Vol. 53 OCTOBER 1977 No. 4
THE PAN-PACIFIC ENTOMOLOGIST
MARI MUTT — The Taxonomic Status of A/loscopus and Redescriptions of its two
Species (Gollembola: Entomobigidae)s 5. «opie Oe a ee ales oe ce ee 241 LANDA — An Unusual Ant Nest Morphology for the Ant Formica fusca Linne in Western Oregon (Hymenoptera: formicidae .........................0.. 250
HADDOCK — Larval Morphology, Behavior and Distribution — useful tools in the separation of two closely related Nectopsyche Caddisflies (Trichoptera:
IEXeY ON WOXCLSTTOREYE) aero cee Madcon te aEIC ocak oh rr aemera Maees fore e en Ain ah ae Rat 253 OETTING — Immature Stages and Biology of Chionodes psiloptera a Pest in Bluegrass Seed Fields (Lepidoptera: Gelechiidae)....................... 258 CHEMSAK — Records and Descriptions of some Mexican Species of the Genus Phaea Newman (Coleoptera: Cerambycidae).............. 2.00 ee eee eee 269 HESPENHEIDE and RUBKE — Prey, Predatory Behavior, and the Daily Cycle of Holopogon wilcoxi Martin (Diptera: Asilidae)................... 0. eee ee eee 277 ALLEN — New Species of Ephemerella (Ephemerella) from Alberta (Ephemeroptera: Ephemerellidae)s etek. 222 nested ee or ee ee Fn 286
IRWIN — A New Genus and Species of Stiletto-flies from Southwestern North America with Close Affinities to Chilean and Australian Genera (Diptera:
MNSKeVIidaeATHEreVinae) ae. sie. sp Lew cae a alegre eel oes Spree reps 287 WHARTON — Exodontiellini, a New Tribe of Opiinae with Exodont Mandibles
(Hymenoptera: Braconidae)s An. 5 fe tia. en ee aes os ks ger eee 297 ROCKWELL and GROSSFIELD — Courtship Behavior of Scaptomyza (Bunostoma)
australis Malloch (Diptera: Drosophilidae)..............-....-.--220--05. 305 SCIENMIBIGINOMES pre rec. Nats Wn nae CaO noe Cees, neces er eaeee 257, 268, 285, 303 REGEN BIER AsiU) Riera ra, acc c.citcs: «cto ttc mer eee ans hae fe a ol ae near este 311 ZO OLOGICAENOMENGEATUIRIE® - ki irae = teen nel Penuninayar ..ps28rs ee atnmee: 252, 276 [S| CIES AP tf Segre i ce at aR An emer 7 Ae Le oi «oN ee PR see. 312 EDIM@REARINOMIGES pa v0. eat, Alsat. SR ieee sae Nee ok ging esol 249
SAN FRANCISCO, CALIFORNIA ¢ 1977
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Vol. 53 OCTOBER 1977 No. 4
The Taxonomic Status of Alloscopus and Redescriptions of its Two Species
(Collembola: Entomobryidae) José A. Mari Mutt
Department of Entomology, University of Illinois and Illinois Natural History Survey, Urbana, 61801
The taxon Alloscopus was erected by Borner in 1906 although the first author to study a member of this group was probably Oudemans (1890). This author studied a specimen which he identified as Temple- tonia (=Heteromurus) sp. The specimen was very deteriorated but Oudemans noted the presence of scales, a single eye on each side of the head and the relative lengths of the third and fourth abdominal segments. Oudemans probably had before him a specimen of Alloscopus tenuicornis Borner (1906) as his sample came from the type locality of that species.
Alloscopus was established by Bérner (supra cit.) as a subgenus of Heteromurus Wankel to house two Javan species: Heteromurus (Alloscopus) tenuicornis and H. (A.) tetracanthus. Handschin (1928) in a list of the Collembola of Indonesia, gave A//oscopus full generic status. The latter action has gone unchallenged to this day.
For sixty years A/loscopus harbored only the two original species until 1966 when Winter described A/loscopus strebeli from Peru and Izarra (1972) added Al/loscopus platensis from Argentina.
| have studied a series of specimens of this taxon from several islands of Micronesia and the results of this study coupled with a review of the literature has allowed me to arrive at the following con- clusions regarding the taxonomic status of Alloscopus and the species heretofore placed therein. First, A/loscopus is, de novo, re- garded as a subgenus of Heteromurus; second, the Philippine specimens reported as A//loscopus tetracanthus by Gapud (1971) actually are Heteromurus (Alloscopus) tenuicornis. Third, Alloscopus strebeli Winter is transferred to the genus /ndoscopus Prabhoo on account of the presence of a postantennal organ in this species; and
The Pan-Pacific Entomologist 53:241-249 October 1977
242 THE PAN-PACIFIC ENTOMOLOGIST
finally, Alloscopus platensis |Izarra is moved to the genus Dicrano- centrus Schott.
The evidence that lead to these conclusions will be treated below and under the discussion of the particular species. Specimens reported upon in this paper have been distributed among the follow- ing institutions: United States National Museum of Natural History, Washington, D.C., the Field Museum of Natural History, Chicago, Illinois; the Bernice P. Bishop Museum, Honolulu, Hawaii; and the Illinois Natural History Survey, Urbana.
The author wishes to thank Dr. Peter F. Bellinger for kindly review- ing the manuscript and for the loan of Hawaiian Heteromurus (Alloscopus) tenuicornis.
The Taxonomic Status of Alloscopus Borner. As noted earlier, Alloscopus was originally conceived as a subgenus of Heteromurus, to be separated from Heteromurus s. str. by the presence of dental spines in Alloscopus. Although Handschin (1928) gave no explanation for his action in raising Alloscopus to generic status, it is easy to infer that he regarded the presence or absence of dental spines as a character of generic rank. This, in fact, is the opinion of most present day workers (see discussion by Salmon 1964: 102) even though it is known that the closely related genus Dicranocentrus harbors species which either have or lack dental spines.
A second characteristic that has been invoked to separate Alloscopus from Heteromurus is the presence of large teeth (wing like teeth) on the unguis and unguiculus. This characteristic is not diagnostic because there are species of Heteromurus with wing like teeth on the unguis (e. g. Heteromurus stannardi Mari Mutt 1977) or on both unguis and unguiculus (e. g. Heteromurus dubius Delamare Deboutteville and Paulian 1952, Barra 1968). Ungual wing like teeth are also present in the genus /ndoscopus.
| have found a feature that can be used to separate A//oscopus and Heteromurus. Alloscopus species lack macrochaetae on the posterior dorsal medial part of the head (Figs. 1 & 2). At least two macrochaetae are present on this area in the species of Heteromurus which | have seen to date (i. e. Heteromurus diommatus, major, nitidus, peyerimhoffi, schoetti (Fig. 3), sexoculatus, stannardi, variabilis, and three unde- scribed species).
In view of the above discussion and the fact that in the genus Dicranocentrus (Mari Mutt, in press) we find a group of Central American and Caribbean species which lack macrochaetae on the posterior portion of the head, | am considering A//ioscopus a subgenus of Heteromurus. The head and body macrochaetotaxy of both sub- genera is completely similar.
Heteromurus Subgenus Al//oscopus
Alloscopus Borner 1906: 177, as subgenus of Heteromurus Wankel. Type Species: Heteromurus (Alloscopus) tenuicornis Borner, l.c.,
VOL. 53, NO. 4, OCTOBER 1977 243
subsequent designation by Salmon 1964: 478.
Member of the family Entomobryidae, subfamily Entomobryinae, tribe Orchesellini. Antennae five segmented due to the subdivision of the first segment (Ant. 1). Ant. 4&5 may be distinctly annulated or at least with many circular, evenly spaced, whorls of short setae along their length. Antennae about half the length of head and body combined. Postantennal organ absent.
