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111111.25 111111.4. 111111.6 111111.25 111111.4_ 111111.6
MICROCOPY RESOLUTION TEST CfiART I-IICROCOPY RESOLUTION TEST CHART NATiIlNA. SUR[A.' Qf SIA~u"RDS·19f<';' NA1IClNAL BURll,U or STA"'DARDS·195j·A TB 1575/11/78
A SYSTEMATIC ANALYSIS OF THE PINE PITCH MIDGES, CECIDOMYIA SPP. (DIPTERA: CECIDOMYIIDAE)
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:;;~; UNITED STATES TECHNICAL PREPARED Br '~\.~~; DEP/\RH.IENT or I3ULLETIN SCIENCE /\ND ,~;, ,\CRICULTURf. NUMI3ER 1')7') EDUCATION ADMINISTR/\TION 011 .Janual'Y :2·1, IH7~, four L'SI>A agpneil'~-.-\gTi('ultural Reseal"Ch Service tARS), Coopprali\"E;' SU,te Hl'Sl'arch St'I'\'it'E' {CSR~;I. EXLPnslOl1 Service (ES), and till' :\'ational Agricultund Libral'Y (0:AL).-nwrgE:'d to become a new org-anizatlOn, the ~. I,'nel' and Education Administration (SEA). C,S, De partmpnt of Agriculture,
Thi:-: publleatiun \\'a:-: pn'pared by the Seienee and Edueation Admini~tra ti()n'~ Federal I{esearch staff. whidl \\'a~ formel'ly the AgTicultural Re ~l'ar('h Sen'lee, ACKNOWLEDGMENTS
I am grateful for the aid of all those who collected and loaned material used in this study. Among them are A. T. Drooz, Forest Service, U.S. Department of Agri culture, Research Triangle, N.C.; B. H. Ebel, Forest Service, U.S. Department of Agriculture, Athens. Ga.; R. T. Franklin, University of Georgia, Athens; J. F. Law rence, Museum of Comparative Zoology, Cambridge, Mass.; J. F. McAlpine and J. R. Vockeroth, Biosystematics Research Institute, Agriculture Canada, Ottawa; J. R. McGraw, North Carolina State University, Raleigh; E. P. Markel, Forest Ser vice, U.S. Department of Agriculture, Olustee, Fla.; K. Valley and A. G. Wheeler, Pennsylvania Department of Agriculture, Harrisburg; J. Wilcox, New York State Museum. Albany; and R. C. Wilkinson, University of Florida, Gainesville. I thank also the staff of Cedarville State Forest, Prince Georges County, Md., for allowing and helping me to collect in the forest. My colleagues in the Systematic Entomology Laboratory, Science and Education Administration, P. M. Marsh and G. Gordh kindly identified the hymenopterous parasites reared from Cecidomy;a spp. CONTENTS
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Washington, DoC. Issued November 1978 A SYSTEMATIC ANALYSIS OF THE PINE PITCH MIDGES, CECIDOMYIA SPP. (DIPTERA: CECIDOMYUDAE)
By Raymond J. Gagne', research entomologistl
ABSTRACT
The genus Cecidomyia contains II species, 8 from by use of a cladogram, and the North America spe North America and 3 from Eurasia. Most are only cies are keyed, described, and their pertinent tax broadly host specific and occur on many species of onomic characters illustrated. New species described pines in North America and also spruce and fir in are C. brevispatula. C. !ortunactus, c. reburrata. and Europe. The larvae feed on pitch and can cause C. tortilis. extensive primary damage. The species are analyzed
INTRODUCTION The genus Cecidomyia contains the "pine pitch dleform, and the hind spiracles are at the extreme midges," whose larvae feed on the resin of conifers, end of the body. They are the only cecidomyiid mainly pines. Eleven species are known-eight from larvae found submerged in conifer resin. North America and Cuba, two from Europe, and This is the first complete taxonomic revision of one or possibly two from the Himalayan region. Cecidomyia, although Barnes (1951) reviewed the They are primary feeders and cause extensive literature to about 1950, and Vockeroth (/960) in his damage to pine~. especially in North America. The study of the species on jack pine (Pinus banksiana iarvae form pits or resinous nodules in twigs, Lamb.) reviewed much of the previous taxonomic branches, and cones, where they live completely sub literature. Vockeroth implied a narrow host specif merged in the accumulated resin except for their pro icity to Cecidomyia species by treating his three spe tnlding hind spiracles. cies from jack pine as distinct from those previously The genus belongs to the supertribe Cecidomyiidi. described from Pinus radiata D. Don., P. rigida rhe adult nics may be separated from those of other Mil!., and P. virginiana Mil!. To the contrary, most genera by the following combination of characters: of the Nearctic species that occur on hard pines (Pin The tarsal claws are shorter than the empodia, us, subgenus Diploxylon) have a wide host range. 1 untoothed, and bent beyond midlength; the basi found six species of Cecidomyia in a small grove of meres are stout and unlobed basally, and the male P. virginiana in Montgomery County, Md. Three of tergum and sternum X are of approximately equal them occur from coast to coast on several species of length: the ovipositor is protrusible although not pine. The other three are also on at least one other very long; the antennal Ilagellomcres 1-11 are not pine species in eastern North America. On the other connate: and the palpi have three or four segments. hand, Cecidomyia candidipes Foote is known exclu For a key to genera of North American sively from white pine (Pinus strobus L.), although it Cecidomyiidi, see Gagne (1973).2 The larvae are spin may yet be found on other soft pines (Pinus, sub genus Haploxylon) in western North America as that region becomes better collected. Cecidomyia larvae are usually easy to find and 'SV\tem,1I1<: FntoQ111lo/:lY I .tomaton. C \l l'.S. "allonal Mmeum ,)( \.Ilurai IIlstur~. \VaShllJ/:l111n. D.l' 20560. rear to the adult stage in early spring, although they -' I hl' ,car III 1l,IIIC ,liter the ;Iuthllr\ name tmI!catcs the relerence may not be present on some trees and stands. Full III Ilter.lIure (·\lcd. p I} grown orange larvae, 5-10 mm in length, can be seen in small pitch-filled crevices or large nodules and outside, a split develops along the dorsum of the pitch masses (figs. 2-3) in early to mid-April near pup~1 thorax. and the adult emerges. Larvae of those Washington, D.C. Larvae in twigs brought indoors s~ec.les t.hat pupate apart from the pitch mass have and placed in plastic bags soon pupat" and adults dIstInctIve ?ors~lI abdominal tubercles (fig. 27), emerge shortly after. The larvae either remain in the whose functIon IS unknown. Each larva forms a pitch masses to pupate or leave depending on the white co~oon ~fig. 4). which. when formed on a pine species. Those remaining in the pitch pupate in a needle, IS f~lrly conspicuous. The pupa escapes silken cylinder, which effectively separates the pupae through a CIrcular cap at the anterior end of the from the resin. When the adult is fully formed, the co~oon, and the adult then breaks out of the pupal pupa pushes through a thin cap of dried resin to the sktn.
