Hennig and N. picea Hen­ 199 1), but only 1J species have been recorded for Central nig. An additional undescribed species 1'rom that cOllntry \Val America. The present study revealed that approximately 22 al so encounterecl. other undescribed species and three undescribed genera are Pyrgota Wiedemann. Five species are known in lhis ge­ present in the region. nus, including two 1'ound in Central America. Pyrgolo /I/gens Most oY the genera occurring in the Americas were found Wulp was originally described from Guatemala, but there Í' in Central America. Only the North American genus Pyr­ P /ongipe\ golel/a Cmran, and the tropical genera Lopadops Enderlein additional material from southel'l1 Mexico, and (Argentina), Tropidolhrinax Enderlein (Bolivia), and Car­ Hendel is knowll from Guatemala and Brazil. rerapyrgota Aczél (Brazil, Paraguay, and Argentina) were Sphecomyiell.a Hendel. This genus ¡neludes two kno\\n not recorded. Most Central American genera were found in species that occur in the USA, S. valida Harris and S. l/e/son; Mexico (10) and Costa Rica (8). Despite the great number of undescribed species so far uncovered in the family, 1 expect Kondratief1' & Fitzgerald. The specimens examineu in thi, that fUlther collecting wi 11 show that the pyrgotids are even study revealed the presence 01' S. valida in central Mexico. more diverse in the region. Within Sphecomyie//a, the l'ollowing species ol' Stirolhril/ilX Enderlein should also be ineluded: S. cribralus Enuerlein. 1\lIapyrgola Sleyskal. Only one species, A. personlo recorded l'ram Mexico and Nicaragua, and S. knudsel/i Mal" (Lutz & Lima), fram Brazil has been recognized in this ge­ el' l'rom Costa Rica. Additional specimens examined ~llgge'l nus. Several specimens examined in this study represent a the presence 01' three undescribed species: one l'rom Mcxico second unclescribed species from Panama. and two others from Costa Rica.

Boreot/¡rirlflx Sleyskal. Three species have been re­ Stenopyrgota Malloch. The genus currently comprise.\ corded l'mm the USA, of which B. dichaetus Steyskal is also two species, S. mexicana Malloch 1'1'0111 Mexico ano S. UiIS· present in Mexico (Durango). In adclition, two undescribed sitibia Aczél known from Argentina. Studied specimens in­ Mexican species were examined l'rom Querétara, Guerrero, dicate that three other unnamecl species occur in Costa Rica and Oaxaca. and Mexico.

Idiopyrgolll Aczél. This is a monotypic genus represented ndcscribed Gcnus A. This genus comprises at least tilO by 1. selivenlris AczéI l'rom Brazil, but at least three other new species, one represented by specimens l'rom Mexico and species occur in Mexico, Guatemala, Costa Rica, Panama, Guatemala, and the other from Costa Rica. the Bahamas, and Ecuador. lJndescribed Genus B. This genus is represenlcd by Lef'fopyrgolll Hendel. (n the cataIog of the Pyrgotidae three undescribed species all occurring in Costa Rica, bUl south of the LfSA, Steyskal (1967) recognized J 2 species, two of which are al so present in Mexico. and more recently Bernardi (1991) described 21 Brazilian species, most of them on the basis ol' only one specimen. In Undescrihed Genus C. One undescribed specie~ is Ín­ Central America, [wo species were previously recorded fram cluded in this genus, represented by a large series (JI' speci· Costa Rica, L. amplipenl1is Hendel and L. undulola Becker, mens from Mexico, Costa Rica, and Panama. PYRGOTIDAE (PYRGOTID FLlES) 69 961