Head with no more than one eye on each side or eyeless. When eyes are present they may or may not be surrounded by red-brown pigment. Head bears upon its anterior dorsal half many macrochaetae and a pair of lasiotrichia (Figs. 1 & 2). Medial posterior part of head without macrochaetae. Head and body covered with apically rounded or truncate scales. Antennae and legs possess scales but the number of these is dependent on age, increasing as the animal grows older. Scales absent from Ant. 4 & 5 and from tibiotarsi.
Tibiotarsi with or without rows of smooth setae along inner margin. Tibiotarsus of
metathoracic legs with a smooth seta (supra empodial seta) opposite the tenent hair. Posterior pair of legs with a well developed trochanteral organ (Fig. 8).
Fourth abdominal segment dorsally about 1.5 times longer than the third. Dentes with a single row of spines, the number of which is variable. Mucrones always without a basal spine.
Diagnosis: The subgenus may be separated from Heteromurus s. str. by the presence of dental spines in A/loscopus and by the absence of macrochaetae on the posterior dorsal half of the head also in AllosSCoOpus.
Key to the Species of the Subgenus Alloscopus
Dorsal macrochaetotaxy of meso and metathorax as in figure 4; base of the labium devoid of scales (Fig. 6); inner margin of tibiotarsi with two rows of smooth setae; shape of inner ungual teeth as in ct RL 1S ee ne a ee ie eee eae tetracanthus Borner
Dorsal macrochaetotaxy of meso and metathorax as in figure 5; base of the labium on each side with a pair of scales (Fig. 7); inner margin of tibiotarsi devoid of smooth setae; shape of inner ungual téeth as (n figure 14 san sa ene ce ede was tenuicornis Borner
Heteromurus (Alloscopus) tetracanthus BOrner
Heteromurus (Alloscopus) tetracanthus Bérner 1906: 177. Handschin 1925: 266.
Heteromurus tetracanthus Borner. Imms 1912: 120.
Alloscopus tetracanthus Borner. Handschin 1928: 267. Yosii 1959: 38. Yosii 1960: 27. Salmon 1964: 478. Yosii 1966a: 50. Yosii 1966b: 370. Winter 1966: 167, 168. Prabhoo 1971: 33. Gapud 1971: 11. Izarra 1972: 539.
This species was redescribed by Yosii (1959) from Malaysian specimens. Prabhoo (1971) reported the species from India, dis- cussed the dorsal body macrochaetotaxy and provided a habitus drawing and other drawings of the claw and mucro. Gapud (1971) reported the species from India, discussed the dorsal body
244 THE PAN-PACIFIC ENTOMOLOGIST
Figs. 1-7, Heteromurus sp. Fig. 1. H. (Alloscopus) tetracanthus-Dorsal head macrochae- totaxy and distribution of lasiotrichia, each dot represents one seta. Fig. 2. H. (A.) tenuicornis-Dorsal head macrochaetotaxy and distribution of lasiotrichia. Fig. 3. H. (Heteromurus) schoetti Denis-Dorsal head macrochaetotaxy and distribution of lasiotrichia based on specimens from Guatemala. Fig. 4. H. (A.) tetracanthus-Dorsal body macro- chaetotaxy. Fig. 5. H. (A.) tenuicornis-Dorsal body macrochaetotaxy. Fig. 6. H. (A.) tetracanthus-Chaetotaxy of base of labium. Fig. 7. H. (A.) tenuicornis-Chaetotaxy of base of labium.
VOL. 53, NO. 4, OCTOBER 1977 245
macrochaetotaxy and provided a habitus drawing and other drawings of the claw and mucro. Gapud (1971) also redescribed the species based on Philippine specimens but it is now evident that he was referring to Heteromurus (Alloscopus) tenuicornis. Gapud (supra cit.) noted several differences between: his specimens and Yosii’s redescription but did not ascribe specific importance to these. The species is redescribed below based on specimens from Micronesia (Ponape). For a diagnosis see the key to the species.
Habitus typical of the genus. Length excluding antennae and furcula up to 2.1 mm. Body devoid of dark pigment, specimens in alcohol white, yellow or brown depending on condition of preservation. Head, body and appendages covered with scales. Apical two antennal segments, tibiotarsi and dorsal portion of furcula devoid of scales. Collophore with very few scales. Ant. 5 distinctly annulated, Ant. 4 not clearly annulated but with setae arranged in whorls as in Ant. 5, latter segment without apical bulb.
Prelabral setae four in number. Labral chaetotaxy follows formula 5,5,4; all prelabral and labral setae smooth. Labral papillae well developed into cone like bodies (Fig. 16). Differentiated seta of the outer labial papilla apically rounded, not surpassing the apex of its papilla. Chaetotaxy of labial base as in figure 6. Arrangement exhibits the following variability: Seta 1 may be smooth or ciliated. Seta 3 may be present or absent, when present it may be smooth or ciliated. Setae A-C and 1,2,4 are always present regardless of the size of the specimen. Setae of venter of head smooth. One eye on each side of the head or the eyes may be absent.
Head and body macrochaetotaxy and distribution of lasiotrichia as in figure 1. The arrangement of these setae is constant in all specimens examined. Metathoracic leg with trochanteral organ of about 15 setae (Fig. 8). All tibiotarsi with two rows of smooth setae along inner margin; the number of these setae varies with age, largest specimen has 15 setae. Claw structure as in figure 13. Shape of ungual teeth and tenent hair constant in all specimens examined for this character (N = 40). In one specimen the unguis of one of the mesothoracic legs bears a median unpaired tooth upon its inner surface. All other ungues of same specimen bidentate. Another specimen has all its ungues quadridentate. A smooth seta present opposite tenent hair of metathoracic legs.
Corpus of tenaculum with a single smooth seta. Rami four toothed.
Manubrium dorsally with four or five pairs of smooth setae (only three pair shown in figures 11 & 12). Proximal portion of dentes bears one of these setae and from 4-7 spines. Mucro bidentate, devoid of basal spine.
Material Examined: Micronesia, Caroline Islands, Ponape, Mt. Tamatansakir, 180 m., Jan. 16, 1953, on Asplenium, J. L. Gressitt. 1 specimen. Ponape, Colonia, Agr. Exp. Sta., Jan. 6-17, 1953, berlese from woody compost, coffee litter, coconut litter and rotten palm, J. L. Gressitt. Over 500 specimens. Both this species and tenuicornis occur sympatrically in Ponape. Since both can be separated only after specimens are mounted in slides, and since it is undesirable to mount all specimens, it is not possible to give the definite number of specimens belonging to this species collected in Ponape. Approximately 90% of the specimens mounted from Ponape are tetracanthus.
Geographic Distribution: Java (Borner 1906), Malaysia (Yosii 1959), Singapore (Yosii 1959), India (Yosii 1966b, Prabhoo 1971), New Britain (Yosii 1960), Micronesia (Ponape) (new record).
Heteromurus (Alloscopus) tenuicornis Borner
Heteromurus (Alloscopus) tenuicornis Borner 1906: 177. Handschin 1925: 244, 266. Handschin 1926: 460. Heteromurus tenuicornis Borner. Imms 1912: 120.
THE PAN-PACIFIC ENTOMOLOGIST
|
Figs. 8-16. Heteromurus sp. Fig. 8. H. (A.) tetracanthus-Trochanteral organ present on metathoracic legs. Fig. 9.H. (A.) tenuicornis-Abnormal metathoracic leg. Fig. 10.H. (A.) tenuicornis-Normal metathoracic leg. Fig. 11. H. (A.) tetracanthus-Dorsal view of the dis- tribution of smooth setae on manubrium and proximal portion of dentes. Fig. 12. H. (A.) tetracanthus-Lateral view of the distribution of smooth setae on manubrium and proximal portion of dentes. Fig. 13. H. (A.) tetracanthus-Claw morphology of metathoracic legs. Fig. 14. H. (A.) tenuicornis-Claw morphology of metathoracic legs. Fig. 15 H. (A.) tenui- cornis-Variation in the relative position and shape of the external labial papilla and its differentiated seta. Fig. 16.H. (A.) tetracanthus-Labral papillae.