Genus CECIDOMYIA Meigen
honida :'>\clgcn. 18(}{/: 19. I ypC-'PC<:IC', J/PUlll (lilll Dc (jeer (Co very large, produced apicolaterally to partially qudktt. /Y/(J: 556) Supprcs>cd by Intcrnatlonal COlllllli,Slon envelop telomere in C. rorrilis and C. reburrara; 01' :\olllcndalllr~ (/963: 339). Zoological telomere short or long, variously shaped, setose ('t'Clc/CJllma :'>1clgen. 11m3: 261 1ypc-'peclcs. 7iPIl/ll pini De Geer (rnonotYPlcl throughout, setulose on basal two-thirds, striate on ('<'etc/oil/ria. subgcnus Dlp/v.w LocII. 1850: 20. ryp~-,peclc>. distal third: tergum X with fold at mid length, cerci lil'lIla /,/111 Dc (;ecr (Rllndal1l. 18611: 2S9) triangular; sternum X narrow to wide. simple to CnJ,lcltftrJ/o.\l\ Kldler. ISl)5 <:.\<:1\. I ypc-spcclcs. 7/pula pilli Dc deeply lobed; aedeagus simple, long to short, Geer (ung. dcsignau\lll) SYIHlll)1ll by Kldler (f913. 211). rounded or blunt apically. Female abdomen: Terga RI!I/ltr}(/tp/(lIl.1 Kldlt:r. 191::.' L 1 ype-spccics. Dlplosi.\ resilllcola Osten Sad.en (ong. dcsign.ltlon). Synonym by Vockcroth I-VII as for male except VII as long as preceding ter (11)0(1 (,S) gum; sterna I-VII as for male; tergum VIII rectan Srelt'l'/tor/lplo.l/s "'16hn, 19551>: \4 I. Type-species, SrelechudiplosLI gular, with basal pair of trichoid sensilla and scat /IIagl/a "'1l)hn (orig. designation). Synonym by Vad.croth tered setae on distal half; sternum VIIl not apparent; /960: (6). ovipositor short, protrusible, with scattered short Adulr.·Head as in figures 6-7. Eyes broadly setae laterally and ventrally; tergum X lightly sclero cont1uent at vertex, facets closely juxtaposed, hexag tized. with scattered short setae; cerci with two short. onal. Postvertical peak absent. Antenna with 12 flag apical sensoria. covered elsewhere with dense, short ellOl11eres, I and II not connate; male flagellomeres setae. binodal, tricircumfilar; female flagellomeres cylindri Pupa.-Antennal horns rounded, crested, or cal. their two circumfila connected by two longi peaked. Pronotum variable, with two pairs of setae. tudinal strands. Palpus four-segmented, except three Frons with two pairs of setae. Face with group of segmented in C. piniinopis. Wing RS curved distally. three papillae at base of each palpus. Prothoracic jotning C posteriad of wing apex; C broken at junc spiracles variously shaped, variable within species. ture with RS' Rs weak; M3+4 fold present; eu Abdomen covered with minute spiCUles. rorked. Legs usually covered with brown scales, but lArva (figs. 27-28).-Head capsule with very C. candidipes white on distal three hind tarsomeres; long apodemes. Sternal spatula of third instar vari tarsal claws simple. curved beyond midlength; ously shaped or absent. Papillae as follows: Laterals empodia slightly longer than claws. Male abdomen: on thoracic segments in two groups of three each per Terga I-VI large. rectangular, with approximately side. two of each group with short seta; four fore two rows of caudal setae, a group of lateral setae. ventrals and two hind ventrals, all unhaired; two and scattered narrO\v scales elsewhere; tergum VlI as haired pleurals per side, in some species abdominal for V I except shorter, caudal setal rows interrupted pleurals situated on tubercles; six dorsals (except mesally; tergum VIII very short, barely sclerotized. abdominal segment VIll with two (homologous with naked except for pair of trichoid sensilla; sterna II middle pair on previous segments», each with seta VII quadrate, with two to three rows of caudal setae, except middle pair sometimes without, lateral pairs scattered setae elsewhere. denser laterally, and scat of abdominal dorsals in some species each situated at tered narrow scales; sternum VIII less sclerotized end of common, long, cephalocaudally flattened than preceding, with only basal pair of trichoid sen tubercle, middle pair of papillae sometimes situated sil1a and apical setae; basi mere unmodified. except on short tubercles: terminals primitively four per
2 side, one to two lost or not apparent in some species, (2) Caudal extensions of terminal spiracles: at least one with cylindrical peglike seta, one with Short, not langer than wide; long, much longer than long, tapered seta. Hind spiracle greatly enlarged, wide. projecting caudad of anal ("terminal") segment, in (3) Size and shape of larval spatula: Large, clove first instar terminating in single point, in second shaped; reduced in size or absent, when present ceph instar in two, and in third instar, in four. alic margin not deeply cleft. Remarks.