Literature cited

Aczél, ¡v!. 1956a. Revisión parcial de las Pyrgotidae neotropicales Bu//e/in o[ /he I/linois Agricu//ura/ EX/Jerimen/a/ S/o/ion 116: y antárticas, con sinopsis de Jos géneros y especies (Dlptera, 447-480 ), Parte 1. Revis/a Brasi/eira de En/om%gia 4: Gardner, T. R. & L.B. Parker. 1942. Investigations of the parasites 161-184. 01' Popi//ia jOjJonica and related Scarabaeidae in the Far East \czél, M. J956b. Revisión parcial de las Pyrgotidae neotropicales y froJll 1929 to 1933 inclusive. Reviel1' olApplied En/om%gy antárticas, con sinopsis de los géneros y especies (Diptera, Aca­ (series A) 30: 51-52. Iyplratae), Parte Il. Revis/a Brasi/eira de En/amo/agio 5: 1-70. Kondratieff, B.e. & S.e. Fitzgerald. ILJ93. A new species 01' ;\czél, M. 1956c. Revisión parcial de las Pyrgotídae neotropicales Sphecomyie//a (Diptera' Pyrgotidae) from North AJllerica. y antárticas, con sinopsis de los géneros y especies (Diptera, Proceedings 01 the En/omolugica/ Society 01 Washington 95: Acalyptratae), Parte rll. Revis/a Brasi/eira de En/amo/agio 6: 554-557 1-38 McAlpine, D.K. 1978. Notes on nomenclatllre and type-speci­ Aczél, 1vI. 1958. Pyrgotidae (Diptera, Acalyptrata). Exp/ora/ion du mens 01' AlIstralian Pyrgotidae (Diptera, ). Aus/ra­ Pure Na/iona/ de /'Upemba /\lfission G.F de Witle (1946-/949), /ian En/om%gica/ Magazine 5: 27-33. IJrusse/s 50: 3-53 l\tlcAlpine, D.K..1989. The taxonomic position 01' lhe Ctenostyli­ Ilcrnardi. N. 1990. Caraclerizar,:ao do género Carrerapytgo/a Ac­ dae (=Lochmostyliinae; Diptera: Schizophora). ¡Vlemol'ias do té!, 1956 e sinonimizar,:ao de Anapytgo/a SteyskaJ, 1967 (Dip­ Ins/i/u/o Oswa/do Cruz 84: 365-371. tera. Pyrgotidae). Bio/emas 3: 101-104. Steyskal, G.e. 1967.56. Family Pyrgotidae./n Papavero, N. (edi­ Ilcrnardi, N. 1991. Notas sobre o genero Lep/opyrgo/a Hendel, com tor). A Cata/ague of /he Dip/ero (JI/he Amerieas Sou/h oj'the dcscrir,:ao de vinte e uma novas espécies do Brasíl (Díptera, pYI'­ Uni/ed Sta/es. Departamento de Zoologla, Secretaria da Agri­ golidac). Revis/a Brasileira de Zo%gia 7: 327-350. cultura, Sao Paulo, 8 pp. Cracker, R.L., L.A. Rodríguez-del-Bosque, \N.T. Nailon, JI'., & Sleyskal, G.e. JLJ72. African Pyrgotidae (Diptera) colleCled by X. \Vei. 1996. Flight periods in Texas of three parasites (Dip­ Willi Richter, Erwin Lindner, and others. Stuflgar/er Bei/I'úge lera: Pyrgotidae) of adult Phy//ophaga spp. (Coleoptera: Scara­ zur Na/urkunde 238: 1-11. baeidae), and egg production by Pyrgo/a unda/a. SOll/hwes/ern Steyskal, G.e. 1978. Synopsis 01' the North American Pyrgotidae En/om%gis/ 21: 317-324. (Diptera). Proceedings of /he En/o/1m/ugica/ Socie/y of Wash­ Davis, ,I.J. 19/9. Contributions to a knowledge of the natural en­ ing/on 80: 149-155. cmies of Phy//ophaga, Bui/e/in o[ /he I/linois Natural His/olY Steyskal, G.e. 1987. Pyrgotidae, pp, 813-816. MND, Volume 2. Survey 13: 53-138. Wolcott, G.N. j 923. parasite introduction in Porto Rico. Forbes, S,A. 1908. On the Jife histary, habits, and economic re­ Journa/ o[the Depar/men/ o[Agricu//ure o/Parla Rico. Rcvie\\' lations of the white-grubs and may-beetles (Luchnosterna). ojl1pp/ied En/om%gy (series A) 1¡: 60-61.