VOL. 53, NO. 4, OCTOBER 1977 247
Alloscopus tenuicornis Borner. Handschin 1928: 267. Handschin 1930: 422. Salmon 1964: 478. Yosii 1966: 49. Winter 1966: 167. Gapud 1971: 12. Izarra 1972: 539.
Alloscopus tetracanthus B8rner sensu Gapud 1971: 11, nec. Bérner 1906: 177.
This species was redescribed by Handschin (1925) from Javan specimens and Gapud (1971) presented a translation of that re- description. Through the kindness of Dr. C. Baroni Urbani and the Naturhistorisches Museum, Basel, Switzerland, | have studied the three topotypical specimens upon which Handschin (1925) based his redescription. The latter were found to be cospecific with a series of Micronesian specimens and upon both sets of specimens | base the redescription presented below.
Habitus typical of genus. Length excluding antennae and furcula up to 1.7 mm. Colora- tion, distribution of scales, structure of Ants. 4-5, labral chaetotaxy, and structure of labral papillae as in tetracanthus. Relative position of external labial papilla and its seta as in figure 15. Chaetotaxy of base of labium as in figure 7. In this species setae 3 & 4 of tetracanthus have given way to a pair of scales. Setae of venter of head smooth. | was unable to locate eyes but Gapud (1971) found one eye on each side of head.
Head and body macrochaetotaxy and distribution of lasiotrichia as in figure 5. Trochanteral organ as in tetracanthus. Tibiotarsi devoid of smooth setae with the excep- tion of the opposite seta to the tenent hair found on the metathoracic legs. Claw structure as in figure 14. Morphology of ungual teeth and tenent hair constant in 40 specimens examined for this character. Three specimens possess a third distal unpaired tooth upon the ungues of all legs. Another specimen had a tridentate unguis in only one of the mesothoracic legs. No quadridentate ungues were found.
Tenaculum, distribution of smooth setae on the dorsum of manubrium, dental spination, and mucronal morphology as in tetracanthus. Javan specimens with 6-10 dental spines. Micronesian specimens with no more than 7 spines.
It may be of interest to note here that two specimens were found with an abnormal metathoracic leg while the other member of the pair was completely normal (Figs. 9 & 10). For adiagnosis of this species see the included key.
Material Examined: Java, Tjibodas, under leaves in tropical rain forest, August 19, 1921. 2 specimens. Tjibodas, May 1922, 1 specimen. Material from Java deposited in the Natur- historisches Museum, Basel, Switzerland. Micronesia, Mariana Islands, Guam, 1 km. S. E. of Asan, 200-250 m., Nov. 1, 1947, H. S. Dybas. 30 specimens. Palau Islands, Yap Group, Yap |., Gagil and Ruul districts and Kanif, leaf litter, Jun. 30-Aug. 21, 1950, R. J. Goss. 23 specimens. Yap. |., Yap Hill, behind Yaptown, 50 m., Dec. 2-3, 1952, J. L. Gressitt. 5 specimens. Babelthuap, E. Ngatpang, 65 m., Dec. 7-9, 1952, J. L. Gressitt. Over 1,000 specimens. N. W. Aulup Tagel, 25 m., Dec. 13, 1952, leaf litter, J. L. Gressitt. 1 specimen. Caroline Islands, Truk, Tol |., Mt. Unibot, Dec. 31-Jan. 24, 1952, leaf litter, J. L. Gressitt. 14 specimens. Ponape-see under material examined for tetracanthus. Kusaie, Mutunik, 22m., Jan. 26, 1953, rotten palm stump, J. F. G. Clarke. 30 specimens. Kusaie, Hill 1010, 300 m., leaves and thrash from forest floor, J. F. G. Clarke. 2 specimens. Kusaie, 22 m., Feb. 1, 1953, rotting coconut log, J. F. G. Clarke. 1 specimen. Kusaie, Jan. 1953, rotting wood of breadfruit tree, J. L. Gressitt. 1 specimen. Hawaii, Oahu, Kuneuche banana plantation, Jan. 4, 1969, P. M. Lee. 5 specimens. Oahu, Pukijaga forest preserve, March 9, 1969, P. M. Lee 6 specimens.
Geographic Distribution: Java (B6rner 1906, Handschin 1925), Sumatra (Handschin 1926), Philippine Islands (Handschin 1930, Gapud 1971, as Allosco-pus tetracanthus), Micronesia (Ngatpang, Babelthaup, Yap, Guam, Truk, Ponape, Kusaie) (new records), Hawaii (new record).
248 THE PAN-PACIFIC ENTOMOLOGIST
The Javan specimens at hand are not well preserved but it was possible to study the claw structure, tibiotarsal clothing, some aspects of the body chaetotaxy and the dental spine arrangement. It is the number of these spines which have been used to separate tenuicornis and tetracanthus.
The Micronesian tenuicornis never have more than seven dental spines. One of the Javan specimens of this species has six spines on one dens and nine on the other. A second specimen has nine and ten spines respectively. Following the lead of Gapud (1971) | regard this difference as geographic variation. Such intraspecific variation | have observed in several species of Dicranocentrus.
Handschin (1930) reported H. (A.) tenuicornis from the Philippine Islands but Gapud (1971) did not encounter the species there even though he collected in the area from where Handschin’s specimens came. It is clear now that Gapud did collect tenuicornis but since his specimens had no more than seven dental spines they were called tetracanthus. Two of the three Javan specimens bear the label ‘‘more of tetracanthus Borner, V. Gapud ’68’’.
Indoscopus strebeli (Winter), new combination Alloscopus strebeli Winter 1966: 165. Izarra 1972: 539.
This species would fall within the definition of the subgenus Alloscopus were it not for the presence of a postantennal organ (PAO). Winter was aware of the significance of this being the first orcheselline species with such structure.
Prabhoo (1971) erected the genus /ndoscopus for two Indian species (I. spinosus and |. aspinosus). The genus was to be separated from Alloscopus by the presence of a PAO and by the absence of wing like teeth on the unguiculus.
Winter’s species differs from /ndoscopus by possessing wing like teeth on the unguiculi. | feel that this difference is outweighed by the presence of the PAO in all three species. /. strebeli is known from Peru.
Dicranocentrus platensis (Izarra), new combination Alloscopus platensis |zarra 1972: 537-539.
In arevision of the genus Dicranocentrus, now in press, | discuss in detail the reasons that justify the placement of this species in Dicranocentrus. Briefly these are: Antennae six segmented, claws without wing like teeth, and eight eyes on each side of the head. The species is known from Argentina.
VOL. 53, NO. 4, OCTOBER 1977 249
Literature Cited
Barra, J. A. 1968. Contribution a l’étude du genre Heteromurus Wankel, 1860 (Collem- boles). Biol. Gabonica 4:105-117.
Borner, C. 1906. Das system der Collembolen, nebst beschreibung neuer Collem- bolen des Hamburger Naturhistorisches Museums. Mitt. Naturhist. Mus. Hamburg 23:147-188.
Delamare Deboutteville, C. & R. Paulian. 1952. Faune des nids et des terriers en Basse Cote d’lvoire. Encyclopedie Biogeogr. Ecol. Vil, Paul Le Chevalier, Ed. 12, rue de Tournon-Paris (VI*) 1952 116 pp.
Gapud, V. P. 1971. Studies on Philippine Collembola, III. Suborder Arthropleona: Entomobryidae, with a checklist of Philippine Collembola. Philip. Entomol. 2:1-50.
Handschin, E. 1925. Beitrdge zur Collembolenfauna der Sundainseln. Treubia 6: 225-270.
Handschin, E. 1926. Ost-Indische Collembolen. Ill. Beitrag zur Collembolen-fauna von Java und Sumatra. Treubia 8:446-461.