-Cecidomyia is distinctive and readily (4) Length of cephalic apodemes of larva: keyed (Gagne, 1973), but I cannot bring its affinities Shorter than head; longer than head. with otha genera into focus except to place the (5) Number of papillae on terminal segment of genus within the large mass of phytophagous larva: Three; two. Cecidomyiidi. Since the adult characters are basic to (6) Larval spatula: Present; absent. the supertribe, although diagnostic, they do not lend (7) Setation of dorsal papillae of larva: Middle themsehcs to clarifying relationships. The adult pair on segments I-VII and two papillae on segment character states that 1 consider plesiomorphic are the VIII with setae; without. absence of a postvertical peak, the large eyes, the (8) Shape of terminal spiracle: Bilaterally sym separate tlagellomcres I and II, the large abdominal metrical; longer mesally than laterally. terga and sterna with their full complement of setae, (9) Size of male terminalia: Normal; extremely the la..ge male tergum X with the horizontal fold large. cephalad of the cerci, the short ovipositor with large, (10) Dorsum of larval abdomen: Unmodified; free, unjuxtaposed cerci, and the presence of a weak with tubercles. Rs wing vein. I believe that the apomorphic char (II) Pupation site: In pitch mass on clear cocoon; acter of the long empodia relative to the length of apart from pitch mass in opaque cocoon. claws is insignificant, because it is a character most (12) Size and shape of larval spatula: Wide ceph conifer cecidomyiids share. regardless of their affini alic margin. tapering caudally to long shaft; trian ties. The larvae of Cecidomyia are basically typical gular, not longer than wide. Cecidomyiidi larvae except insofar as they are mod (13) Number of adult palpal segments: Four; ified for life in conifer resin. three. Cecidomyia species are easily distinguishable. The only group of Cecidomyia species common to Moreover, enough characters are definable into both North America and Eurasia has larvae with piesiolllorphic and apomorphic states that a cladistic dorsal tubercles. It appears to me that the genus had analysis uf the species is possible. The cladogram its genesis in North America and that a section was (fig. I) is based on the following 13 characters. Each able, for whatever reason, to colonize Eurasia, where character is followed first by what I consider to be speciation occurred before subsequent isolation. This the plesiomorphic state and then by the cor pattern is reflected in another group of cecidomyiids, responding apomorphic state. the large tribe Lasiopterini. The subtribe Alycaulina (I) Number of papillae on terminal segment of occurs entirely in the New World, and its sister larva: Four, one without seta, two with long, pointed group, the Lasiopterina, is found throughout most of setae: fewer than four, only one with long, pointed the world (Gagne, 1969). seta.
THE NEARCTIC SPECIES The Nea.rctic species are treated here alphabet (Chapm.) Vasey, P. conlorta Dougl., P. echinala ically. Locality data for previously described species Mill., P. elliottii Engelm., P. glabra WalL, p, palu are broadly summarized by host and State or Prov slris Mil\', P. ponderosa Doug\. ex P. and C. Law mce. The complete collection data are on file in the son, p, radiala D. Don., P. resinosa Ait., P. rigida Diptera Unit, Systematic Entomology Laboratory, Mil\., P. serolina Michx., P. slrobus L, P. taeda L, L' .S. Department of Agriculture, but eventually will and P. virginiana Mil\, They include all the pines be deposited III the Smithsonian archives. found in North America east of the Mississippi River Cecidomyia species are recorded from the fol except P. pungens Lamb., but only four from west of lowing native American pine species: Pinus bank the Mississippi. sIGna Lamb., p, caribaea Morelet, P. c/ausa
3 KEYS TO NEARCTIC SPECIES
The adult stage of C'. fortlillClctlis Gagne is ments. and C. c{Jnclidipes, on which distalmost hind unknown. Females of Cecidolllyia are too similar for tarsOl11eres arc covered with White instead of brown etfcl:ti\c dillcrimination except those of C. piniillopis, scales. whil:h have three instead or the usual four palpal seg-
Males
L IcrmlOalta Icry larg\!. much wider than abdominal segments VI and Vii...... 2 lerm11lalla ~maller (normal). narnmer than abdominal segments VI and VII ...... J , Ha~lInere (fig, S-9) rounded distally, with dense apical bristles cm'ering telomcre in caudal VICW; telomere stnught .....•...... ••....••...... •.... C. relillrr(](a Gagne Ba,lIncrc (lig,. 10·11) bilaterally narrowed di~tally. without dense bristles; telomere twisted . . . • . . • . •...... •...•..•...•..•.....•.•....•..•....•...... C. tOrlilis Gagne Palpll, three-~eg111ented . ..•. .. •.•...... • ' ...... C. I'illiillopis Osten Sacken Palpus four-'egmentcd...... •...... •...... 4 4. Sternu111 X about ih W tde as acdcagus. Simple, rounded apically (lig. 16): hind tarsomercs I_II bnmn. III-V lIhue, .••.•.....•.....•....•..•...... •.....•...... C. c{ll/(lic!ipes Foote Sternum X delllHtcl) Ivider than aedeag1l5, at least weakly hilobcd apically (figs. 12. 15. 17): him! tar\u, unllortnly brown .•...... •...... •...... •.... 5 5 ·\edeagm Illdc~t beyond middle; sternum X deeply btlobcd (fig. 12) .• C. resil/ico/a (Osten Sacken) AClkagu\ Illdest at base; sternum X weakly hllobcd (l1gs. 15.17)...... 6 6. Sternul11 X lunger than aedeagus. paralled-slded (fig. 15) ...... C. resil/ic%ities Williams Sternum :\ Ihorter than aedeagus. IIlder basally than .\pically (l1g. 17) ...... C. brevi.l'{JilIli/a Gagne
Last-Instar Larvae
Abdomen I~lth dorsal tubercles (fig. 27)...... • ...... 2 Ahdomen Illthout dorsal tubercle, (lig. 28)...... 4 .; ~liddlc pair 01 dorsal abdominal setae not set on tubercles (fig. J7): spatula (fig. J6) almost en tirely black; Irom 1'. \'IronIl5 ••••• ••.••..•••••••••••••.•.••.•.••.••.••. C. c{ll/dit/jlll's I"oote ~llddle pUlr ot dorsal abdominal "etae set on tubercles (figs. 39.42); spatula (figs.40,41)brown; Irom hard PITlCS. ••••. ••••.••••••••••..••.•••••.•.••.•.••••••••• .•.••••••..•••••••• J Spatula long (Iig. 40); lateral pairs of dorsal abdominal tubercles weakly bifid (fig. J9) •.•...... •...... •.... C. {Jiniil/opis Osten Sacken Spatula short. tl'langular (Iig. -ll): lateral pairs of dorsal abdominal tubercles deeply bifid (fig. 421 ...... •...•...... •...... •...... •...... C. hrevi.l'{J{(IlI/{I Gagne 4 Spatula prc,cJ1l (fig~. 32. J5): middle p,ur of dorsal abdominal papillae with setae: abdominal ,cgment \'111 (beal'lng hind spiracles) with three pairs of dorsal-pleural setae (figs. JJ. J4)
• ••••••••••••••••• a , ...... ~ ••• " •••••••••••••••••••••••••••••••••••••••••••••••••• 5 SpalUla ab,e[\(; middle p.llr 01 dorsal abdominal papillae without setae (fig. 29): abdominal segment VIII with tWll P:lIfS or dorsal-pleural setae (rigs. JO. 31) ...... 6 5. Spatula large. deeply bifid cephalad (fig. 32); caudal prongs of hind spiracles short (fig. JJ) ..•..••...••...... •...... ••...... •...... , ...... c.jortunactu.l' Gagne Spatula shorter. weakl} bifid cephalad (fig. 35): caudal prongs or spiracles long (fig. J4) · .• " •.••...•.•.•...... , .•...... •...... •...... C. resinic%ides Williams /1. Hind splraeb longer mesally than later:tlly (fig, 30) .•...••...... C. resinico/a (Osten Sacken) HlIld ~plraclc, of c4uallength mesally and laterally (fig, JI)C'. relillrratlJ Gagncand C. tortilis Gagne
DESCRIPTIONS OF NEARCTIC SPECIES
Cl!cidomyia hrevispatu/a Gagne, new species Last-instar larva (figs. 41-42).-Spatula brown, triangular. tiny. Pleural and dorsal abdominal Adult. Palpus four-segmented. Legs covered papillae with long setae. situated on tubercles, two \\lth bt\)\\11 scale,. ~lale terr11lnalin (fig. 17): Sternum lateral pairs of dorsal papillae each on common, \ eol1l:iI\t' ilpll:alh: acdcagll~ short. wide. slightly forked. anteroposteriorly flattened lobe. Two ter notl:hcd aptl:alh minal papillae (as in fig. 39): One with long, tapered Pupa. l!nknown. seta; one with long, peglike seta.
4 Holotype, larva, ex P. elliottii, Lykes Fire Tower, Maryland. l"ew York, Ontario. Pennsylvania, and Glades Co., Fla., 31-V-1973. J. R. McGraw, USNM Quebec. Felt (19/8) stated that the larvae trans Type No. 66650. Paratypes: 10 larvae, same data as formed to pupae within the pitch masses, but his holotype; malt:, 3 larvae, P. elliottii, Mile Post 65, notebook (unpub.), where complete collection data Interstate 75. J. R. McGraw; 2 larvae. P. echinata, were generally kept. notes that "larvae of [albita.rsis] Murphy, N.C.. 7-15-1970. J. L. Rauschenberger; 2 occur in the pitch masses and 3 cocoons, 2 on nee larvae from pitch mass on branch. P. cla usa , U. of dles and I on bark. were separated." If larvae of C. Fla. campus, Alachua Co., 31-XII-1970, J. R. co.ndidipes do leave the pitch masses to pupate, their McGraw: J larvae. P. virginiana, Wheaton Park, behavior would fit that of the other two Nearctic Montgomery Co., Md., Oct. 15. 1965, R. J. Gagne. species with larval tubercles, C. piniinopis and C. Several paratypes will be deposited in the Florida hrel·ispo.lLl/o.. ( cannot conclude that the presence of State Collection of Arthropods, Gainesville. the tubercles and the habit of leaving the pitch mass Etymology.-Brevispawla. a noun in apposition. to pupate are correlated, because one European spe refers to the short. larval spatula. cies, C. magna (Mohn), that has abdominal tubercles Distribution and biological notes.-c. brel'i reportedly pupates in the pitch mass. spl/w/a wa~ found on several species of pine in the Southeastern United States: P. c/ausa (Fla.). P. echi Cecidomyia !ortunactus Gagn~, new species flaw (!':.c.), P. ellioLlii (Fla.). and P. virgifliana (Md.). The specimens from P. elliottii were collected Adult and pupa.-Unknown. near needle bases of new foliage on expanding tenni Last-instar larva (figs. 32-33).-Spatula large. nals in the area of feeding injury of Rhyaciollia sub strongly sclerotized, deeply bifid cephalad. Pleural £ropica Miller (Lepidoptera: Olethreutidae): those and dorsal abdominal papillae with short setae, no from P. I'irgifliana were found in a small pit near the papillae on tubercles. Four terminal papillae: Two base of a needle fascicle. each with long. tapered seta; one with long, peglike seta; and one with short. conical seta. Terminal spi Cecidomyia candidipes Foote racles bilaterally symmetrical with short, caudal prongs. Rel1llo,III'{,,\l' albue/rlll h.!l,. /'i18: 3SJ (set'ondar~ homonym: Holotype. larva, in resin drop on twig, P. I'irgin prcn~t'uplt:d 111 ('''<"It/Oll/llt! h} :--.tclgell (lS03)). iano.. Wheaton Park. Montgomery Co.. Md., Apr. I, (t'lIdOIllIItI "m,lId,pt', h>lHt:. l'in5. 