Handschin, E. 1928. Collembolen aus Java, nebst einem beitrag zu einer monographie der gattung Cremastocephalus Schtt. Treubia 10:245-270.
Handschin, E. 1930. Philippine Collembola, Il. Material collected by the late Charles Fuller Baker. Philip. J. Sc. 42:411-428.
Imms, A. D. 1912. On some Collembola from India, Burma, and Ceylon; with a catalogue of the oriental species of the order. Proc. Zool. Soc. London 6:80- 125.
Izarra,D.C.de 1972. Alloscopus platensis, una nueva especie de La Plata, Provincia de Buenos Aires (Collembola, Entomobryidae). Physis 31:537-539.
Mari Mutt, J. A. 1977. A new species of Heteromurus from the Solomon Islands (col- lembola: Entomobryidae). Pan-Pacific Entomol. 52:326-330.
in press, A Revision of the genus Dicranocentrus Schott (Insecta: Collembola: Entomo- bryidae). Bull. Agr. Exp. Sta. Univ. Puerto Rico.
Oudemans, J. T. 1890. Apterygota des Indischen Archipels. In Weber, Zool. Ergeb. Niederlandisch Ost-Indien 1:73-91.
Prabhoo, N.R. 1971. Soil and litter Collembola of South India I|-Arthropleona. Oriental Insects 5:1-46.
Salmon, J. T. 1964-1965. An Index to the Collembola. Bull. Roy. Soc. New Zealand 7:1-651.
Winter, C. 1966. Beitrage zur kenntnis der neotropischen Collembolenfauna. Entom. Zeit. 76:165-169.
Yosii, R. 1959. Studies on the Collembolan fauna of Malay and Singapore, with species reference to the genera: Lobella, Lepidocyrtus and Callyntrura. Contr. Biol. Lab. Kyoto Univ. 10:1-65.
Yosii, R. 1960. On some Collembola of New Caledonia, New Britain and Solomon Islands. Bull. Osaka Mus. Nat. Hist. 12:9-38.
Yosii, R. 1966a. Check list of Collembolan species reported from Indonesia. Treubia 27:45-52.
Yosii, R. 1966b. On some Collembola from Afghanistan, India, and Ceylon collected by the Kuphe expedition, 1960. Results Kyoto Univ. Sc. exped. Karakoram and Kindukush, 1955 8:333-405.
EDITORIAL NOTICE
The editors would like to acknowledge with our great appreciation the many entomologists who served as reviewers for the past year.
An Unusual Ant Nest Morphology for the Ant Formica fusca Linne in Western Oregon’
(Hymenoptera: Formicidae)
Edward R. Landa Oregon State Univ., Corvallis, 97331
The perturbation of soils by ants has long been recognized. Surfi- cial ant nests developed in soils are generally conical - or crater- shaped mounds. | have observed large ant nests in western Oregon which have a cylindrical morphology that has not to my knowledge been previously reported.
These indurated soil structures (Fig. 1) seen in the Willamette Floodplain Natural Research Area of the William L. Finley National Wildlife Refuge in Benton Co., Oregon (T13S, R5W, Section 28), are developed on an uncultivated Dayton silt loam (Typic albaqualf, fine, montmorillonitic, mesic) which shows distinct mottling at 5 cm depth, indicative of poor drainage. Standing water is frequently seen on this iste during the winter, high rainfall period. The density of ant galleries is greatest in the inner portion of the cylinder, and the inha- biting ants were identified as Formica fusca L., the silky ant common across sub-arctic Canada and the northern half of the contiguous United States.
The structures observed are up to 50 cm high and 130 cm in circum- ference, and support grasses rooted in both the top- and side-walls. Excavation of several of the nests failed to reveal any cylindrical precursor which may have acted as a template for the construction. The high areal density of these nests became apparent following an experimental vegetation burning trial in 1976 (Fig. 2). AS many as 106 nests/1000 m? have been observed in the burn area. However, similar nests are seen in the adjacent undisturbed area.
Formica fusca is Known to nest in soil, under logs and rocks and in wood (Wheeler and Wheeler, 1963). The earthen mound nests of this species are generally low relief structures. Wheeler and Wheeler (1963) report F. fusca earthen mounds in North Dakota to measure from 15 to 150 cm in diameter but only 10 to 12 cm in height. Wiken et al. (1976) have described low conical structures developed by F. fusca in a well-drained gravelly loam in southwestern British Columbia. The elevated nest structures observed on this Willamette Valley site are probably adaptations to the poorly-drained nature of the area, with the upper portion of the mound affording the inhabitants shelter from the periodic flooding of the lower reaches of the nest. In addition,
1Contribution from the Oregon Agr. Expt. Sta. Technical Report Series No. 4679.
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VOL. 53, NO. 4, OCTOBER 1977 251
# NY ;
: hoe : : % i Eanes 3 abies ee SESE Roe
Fig. 1. Ant nest (each unit on scale is 10 cm). Vegetation on side-walls has been trimmed to show nest structure. Fig. 2. Distribution of nests on the landscape.
252 THE PAN-PACIFIC ENTOMOLOGIST
such nesting structures may provide for a more favorable tempera- ture regime than found in the adjacent soil (Cole, 1932; Scherba, 1962).
Acknowledgements
The invaluable assistance of Dr. George C. Wheeler Desert Re- search Institute, University of Nevada-Reno, and Drs. Paul O. Ritcher and George R. Ferguson, Department of Entomology, Oregon State University is gratefully acknowledged.
Literature Cited
Cole, A.C., Jr. 1932. Observations on semi-desert ants. Ohio J. Sci., 32:533-537.
Scherba, G. 1962. Mound temperatures of the ant Formica ulkeli Emery. Am. Midland Na- turalist, 67-373-385.
Wheeler, G.C. and Wheeler, J. 1963. The ants of North Dakota. Univ. N. Dak. Press; Grand Forks, ND.
Wiken, E.B., Broersma, K., Laukulich, L.M. and Farstad, L. 1976. Biosynthetic alteration in a British Columbia soil by ants (Formica fusca Linne). Soil Sci. Soc. Amer. J., 40:422-426.
THE INTERNATIONAL CODE OF ZOOLOGICAL NOMENCLATURE
The draft third edition of the International Code of Zoological Nomenclature is now abailable for comment by zoologists. Copies may be obtained (price £2.50 surface mail, £5.00 air mail) from the Publications Officer, International Trust for Zoological Nomencla- ture, c/o British Museum (Natural History), Cromwell Road, London SW7 5BD, U.K. Comments should be sent as soon as possible, and in any case before 30 November 1978, to the Secretary, International Commission on Zoological Nomenclature, at the above address.
A paper explaining the major changes proposed by the Com- mission’s Editorial Committee to the existing Code has been publish- ed in the Bulletin of Zoological Nomenciature, vol. 34, part 3. Copies may be obtained (price 50p) from the same address as copies of the draft Code.
Larval Morphology, Behavior and Distribution — useful tools in the separation of two closely related Nectopsyche Caddis Flies
(Trichoptera : Leptoceridae) James D. Haddock Indiana University-Purdue University at Fort Wayne, Indiana 46805 The genus Nectopsyche, Muller 1879, recently known as (Lep- tocella, Flint, 1974) is presently represented on the north american continent by twelve species (Haddock, in press). The majority are lowland inhabiting species having a broad, transcontinental distribution as in Mystacides (Yamamoto and Wiggins, 1964) and related genera. Ross (1944) has discussed the eastern members of Nectopsyche occurring in Illinois. A clear delineation of species limits had not been previously understood because of the lack of diagnostic information provided by adult genitalia and the paucity of larval material and pinned adults — particularly from the western U.S.