2S7 {ncII name tor alhjlar.lis 1976, R. J. Gagne, USNM Type No. 66649. Para l'cIt I types: 2 larvae, same data as holotype, except IX-22 Adult.·Palpus four-segmented. Legs with white 1976; larva. ex P. ellio{(ii. Lykes Fire Tower, Glades ~cales covering hind tarsomeres Ill-V. brown scales Co.. Fla .. 31 July 1973, J. R. McGraw. Additional elsewhere. Male terminalia as in figure 16. record: Larva. ex P. elliottii. ex pitch canker diseased Pupa. Antennal horn pointed anteriorly; pro terminal. Flagler Co., Fla., 30 July 1974, R. C. notum as in figures :22-23. Wilkinson (this specimen lost during slide mount Last-instar larva (jigs. 27. 36-37).·--Spatula ing). mostly black. occasionally brown anteriorly. Pleural Etymology.-Fortur.actus is a telescoped com and dorsal abdominal papillae with long setae, no pound name from fortuna (lucky) and nactus (a papillae on tubercles except two lateral pairs of dor find). We are fortunate to know this species, if only sals. each situated on common, forked. ante the larval stage. because it is important in estab roposteriorly tlattened lobe. Three terminal papillae: lishing the ground plan of th~ genus. Fortunactus is One with long. tapered seta; one with long, peglike a noun in apposition. ~eta; one sometimes not apparent. without seta. Ter Distribution and biological notes.-c. Iortu minal spiracles bilaterally symmetrical. with long, nactus is known only from Maryland on P. virgin caudal prongs. io.na and from Florida on P. elliottii. One of the Lectotype, male. here designated. from larva in Maryland paratypes was found in a droplet of pitch pilch mass on P. strobus. Round Lake. N. Y ...June that also contained two dead, last-instar larvae of C. (91X. a2917. in Felt Collection. New York State piniinopis. The five known specimens of this species :-Vl.useum. Albany. Paralectotypes: 4 males, 4 females, were all found singly. 4 larvae. same data as lectotype. Distribution and biological notes.-c. candid Cecidom.via piniinopis Osten Sacken ipes IS known only from white pine (P. strobus). I ("<.'('11/(11111111 pill/inC/flis (as {Jilli II/Opis) Ostcn Sackcn. 1815:!.' 196. ha\ e seen specimens from Connecticut. Illinois, (/!('ltlo/lll ill !>lIlllw(//l(Je Vockeroth. No().· 73. ;-'C\I synonym.
5 Adult.Palpus four-segmented. Legs covered reared adults in September so that there are at least with brown scales. Male terminalia (figs. 18-19): two generations in Maryland. Generations probably Sternum X concave apically; aedeagus approxi overlap. I found one larva of each of the three instars mately as long as sternum X, parallel-sided, blunt in September in Maryland in a small twig wound. apically. The species survives the winter in the larval stage, Pupa. .Antennal horn pointed anteriorly; pro and cocoons arc much more numerous and larvae notum as in figure 24. more readily found in early spring than at other Last-instar larva (figs. 38-40).~Spatula brown, times. Reeks (/960) noted that infested buds or widest cephalad, tapering abruptly behind, cephalic shoots may be killed by this species, but he did not edge straight. Pleural and dorsal abdominal papillae find it common enough to consider economically with long setae, situated on tubercles, two lateral important. pairs of dorsal papillae each on common, oblong, 1 reared a series of the small, black Plalygasler anteroposteriorl), f1attened lobe. Two terminal papil dip/osidis (Ashmead) (Platygasteridae) specimens lae: One with long. tapered seta; one with long, peg from each of several cocoons of C. piniinopis and, like seta. Terminal spiracles bilaterally symmetrical, on one occasion, a single large, green-metallic Hab with long. caudal prongs. racyllis sp. (Pteromalidae). Types of names in this taxon.- C. piniillo{!is O. S.: Syntypes, 5 cocoons, deposited Cecidomyia reburrata Gagne, new species in Museum of Comparative Zoology, Cambridge, Adult.---Palpus four-segmented. Legs covered Mass., and an undetermined number of larvae (lost, with brown scales. Male terminalia (figs. 8-9): if originally kept) taken on P. virginiana (as P. Greatly enlarged; basimere with dense apical patch inops) IV-V, 1861, \Vashington, D.C., Osten Sacken. of long setae hiding telomere i.n caudal view; C. banksiaflau Vockeroth: Holotype, male. from P. telomere situated mesoapically on basimere, bilater bal7ksial/(J, Stead, Manitoba, VI-1957. W. A. Reeks, ally compressed. rype No. 6833, in Canadian National Collection, Pupa. --Antennal horn rounded anteriorly; pro Ottawa. notum as in figure 31. Remarks.--All that exist of Osten Sacken's Last-instar !arva.-Spatula absent. Pleural type-~eries of C. piniinopis are five cocoons that arc papillae and lateral pairs of dorsal papillae of not diagnostic and arc the same as formed by C. abdomen with short setae, no papillae on tubercles; bruI·isplJlU/a. which is also found on P. virginiana. middle pair of dorsal papillae without setae or, on Ostcn Sacken noted that the dorsal tubercles of the abdominal segment VIIl, not apparent. Three ter larvae wac oblong. That character will separate C. minal papillae: One with long, tapered seta; one with pin.inopis from C. brel·ispawla. which has forked long, peglike seta; and one with short, conical seta. tubercle~. Terminal spiracles (as in fig. 31) bilaterally sym Distribution and biological notes.-C pini metrical, with long, caudal prongs. inopis and C. resinico/a are the two most widely dis Holotype, male, emerged May 19, 1975, from tributed species of Cecidomyia. Both occur on many larvae cullected April 18 in wound on P. taeda, Ced species of hard pine (Dip/oxylon) in eastern North arville Sta',e Forest, Charles Co., Md., R. J. Gagne, America and in the West. I have seen specimens of USNM Type No. 66647. Paratypes: 2 males, 2 pupal C pilliillopis from P. ballksialla (Mich.. Ontario, exuvia, 6 pupae, 6 larvae, same pertinent data as Wis.), P. conlorta (Brit. Col., Yukon TeL), P. holotype; male, Florida Caverns State Park, Jackson cl/iotlii (Fla.). P. ponderosa (Ariz., Brit. Col., Calif., Co., Fla., 26-V-1973, malaise trap, W. W. Wirth; Idaho, Oreg .. Wash.), P. rigida (Maine, Md., N.Y., larva, 3 females, 3 pupal exuvia, P. taeda, 10-11 mi. N.C.), P. s(rohlls (Md.), P. I'irginiana (D.C., Md., sw. Alexandria, La., 26-II1-1960, J. G. Chillcott; 2 Va.), and P. sp. (Ark.). males, female, P. taeda. Grangeville, La., IV-2-1968, Larvae of C. pilliinopis usually occur singly in D. K. Pollet; male, female, P. taeda. Livingston, La., small pitch droplets and sometimes in inconspicuous IV-16-l968, D. K. Pollet; I larva, 4 males, 4 pupal pockets at the base of needles. Full-grown larvae exuvia, P. taeda. Turkey Creek, La., II1-28-1960, J. leave the pitch mass, weave a white cocoon (fig. 4) G. Chillcott; 20 larvae, 4 males, 4 females, P. car some distance away on the twig, candle, or needle, ibaea, Punta del Rio, Cuba, 1I-23-1928, S. C. Bru and pupate immediately. Adults emerge within a few ner. Several paratypes will be dep0f'.ited in the days. Miller (/958) suggested that there were up to Florida State Collection of Arthropods, Gainesville. four broods yearly on P. rigtda in Ohio. I have Etymology.-Reburrata is an adjective meaning
6 bristly, in reference to the strong, dense setae on the species of this genus. Both occur on almost all the male terminalia. hard pines (Diploxylon) in eastern North America Distribution and biological notes.-C reburrata and on several species in the West. I have seen speci is known from southern Maryland to Florida on P. mens of C resinicola from P. banksiana (Mass., laee/a and from Cuba on P. caribaea. It is responsible Mich., N.W.T., Ontario), P. eontorla (Brit. Col., tor large, pitch-filled nodes on twigs and branches in Oreg.). P. echinQta (La.), P. elliottii (Fla.), P. glabra which the larvae. up to I cm in length. live gregarious (Fla.). P. palustris (Ala.), P. ponderosa (Calif., ly. Pupation occurs in silken tubes within the pitch Colo., Idaho, N. Mex.), P. radiata (Calif.), P. rigida masses. (Md., Mass .. N.J., N.Y., N.C., Ohio, Ontario, Pa.), P. serotina (N.C.), P. taeda (Ga., Md., Miss., Va.), and P. virginiana (Md., Va.). Cecidomyia resinicola (Osten Sacken) C resinicola may occur singly in small droplets of pitch (fig. 5) or gregariously in larger pitch masses 7 D1f'it'\l1 n'\11licolu Osten Sad.en. 18 }; 345. (fig. 3) as well as pitch-filled nodules. The species Rt!illl 7 A dull. Pal pus four-segmented. Legs covered nodule. He reported that Syntasis diplosidis Eckel with brown scales. Male terminalia (fig. 15): Sternum (Pteromalidae) was reared from this fly. X long, wider than aedeaglls, parallel-sided, concave apically; aedeagus shorter than sternum X, parallel Cecidom.via tortilis Gagne, new species sidcd, broadly rounded apically. Adult. -- Palpus four-segmented. Legs covered fupa.- Not seen. with brown scales. Male terminalia (figs. 10-11): Last-instar larva (figs. 34-35).-Spatula short, Greatly enlarged; basimere with apicolateral margin narnm. weakly bifid cephalad. Pleural and dorsal expanded to partiaily cover telomere in caudal view; abdominal papillae with short setae, no papillae on telomere situated mesoapically on basimere, weakly tubercles. Two terminal papillae: One with long, sinuate and twisted. tapered seta; one with long, peglike seta. Terminal Pupa. Antennal horn rounded anteriorly, pro spiracles bilaterally symmetrical, with long. caudai noLum as in figure 26. prongs. I.L1st-instar larva (figs. 28, 3l).