Adults of the genus Nectopsyche are easily recognized in the field because of the narrow, elongate, conspicuously patterned wings that are characteristic of most species. The antennae in the males are longer than the body length. Two frequently confused species, as evidenced by misidentified museum material are Nectopsyche diarina (Ross) and Nectopsyche albida (Walker) due to the very close similiarity in adult structures including wing color patterns, genitalia and the fact that they are sympatric over much of their range in the midwestern United States. In this paper easily identifiable characters present in the larvae and larval cases are given, Table 1, which readily identifies these two easily confused species. Complete taxonomic treatment of the 12 species in the genus is in press.
Larvae of Nectopsyche diarina can be identified by the presence of two dorsal, longitudinal, black stripes on the head, pronotum and mesonotum and the presence of trachael gills on the abdomen (Fig. 1). Nectopsyche diarina has a metathoracic swimming brush con- sisting of many setae located on the lateral and ventral margins of the femur and tibia which is used in locomotion. Larval case design is highly variable and consists of sand with plant fragments arranged either in a spiralling on non-spiralling whorl (Fig. 2). It is interesting that the spiral case is similar to that of some species in the related genus Triaenodes (Hickin, 1967, and others).
Nectopsyche albida larvae have two dorsal, longitudinal black stripes on the head only and do not possess abdominal tracheal gills (Fig. 3). Larvae are non-swimmers due to an absence of a metathoracic swimming brush. Larval cases are composed of sand grains and are therefore non-buoyant (Fig. 4).
Nectopsyche diarina occurs in lentic and lotic habitats east and west of the Rocky Mountains in the northern United States and
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254 THE PAN-PACIFIC ENTOMOLOGIST
Figs. 1&2; Nectopsyche diarina. Fig. 1, mature larva, dorsal view, X8.Fig. 2, larval cases, X6.
Canada. Many records occur from the Snake and Missouri river drainages. The headwaters of both systems are within a short distan- ce of the borders of Idaho, Montana and Wyoming. Miller (1958) has shown that a “well established” two-way faunal exchange has oc- curred between the headwaters of the two systems among certain primary freshwater fishes such as the mountain white fish, longnose dace and mottled sculpin.
VOL. 53, NO. 4, OCTOBER 1977 255
4
Figs. 3&4; Nectopsyche albida. Fig. 3, mature larva, dorsal view, X8. Fig. 4, larval case, X7.
Nectopsyche albida is almost entirely confined to lentic habitats and has been collected only east of the rocky Mountains. The southern extreme of this cold-water species occurs in central Missouri along the Missouri river, which is congruent with the roughly. east-west line that marks the maximum advance of the glaciers during the Pleistocene. | believe that following the Wiscon- sin glaciation, N a/bida dispersed northward into north central United States and southern Canada. The effect of glaciation on drainage reversals such as that suggested for the northward flow of the Upper
256 THE PAN-PACIFIC ENTOMOLOGIST
Table 1. Comparative Diagnostic Chart of Nectopsyche diarina and Nectopsyche albida.
N. diarina N. albida Larval Morphology dorsal color patterns black stripes on head black stripes on head and thorax only swimming brush on present absent metathoracic legs Larval Behavior short-term swimming yes no ability case design and sand grains and plant frag- sand grains only materials ments, arranged in spiral whorl or not Distribution habitat lentic and lotic primarily lentic dispersion east and west of Rocky east of Rockies only
Mountains
Missouri River (Thornbury, 1954) may also explain the distribution of both N. a/bida and N. diarina throughout central Canada today.
Acknowledgements
The assistance of Dr. D.G. Denning, Moraga, California in providing suggestions and material for study is appreciated. Additional material pertaining to the species discussed was loaned from the University of California, Berkeley (C.1I.S.), California Academy of Sciences, Illinois Natural History Survey, U.S. National Museum, Royal Ontario Museum and the Museum of Comparative Zoology.
Literature Cited
Flint, O.S. Jr. 1974. The Trichoptera of Surinam. Stud. fauna of Suriname and other Guyanas. Vol. XIV No. 55. 151 pp.
Haddock, J.D. The Biosystematics of the Caddis Fly Genus Nectopsyche in North America with Emphasis on the Aquatic Stages (in press Amer. Midl. Natur.)
Hickin, N.E. 1967. Caddis Larvae — Larvae of the British Trichoptera. Hutchinson and Co., LTD., London. 476 pp.
Miller, A.H. 1958. Origin and affinities of the freshwater fish fauna of Western North America in Zoogeography. Publ. No. 51 of Am. Assoc. Advan. Sci., Wash., D.C. 509 pp.
Muller, F. 1879. Ueber Phryganiden. Zool. Anz. 2:38-40.
Ross, H.H. 1944. The Caddis Flies or Trichoptera of Illinois. Bull. Ill. Nat. Hist.Surv., 23 (1). 326 pp.
VOL. 53, NO. 4, OCTOBER 1977 257
Thornbury, W.D. 1954. Principles of Geomorphology. John Wiley & Sons, Inc. New York. 618 pp.
Yamamoto T. and G.B. Wiggins. 1964. A Comparative Study of the North American
species in the caddisfly genus Mystacides (Trichoptera, Leptoceridae). Can. J. Zool. 42:1105-1126.
SCIENTIFIC NOTE
Notes on the 1973 Migration of Vanessa cardui (Lepidoptera: Nymphalidae). Notes on the 1973 Migration of Vanessa cardui (Lepidoptera: Nymphalidae). — Spring 1973 produc- ed an extensive northward migration of this butterfly through souther California and Nevada.
On 26 days, from March through May, the directions of flight of 5,361 individuals were recorded at Big Pine, Inyo County, California. At times, three or four directions were simultaneously involved with little or no activity between. The insects’ preferred direction of flight appeared to rotate nearly 135 degrees clockwise from west to north-northeast in
ten weeks. This trend is illustrated in the percentages that flew within 45 degrees of these two directions during six time-periods:
Mar. 18 Mar. 30 Apr. 8 Apr. 23 May 6 May 19 to to to to to to
Mar. 23 Apr. 4 Apr. 12 May 1 May 11 Jun. 2 Ww 89 51 39 6 9 6 NNE 0 4 40 53 66 78
No southward movement could be observed in summer or fall. Williams (1970, J. Lepidop. Soc., 24:165) states that southward migration is most often reported in the more easterly states. An increasingly eastward flight vector during late spring may carry large numbers away from the far west before southerly flight begins.
Occasional migrators were observed in Inyo County as early as February. The number crossing a 50-foot line reached a peak of over 50 per minute at Big Pine by mid-April; 30 per minute continued through 2 June beyond which they drifted undirectionally. Migratory flights persisted longer at elevations above 2500 m.
Numbers decreased as cooler temperatures were encountered; winds alone did not pre- vent flight. On 25 April, near Lida, Esmeralda County, Nevada, they flew northward against near-gale winds. Apparently having difficulty topping an east-west ridge they were funneled through a low pass and fanned out over the road within a foot of the surface ata groundspeed of less than 1 mile per hour. Counts across 10-foot lines showed that about 3 million per hour were crossing a mile’s length of road.
The insects were observed settling to the desert floor and low shrubs before nightfall. On 20 May, in Huntoon Valley, Mineral County, Nevada, | happened near dusk, to be driving northeastward, the preferred direction of flight that day. Those within 50 feet rose into the air and flew alongside. When | drove at their flight speed, about 15 miles per hour, a cloud of butterflies formed on either side which soon became too dense to see through. They settled back to earth when the car speeded ahead or turned away from their flight direction.
Large migrations seem to occur every 4 to 7 years. The previous mass movement was in 1968. Spring 1977 produced only very small numbers. The next heavy flight is likely to 0 c- cur in 1978 or 1979 and it is hoped that many persons will prepare themselves to accurate- ly record numbers and flight directions. Such counts are needed from many locations and dates before the insects’ behavior can be properly interpreted. — DERHAM GIULIANI, 770 Flower Alley, Big Pine, California 93513.