-Spatula absent. Types of names in this taxon. Pleural papillae ,ltld lateral pairs of dorsal papillae of e. rf!siwwloicles Williams: Lel;tlltype. here desig abdomen with short setae, none on tubercles; middle nated. male. "Stan. Uni, .. Cal.. coil. by R. W. pair of dorsa! papillae without setae or, on abdom Duane, aI930," in Felt Collectiun, \iew York State inal segment VIlt, not apparent. Three terminal ivfuseum. Albany. Paralel:totype: Female. same data papillae: One with long, tapered seta; one with long, a~ male. l'ht.:se two specimens are evidently the only peglike seta; and one with short, conical seta. Ter two extant specimens or the original type-series (Ar minal spiracles (as in fig. 31) bilaterally symmetrical, naUd, 1c)7U). which also induded at least eggs, larvae, with long, caudal prongs. and pupae it not other adults. Holotype, male, emerged V-2-1976, from larvae C. accola Vocke roth: Holotype. male. Espulloia, collected IV-I-1976 in wound on P. virginiana, Ontario, June 2, 1951:{, Type No. 6834. in Canadian Wheaton Park, Montgomery Co., Md., R. J. Gagne', !' THE PALEARCTIC SPECIES Cecidomyia pini (De Geer) cocoons, not seen; not in Swedish Museum of Nat ural History (P. I. Persson, pel's. commun.). 1Il'u{a {'1I1I Dc (iet:r, 1~"().. 417. C. pinimaritimae Dufour: Syntypes, 2 Jema.les, 2 CnldO/l/l/tl 1'/I/llllrlrJIIII1I1' (a, pilli lIIariril11(1e) DUrllur. 1838: 294. !->\n,lIl\ mill Burries (/89/) and authors. cocoons, Jrom resin of Pinus pinaster Ail. (P. mar Ct'cldomrirl pi/O.ICl Brcl11l. 184': J I. 61. Sy,nonym oy Borrics itima Mill.), sw. France. I do not know if types exist (/SIII) and duthor,. or, if so, where, but females and cocoons are not diagnostic. For illustrations or all stages, see Tubeuf (/930); of C. pi/osa Bremi: Type in poor condition, covered male terminalia, Molm (/955b); of larva, Mohn with fungal hyphae, sex undetermined, "Bre./ Lin (I 955a). dau" [nr. Zurich], in Entomologisches Institut, Types of names in this taxon.- Eidgenossischen Technische Hochschule, Zurich. C. pini (De Geer): Syntypes, all life stages and Information provided by W. Sauter of that institute. 8 Remarks. -- According to the literature on C. Museum of Natural History. They are from Pinus pini as summariLed in Barnes (1951), this species roxburghii Sarg. in Pakistan. The larvae have dorsal occurs on pines, fir, and spruce. Possibly more than tubercles as do the other non-American species, and one species is represented under this name. Syn the male terminalia are distinct from the two Euro onymy of the three available names has been based pean species as illustrated in Mohn (/955b). I have on similarities of the biology and gross anatomical not described this species here because the male geni characters, not comparison of male terminalia of talia are slightly distorted on the slide, and I do not specimens from the various hosts. Unlike C. magna know whetht::r the larvae leave the resin mass to (Mohn), larvae of C. pini leave the pitch mass to spin pupate. cocoons in which they pupate. ?Cecidomyia species Cecidomyia magna (Ml.ihn) C"dc/oll/ria ~p. Stehhing, 19()5: 430. This Indian species may be a Cecidomyia, but one Slelechodiplo.li.l magna :vltihn. 195511: 141. cannot be sure from the description and illustrations Description in Mohn (l955b), including illustra in Stebbing (1905). The larva has pleural and lat tions of male terminalia. larval spatula, and larval erodorsal tubercles the length of the body, but the terminal i>egments. conspicuous hind spiracles one would expect to find H(,lotype. male, "an Fichte," HebertshBusen (nr. on a Cecidomyia are not iIl.ustrated. Instead, Munich). Germany, IV-22-1954, Nr. 552-T, in E. Stebbing notes that the larva is pointed at both ends. Mohn Collection. Also, known Cecidomyia species with tubercles do Remarks. - This species is known only from not have them on the thorax as Stebbing's illustra spruce. It pupates in the resin mass unlike all the tion shows. The larvae of this species feed on resin other species whose larvae have dorsal tubercles. and are responsible for a gall on Pinus roxburghii Sarg. (P. fongifolia Roxb. not Salisb.) in northern Cecidomyia, new species India. They cause a foreshortening of the growing tip that resembles a pine cone. When fully grown, the This undescribed species is known from a male, a larvae leave the gall to pupate nearby in white female, and three larvae in the U.S. National cocoons. LITERATURE CITED Arnaud. P. H .• .lr. Eckel. l.. S. I'.)70. LIsts of the 5cicnllfio;: puhlications and insect taxa 1903. rhe resin gnat diplosis and three of its parasites. Ent. descrIhed hy franCIS Xavier \Villiam~ (1882-1967). Calif. News 14: 279-284. pI. xiI. Acad. SCI. Occas. Papt:rs "0. SO, j3 pp. San francisco. Felt, E. P. Barnc~. H. f. 1915. ;';ew gall midges. Jour. Econ. Ent. 8: 405-409.. 1951. Gall midges of economic importance. Vol. V. Gall midge, of trees. 270 I'p. Croshy, Lockwood & Son Ltd .. 1918. :-';ew gall midges (Dipt.). Jour. Econ. Ent. II: 380-384. London. Foote. R. H. Borne,. H. 1965. Family Cecidomyiidae. III Stone et al.. A Catalog of the 1891. C\ogle nye lagttagebcr over danskc Naaletrae-Insektcr. Diptera or America "orth of Mexico. U.S. Dept. Agr. Agr. I Cecidomria pilli. De Geer. l'idsskr. for Skogbruk 1891: Handb. 276. 1696 pp. 2.19-249. Gagne, R. J. Brcml.,1 J 1969. A trihal and generic revision of the Nearctic Lasioptcridi 184-. Bcitraegc lU ciner Monographic der Gallmiickcn, Ce (Diptera: Cecidomyiidae). Ent. Soc. Amer. Ann. 62: 1348 ciclo/llviU Meigel1. 71 ppo, 2 pis. ;O-;cucnherg. 1364. (\lljlllht:tt. D. H. '1973. A generic synopsis of the Nearctic genera of 1910. Ihe type-~pecies o( the North AmerIcan genera of Dip tel'lL LS. \atl. 1-.lu,. Proe. 37; 499-647. Cecidomyiidi (Diptera: Cecidomyiidae: Cccidomyiinae). Ent. Soc. Amer. Ann. 66: 857-889. De Geer. C. International Commission on Zoological 1'omenclature. 1"~6 ~lcnHlIfe:i pour scrvlr a I'hi~toire des inscctes. V. 6. 523 1963. Opinion 678, the suppression under the plenary powers pp • .10 pIs. Stockholm. of the pamphlet puhlished by Meigen, 1800. Internatl. Dufour. L Comn. Zool. 1'omencL 20: 339-342. 183~. \ote.' pour servir a I'histoire des cecidomycs, et de~crip Keen, F. 1'. lion d'unc nnuvdie espccc de ce genre de dipleres. Ent. 1938. Insect enemies of western forests. U.S. Dept. Agr. Misc. Soc. hance Ann. 7: 293-296. Pub. 273, 210 pp. (Revised, 1952.) 9 Kldkr. J,.I. Osten Sacken. C. R. 1895, :\lltl\cllc, ob,cnations sur Ie groupe de, Difllusis ct 1862. V. On the :'I:orth Amt!rican Cecidomyidac. III L.ocw. H .• descrlptlon de cinq genres nouveaux (Dipt.). Ent. Soc. Monographs or the Diptt!nl of ;>.'orth America. pt. I. pp. hanee Ann. 64 (Bul.): CXCil-cxci\'. 173-205.2 pIs. Smithsn. Misc. Collect. 6 (I pub. 141): 1 221. 2 pis. 1912. Ncue Uallmucken-Gallungen. 2 pp. Bitsch, France. 1871. Biological notes on Diptcra. (Article 3) Amer. EnL Soc. 191 J Dlptera. Fam. Cecidomyidae. Genera Insectorum. Fasc. Trans. 3: 345-347. 152. 346 pp,. 15 pb. Bruxellt:s. l.l\t!\\. 1-/, Reeks. W. A. IH50 Diptcrologi,che beitrage. Viertel' Theil. K. hied rich 1960. Ob8t:rvations on the life history. distribution. and abun Wilhellll\-Gymna,iulIIs iU Po;en. OITent!. PrM d. Schuler dance or two species of Cecidol/l,l'ia (Diptera. IH50: 1-40. I pI. Berlin. Cecidomyiidac) on jack pine in Manitoba and Sas I\klgcn. J, \\". katchewan. Canad. Ent. 92: 154-160. ISOO. Nou\elle da,sificalion des mouche; ;\ deux aile, (Dip Rondani. C. tera 1..J d'apri:s un plan tout nOUleau. -10 nr,. Paris. li!60. Stirpis cecidomyarum. Genera revisa. Nota undecima. pro dipterologia italica. Soc. Ital. di Sci. Nat. Atti (1859- IStU Ver,ueh ciner nCLJcn Gattungseintheilung des euro IH60) 2' 286-294. I pI. piibehen 1\\ eillligciigcn Insckh:n, Mag, Inscktcnkunde 2: Stcbbing. E. P. 259-2li L 1905. On the cecidomyid (Cecidol/l)'ia e!) sp.) forming the "hiler. \\'. F galls or pseudo-cones on PillliS /ollgifolia. Indian Forester 195H, EcologIcal notes on the pine midges Relil/oc/iplosis (Allahabad) 31: 429-434. pI. xxxviii. re,I//:;"ola (Osten Sad,en) and R. il/opis (Osten Sacken) in TUbeuf. C. von. ,l1uthcrn OhIO (Diptera. Itonididae), Ohio Jour. Sci. 5S: 1930. Diplosis pilli (De Geer). (Cecido myiden-Studien). 231-235. ;>'Illhn. F. Ztschr. r. Pllanzcnkrank. 40: 375-390. IlJ5Sa. Beitrage lur Sy,lematik der Lanen der Itonididae (= Vockeroth, J. R. Cecldomyildae. Diptera). I. reil: PorricolJf'ylinac und 1960. Taxonomy of the genus Cecidolll)'ia (Diptera: ltonld'lnae .\tlttclcuropas. Zoologica 105 (I and 2): 1-247. Cecidomyiidae) with special reference to the species occur 30 pIs, ring on Pinlls bllnksianll Lamb. Canad. En!. 92: 65-79. Williams, F. X. 1955b. Neue frdlebcnde GallmOcken-Gattungen. Deut. Ent. 1909. The Monterey pine resin midge - Cecidolllyia resin i Llsehr. 2: 127-151. colaides n. sp. Enl. News 20: 1-8. I pI. 10 NEARCTIC PAltARCTlC ( FOR rUNACTUS RESINICOLA REBURRATA TORTIUS RESINICOLOIOES UNDESCR SP \~ / hgurc I CladQgrarn for Cecidomyia species. II 1'\·{l42I h!!'H~' :-~ (nl!, -" (nld'lIIlIllI 1\lIh pupal C\.U\I.l .\Od C,'CO\\() ::, c. IOffl/(' Ilodule Oil PIIlW dlll/slI. Panama ('Ill, 1-1<1.; J. C "'.1//11 ""Ii I'lldl n1.\",,11 I'U/II\ rtll/d.,. llurhl1!'(llll ("'Ul1h. \..1 _4. C. /1/111"'''1'/1 ~IIC"II!I 011 P J/f()/>U\. Charlc\ Counly. ~Id .• 5, C. rt'.III/1 , ••;',; 1'11.;1\ 11),,,, .>11 l' I/r\llfllllllf/. Belt-\t!ie. '.Id. 12 6 " . ,,~ '" to " , , ' ' , / , ' , , , ..' , , . o. o , , . " , " 8 9 \ - I \ \ ,',' 'J' , '. <, V /. / ,:: r ) ,.~ 11 10 Head lind n'l'Me Icnl1ll1aha: 0·9• (,' re/>/lrraw. 10·11 • ( . tori1111, IJ .~ • .. "1) . : , > ., , • ,"JI , .. ~ :.; J 12 13 , ,J \ 14 !'Igurc, t2-15 .\Ialc \crll1lOalia: 12-14. C. re.\i/1/co/IJ. 15. C. r("I/ll/c%ide,\. 14 "\' \,' , , \~ , ~ 16 19 I' :,'1 j /' "/ 21 20 hgurc\ 10·21 :'>Ialc lCl'nll{1.1ha ,Inti O\'lposllor: 10. C. Cafl{/ic/tpl!l; 17, C bredspalll/a; 111·19. C. plllii/1o{iis; 20-21. C. resillico/a, 15 23 24 25 26 hgure, :'.~-2(i Pupal head (\cntral VIC\\ I and pronl1(um (anterior VICW): 22-23. C. candidipes; 24, C. piniinopis; 25. C. resinicola: 26. C. (Prli!t.\ 16 -!'- 27 ,;.. ",. "' I" ..... -~ 28 30 l I I 29 31 hgurt:~ 2;·.\1 1anat: and lanai tcrl11l1lal s 17 33~ 34 j \ 37 41 \? 42 hgurc\ ~2-12 1anal terminal ~t!g!11cnts and ~ternal spatulac: 32-33. C.l()r/III/lleltls; 34-35. C. res/nic()lo/des; 36-37. C. ct/luJ/dip<,s; JX-40. C p",iil1opis; 41-12. C. hr(ll'ispafllia.