The Pan-Pacific Entomologist 53:257 October 1977
Immature stages and biology of Chionodes psiloptera a pest in bluegrass seed fields’
(Lepidoptera, Gelechiidae)
Ronald D. Oetting? Department of Entomology, Washington State University, Pullman, 99163
Chionodes psiloptera (Barnes and Busck) is a localized pest of Kentucky bluegrass (Poa pratensis L.) in the United States. The only record of C. psiloptera being a pest of an agricultural crop is in Spokane County, Washington. Damage in this area was first reported by Crawford and Harwood (1959). Prior to that time it was only reported from Canada and the host plant was unknown (Barnes and Busck 1920). In Spokane County it is an important pest of the Kentucky bluegrass seed producing industry. New plantings are especially vulnerable to damage by larvae feeding near the crown and established stands are also weakened and damaged to the point where seed can no longer be produced economically.
Materials and Methods
Field collected larvae and adults were brought into the laboratory for study. Rearing cages were 3/4 oz. wax lined paper creamers placed in environmental growth chambers. Growth chambers were set with a daily photoperiod of 16:8 LD with coincident temperatures of 21:10°C or 21°C and 30°C constant. Larvae were reared individually on germinated bluegrass or artificial diet similar to the diet described by Dupnik and Kamm (1970). Diet was cut into 1 cm cubes and placed on the bottom of the rearing cup.
Adults were kept in 1 qt. paper cartons with a clear plastic screen cover. The bottom of the carton was removed and the carton placed in a petri dish bottom containing a sheet of filter paper. Adults were fed a liquid diet consisting mainly of water, beer, and honey, (described by Reese et a/. 1972). The diet was placed in a glass vial containing a dental wick protruding through an opening cut in the plastic snap Cap.
An experiment was conducted to determine if females had a pre- ference for a particular type of surface for egg deposition. Strips of different oviposition substrates ca 2.5 x 10 cm were hung from the top of the carton sides. The substrates were wax paper, paper toweling, filter paper, bond paper, and corrugated cardboard. Eggs were re- Sees litic aabernerober 4714, Project 0236, Washington State University, Agricultural Research Center.
2Assistant Entomologist. Present address: Department of Entomology, University of California, River- side, California 92502.
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moved from the cartons and placed in stender dishes with moist filter paper bottoms. Newly hatched larvae were maintained in stender dishes with moist filter paper bottoms and entire newly germinated bluegrass plants were used as food source. Larval developmental data were obtained from individually reared specimens and number of instars determined by daily observation for shed of head capsules.
The site of larval feeding was studied in 2 containers. In the first container bluegrass was planted in soil in 10 glass containers with glass sides enclosing 1 cm of soil ca 15 cm deep and 30 cm long. Larvae were placed on the soil surface after the grass grew until roots reached the bottom of the container. In the second, washed plants with ca6 cm of leaf and stem and 3 cm of roots were placed in petri dishes and covered with moist filter paper. Ten larvae were placed in | each dish containing 6 plants replicated 4 times. Location of larvae and feeding was recorded at least 3 times daily.
Potential male sex attractant tests were obtained from Farchan Research Laboratory, Willoughby, Ohio and prepared immediately before placing them in the field. Ten p1 of each chemical were injected into OS-6 natural polyethylene closures for 60975-L vials (Kimble Products, La Mirada, California) or dissolved in 100 ,p1 methylene chloride and placed into the cup of a No. 1 red rubber sleeve stopper. Both carriers worked equally well during the test period. The carriers were placed in the center of a Howell trap coated with Stickem® (Howell 1972). Traps were placed in bluegrass seed fields ca 10 cm from the ground 50 m apart in one area and 25 m apart in a second location. Traps were charged with new carriers every 2 wks, and with new Stickem® every wk.
Descriptions were made from live and preserved specimens. Averages and ranges given in the descriptions were based on 10 measurements unless stated otherwise. Head capsule widths were measured at the vertex as seen from above.
Descriptions
There are usually 10 or 11 larval instars but occasionally 9 or 12 are present. Head capsule measurements of the instars are presented in Table 1. The last-instar is described along with a brief description of the egg, first-instar, and pupa.
Egg: 0.65 mm (0.56-0.81 mm) long, 0.35 mm (0.30-0.41 mm) diam. at widest point; generally oval but irregular, pearly white initially but changes to yellow.
1st instar: 1.77 mm (1.35-2.15 mm) long, head capsule width 0.20 mm (0.19-0.21 mm); head capsule and cervical plate yellowish-tan, shiny, body transluscent white, faint reddish bands on dorsum not noticeable in first instar but more distinct in 2nd and 3rd instar.
Last instar: Length 9.74 mm (8.34-11.83 mm), width 1.94 mm (1.84-
260 THE PAN-PACIFIC ENTOMOLOGIST
Table 1. Head capsule widths in mm of C. psiloptera larvae requiring 10, 11, or 12 instars for development.
Larvae with 10 instars Larvae with 11 instars Larvae with 12 instars Instar Indiv Range Mean Indiv Range Mean Indiv Range Mean
1st 50 = =0.19-0.21 0.20* 2nd 50 0.24-0.27 0.25* 3rd 40 0.28-0.34 0.29* 4th 40 0.37-0.44 0.40* Sth 40 0.45-0.58 0.50*
6th 17 =0.55-0.72 0.63 18 0.60-0.69 0.64 5 0.53-0.69 0.60 7th 17 0.69-0.83 0.78 18 0.69-0.84 0.76 5 0.63-0.83 0.73 8th 17 = (0.78-0.94 0.88 18 0.79-0.99 0.90 5 0.80-0.93 0.85 9th 17. 0.99-0.10 1.05 18 0.90-1.12 1.00 5 0.92-1.05 0.98 10th 17) -1.18-1.26 1.23 18 0.99-1.25 1.08 5 1.05-1.07 1.07 11th 18 9 =1.10-1.33 1.22 5 1.10-1.18 1.16 12th 5 1.22-1.34 1.30
*Head capsule widths for the first five instars were the same regardless of the total number of instars. Therefore the widths are all presented together.
2.00 mm); head capsule width 1.23 mm (1.10-1.34 mm, n = 40). Head capsule orange-brown blending into dark brown on posterior edge, around mouth parts, and on clypeus; conspicuous black area in ocellar region; ocelli arranged as in Fig. 1 and 2. Labrum width twice the length, notched (1/3 length) medial anterior margin, each lobe with 2 stout submarginal setae anterior, 2 lateral setae farther from the margin, 4 central setae. Mandible orange-brown, darker and reddish apically; teeth 5 (Fig. 3), 2nd and 3rd longest. Spinneret orange-brown, 7 times as long as diam, slightly longer than palpi. Maxillary cardo and stipes very light brown.
Body grayish white to cream, with reddish irregular blotchy bands covering half of the dorsal half of each segment from 2 posteriorly. Spiracles circular, peritreme fuscous, thoracic ca 0.06 mm diam, ist 7 abdominal ca 0.03 mm diam. Prothoracic shield sclerotized, yellowish-brown to brown, divided by non-sclerotized midline on dorsum. Anal plate light brown, setae arranged as in Fig. 5. Setae pale reddish, pinacula brown, ca 0.55-0.85 mm long, longest on sclerotized shields, arrangement as in Fig. 4. Thoracic legs light brown; ventral prolegs concolorous with body, crochets yellowish brown, uniserial, biordinal, arranged in acircle, about 12 large and 20 small (Fig. 6); anal prolegs light brown laterad, crochets biordinal, uniserial transverse band on anterior of proleg, directed caudad, ca 8 large and 8 small.
Pupa: Length 6.95 mm (6.12-7.32 mm), width 2.20 mm (2.04-2.46 mm). Head, thorax, and abdomen yellow-brown, shiny, changes to fuscous before adult emergence. Head smooth to rugulose, exposed maxillae 2.66 (2.40-2.82 mm) long; labial palpi very small (ca 0.03 mm) triangular area caudad labrum, maxillary palpi ca 0.53 mm long (Fig. 7). Thorax smooth to rugulose. Abdomen rugulose, spiracles ca 0.06 mm diam.
VOL. 53, NO. 4, OCTOBER 1977 | 261
Figs. 1-7. Last instar of C. psiloptera: Fig. 1. frontal view of head capsule; Fig. 2. lateral view of head capsule and anterior thorax; Fig. 3. mesal view of right mandible; Fig. 4. setal map (t?, T?, A’, A?, A‘); Fig. 5. dorsal view of anal plate; Fig. 6. ventral view of proleg.; Fig. 7. ventral view of cephalic segments of pupae.
Biology — Laboratory Studies
Adults. In the laboratory females preferred to lay eggs in masses attached to the grass stem near the ground. In some instances only single eggs were deposited but usually a female either scattered eggs singly or in small masses in the same night. The number of eggs
262 THE PAN-PACIFIC ENTOMOLOGIST
ranged from 1 to 77 and there was no trend for numbers to be high or low. However, egg masses usually ranged from 5 to 25 eggs and the 2nd largest egg mass contained 34 eggs. The duration between emergence to 1st egg deposition ranged from 3 to 15 days (n = 15) but most were 7 days or less with a mean of ca 4 days. A female laid more eggs, usually depositing eggs every night, the 1st 4 or 5 nights after oviposition commenced and less frequently thereafter. One female laid her last eggs 19 days after emergency. Fourteen females were dissected before egg laying commenced and the ovaries con- tained from 126 to 184 eggs (x = 147).
There was no preference for the different artificial substrates presented to females for egg laying sites. All substrates were used occasionally. However, the majority of the time eggs were oviposited at the bottom and under the side of the cage. The eggs were attached either to the bottom or the lower section of the side. A stem of bluegrass was attached to the side next to the bottom of the cage by tape. Females began attaching egg masses on the stem or next to the stem usually beneath the tape. A few egg masses were still deposited under the side as before. When females were placed in cages with growing plants, eggs were deposited on the stem near the ground.
Eggs. The egg stage required an average of 17.5 days (12 to 25 days) before hatching when temperatures were 21° day 10° night. Ata constant temperature of 21° only 10.3 days were required with a range of 8 to 15 days. Eggs maintained at 30° constant temperature hatched in 6.5 days, range 5 to 10 days. The hatching success was greater than 90% for all temperatures.
Larvae. First instars were placed in cups with artificial diet and reared at 21°. These larvae were placed on the bottom of the cup next to the diet cube. Many of the larvae crawled under or onto the diet or died apparently trapped by excessive moisture, 54% mortality was observed in the first instar and 51% in the second. A 27% mortality was observed for third instars and minimal mortality in later instars.
Mortality was decreased in the early instars by feeding these larvae on germinated bluegrass with a new leaf ca 2.5 cm long. The small larvae fed on the roots and leaves of the bluegrass but more often they burrowed into and fed within the seed coat. Larvae then re- mained within the seed coat through the third instar. If bluegrass seed was placed nearby, small larvae also burrowed into these seed and fed. When fourth instars were observed they were transferred to artificial diet. When held at 21° larval development took ca 140 days. The first two stages averaged from 6 to 8 days. From the third stage through the twelfth stage there was a broad range in the days re- quired to complete each stage. The average time required in each stage was between 10 and 18 days. The number of instars ranged from 9 to 12 but usually 10 or 11 instars were required to complete development. Larvae were also reared on germinated grass to compare development on natural food and artificial diet. There was
VOL. 53, NO. 4, OCTOBER 1977 263
no significant difference between larval size and duration of instars among larvae reared on these two food sources.
Larvae did not necessarily remain on the diet or grass throughout the day. They spun a silken tube which usually extended from the diet out foracm or less. This tube would sometimes be free from the diet along the edge of the bottom of the cup. During the day larvae were observed within this tube and left only when disturbed. Also silken chambers were spun in the diet where larvae had burrowed, and a silk cap covered the cavity. When this occurred grass was incorporated into the webbing, camouflaging the chamber. Pupation occurred either within webbing or free from webbing on the bottom of the cup.
The location of larval feeding on the plant and depth of larval move- ment in the soil are of major concern to effectively survey for damage and control these pests in the field. Larvae were observed in petri dishes containing bluegrass plants in the laboratory. In these obser- vations 56% of the time larvae were found in the root zone either in silken tubes or free. Twenty five percent of the time larvae were in the crown area and 19% of the time larvae were in the stem and leaf area. Feeding was observed 41% on the roots, 39% on the crown, and 20% on the leaves and stems. Feeding on the leaves and stems was both external and by burrowing into the stems, indicating that feeding is not limited to a specific area of the plant but most damage is next to the soil surface or just beneath the surface.
Larvae were also placed on growing plants in glass containers to observe feeding behavior. Most of the larvae fed on secondary roots just beneath the surface and remained in the upper 2.5 cm of soil in silk-lined tubes. A few plants were cut off at the surface and a few beneath the soil surface. No signs of feeding on the plants over 5mm above the ground were observed. One larva was observed 7 cm below the surface feeding on the root system. Most larvae pupated on the surface but a few were within acm of the surface. Pupae were usually free of webbing.
Pupae. Pupae were maintained at the same temperatures as the eggs and an increase in duration with decreasing temperature was similarly observed. At 21° day 10° night conditions the pupal stage averaged 21.4 days with a range of 17 to 23 days (n = 88). There was an 85% emergence success with a sex ratio of 69% males and 31% females. At 21° constant temperature the duration of the pupal stage averaged 12.4 days with arange of 10 to 14 days (n = 113). Emergence was 89% successful and 61% males and 39% females emerged. At 30° the average duration was 8.0 days with a range of 7 to9 days (n = 60). Seventy five percent emergence was obtained with 64% males and 36% females.
The duration of the egg and pupal stage in the field is affected by the temperature. Banerjee (1969) found similar results with another pest of bluegrass Crambus trisectus (Walker) with a shorter duration of each stage with increased temperature. Crambus trisectus is a
264 THE PAN-PACIFIC ENTOMOLOGIST
A STE
P a) 8-11 a ay ts a 3-4 eos se ke a
o sar]
1 Serre cea]
E Saaarw)
J F M A M J J A S 0 N D
Fig. 8. Field occurrence of life stages of C. psiloptera; (E) egg, (1-11) larval instars, (P) pupae, (A) adult.
univoltine species in the Pacific Northwest and a bivoltine species in Illinois. The temperature at the soil surface in the Spokane area in 1975 and 1976 was ca 15 to 20°C when pupation occurred and 21 to 25°C during egg laying.
Field Life History
Chionodes psiloptera is an univoltine species which overwinters as larvae (Fig. 8). Adults in the Spokane, WA area (latitude 48°N, altitude 738m) begin flying in the field from mid-May to early June, depending on temeperature, and are present for a month, usually reaching a peak in mid-June. In 1975 observed adult activity peaked on June 17 and in 1976 from June 11 to 17. Eggs are attached to stems in early June to early July and the first instars are present in mid-June to early August. Larvae feed on the roots, crown, and lower stems through summer and fall. In October or early November larval activity stops and they remain in their overwintering silk-lined casing within a stem, between stems in the crown, or in the thatch or upper 2.5 cm of the soil. AS soon as the upper soil reaches ca 15°C in the spring larvae start to feed. In late April or early May the last instars change into pre- pupae and pupate in 1 to 3 days.
Larvae have been collected from soil cores primarily from the foot- hills area northeast of Spokane, Washington. This is a nonirrigated area and the larvae are most prevalent on the dry slopes rather than
VOL. 53, NO. 4, OCTOBER 1977 265
Table 2. Chemicals tested for attractancy for C. psi/loptera in the field.
Avg catch/trap/week
Avg total Chemical June 11 June17 June 24 July 1 catch/trap Area 1 (Z,Z) & (Z,E)-7, 11-Hexadecadieny! Acetate 12.5 8.5 3.5 _— 8.79 Z-7, E-11-Hexadecadieny| Acetate 8.8 5.8 3.3 — 6.36 E-9-Dodeceny| Acetate 10.0 1.8 1.3 — 5.14 Check 7.0 1.0 0.0 — 3.29 Z-7-Dodecenyl Acetate 3.8 2.3 2.3 — 2.91 Z-7-Hexadeceny! Acetate 2.5 3.5 1.3 — 2.43 Z-9-Tetradecenyl Acetate 1.7 3.8 1.5 — 2.23 Z-11-Tetradecenyl Acetate 3.0 1.8 1.0 — 2.07 Z-9-Dodecenyl Acetate 2.8 3.3 2.0 — 1.21 Area 2 Z-7, E-11-Hexadecadienyl Acetate 0.0 0.5 6.0 4.5 3.67 Check 0.0 0.5 2.5 2.5 1.83 Z-7-Hexadecenyl Acetate 0.0 0.5 0.5 1.0 0.67 7,11-Hexadecadienyl Acetate 0.0 0.0 1.5 0.0 0.50 (Z,Z) & (Z,E) E-9-Dodecenyl Acetate — 0.5 0.0 1.0 0.50 Z-9-Tetradecenyl Acetate — 0.0 0.5 1.0 0.50 Z-9-Dodeceny! Acetate — 0.5 0.5 0.0 0.33 Z-7-Dodeceny| Acetate — 0.0 0.0 0.5 0.17 Z-11-Tetradecenyl Acetate —_— 0.0 0.0 0.0 0.0
the lower wetter areas. Larvae have also been collected in the irri- gated area of the Spokane Valley east of Spokane, Washington and nonirrigated fields around Freeman and Rockford, Washington. Larvae have been collected from several named varieties of Kentucky bluegrass including Adelphi, Merion, Fylking, Garfield, Nugget, and Glade.
When disturbed, adults fly short distances (usually less than a meter) and conceal themselves in the grass crown, under clumps of sod, under straw, or in cracks of the soil. They move quickly and are hard to detect. They fly close to the ground, seldom above the grass, and are hard to capture with a sweep net. These moths are most eas- ily observed fluttering from row to row and crawling around the young plants. |
No host other than grass is known for this species. In addition to bluegrass it has been reported to feed on creeping red fescue (Crawford and Harwood 1964). In the fall, areas around grass fields were examined to see if an alternate host could be found. Adults were not found in any areas other than within and near the borders of seed fields. Larvae were not found in any weedy areas or under deciduous
266 THE PAN-PACIFIC ENTOMOLOGIST
Fig. 9. Strip across bluegrass field treated with insecticide, with remainder of field show- ing damage by C. psiloptera.
shrubs. However, larvae which appeared to be Chionodes were found in debris under ponderose pine near grass fields. They were not found under pines further from seed fields. The following spring no gelechiid larvae were found in these areas.
Unsaturated acetate pheromones of some other gelechiids and moths were tested for attractancy to C. psiloptera (Table 2). There was consistently a significantly greater attraction to the traps containing gossyplure (Z,Z and Z E-7, 11-hexadecadieny| acetate in area 1. This is known to be the sex pheromone of another gelechiid, the pink boll- worm, Pectinophora gossypiella (Saunders) (Hummel et at., 1973). The sex pheromone of a second gelechiid, the angoumois grain moth (Sitotroga cerealella [Olivier]), Z-7, E-11-Hexadecadienyl acetate also consistently attracted C. psiloptera in areas 1 and 2. The check traps contained a high number of C. psi/loptera but the average was in- creased by a few traps containing a large number of this species. This could be the result of a female randomly being caught in the trap and attracting males in by the release of pheromones. A female moth was found in one replication of the check in area 1 on June 11. This check contained 27 moths. The other chemicals did not result in any signif- icant attraction. Gossyplure could be used in bluegrass fields as a monitoring tool for C. psiloptera activity and was a better indicator than a black light trap for monitoring populations in 1976.
Damage
Chionodes psiloptera feed in the crown area of Kentucky bluegrass similarly to other soil pests of this crop. However, damage is not as noticeable as that caused by pests such as sod webworms. Irregular
VOL. 53, NO. 4, OCTOBER 1977 267
brown areas typical of sod webworm damage are not present with C. psiloptera feeding. Instead there is a gradual weakening of the stand because only part of the crown is damaged by larval feeding. Signs of C. psiloptera feeding are thinning of the stand accompanied by the in- vasion of undesirable plant species. Damage is most noticeable in the fall when regrowth has started or in the spring before it is warm enough for regrowth to surpass damage. This was especially notice- able when part of a field was treated with an insecticide reducing damage (Fig. 9). Insecticide screening has been conducted to find a control for this pest (Oetting 1976).
The larvae of this species can damage newly planted Kentucky bluegrass severely. Bluegrass which is planted in the spring is still small in the summer (ca 10 cm high) and has not developed an exten- sive root system or crown. Damage at this time is more severe than when a stand has matured because the smaller plants are more af- fected by larval feeding.
Natural Enemies
There is no data on predation on this species. However, it was noted that numbers were not reduced by birds while sod webworms in the same field were significantly reduced. Among several hundred larvae brought into the laboratory and reared a single Copidosoma sp. (Encyrtidae) parasite was reared from a larva collected from the foot- hills area northeast of Spokane, Washington in April 1975. From this ca 30 adult Copidosoma emerged in May 1975. There are no other re- cords of parasites from this species.
Acknowledgements
| thank Drs. R.F. Harwood and R.D. Akre for critically reading the manuscript. | also thank Dr. G. Gordh of the USDA Insect Indentifica- tion and Beneficial Insect Introduction Institute for determinations. Financial support was provided by the Intermountain Grass Growers Association, Washington State Department of Agriculture, And Washington State University.
Literature Cited
Banerjee, A.C. 1969. Development of Crambus trisectus at controlled temperatures in the laboratory. J. Econ. Entomol. 62:703-705.
Barnes, S.B.W., and A. Busck. 1920. Notes and new species. Contributions to the natural history of the Lepidoptera of North America. 4(3):230.
Crawford, C.S., and R.F. Harwood. 1959. Lepidoptera associated with grasses grown for seed in Eastern Washington. J. Econ. Entomol. 52:966-969.
268 THE PAN-PACIFIC ENTOMOLOGIST
Crawford, C.S., and R.F. Harwood. 1964. Bionomics and control of insects affecting Washington grass seed fields. Wash. Agric. Exper. Sta. Tech. Bull. 44:25 pp.
Dupnik, T.D., and J.A. Kamm. 1970. Development of an artificial diet for Crambus trisectus. J. Econ. Entomol. 63:1578-1581.
Howell, J.F. 1972. An improved sex attractant trap for codling moth. J. Econ. Entomol. 65:609-611.
Hummel, H.E., L.K. Gaston, H.H. Shorey, R.S. Kaae, K.J. Byrne, and R.M. Silverstein. 1973. Clarification of the chemical status of the pink bollworm sex pheromone. Science (Wash., D.C.) 181:873-875.
Oetting, R.D. 1976. Bluegrass seed fields, Chionodes psiloptera control; SveSHINgIon; 1975. Ins. and Acar. Tests 1:109-110.
Reese, J.C., L.M. English, T.R. Yonke, and M.L. Fairchild. 1972. A method for rearing black cutworms. J. Econ. Entomol. 65:1047-1049.
SCIENTIFIC NOTE
Report of anew host Species for the Long-Tailed Sawfly, Sirex longicauda (Middlekauff, 1948.) (Hymenoptera: Siricidae) — On 23 August 1977 a newly emerged adult female Long-tailed Sawfly, Sirex longicauda (Middlekauff, 1948.) was collected alive from a 7.6 cm cubed sample of Incense Cedar, Libocedrus decurrens Torr.. This wood sample was collected at the sawmill of California Mills, Inc. at Pioneer, Amador County, California from lumber milled from trees originally felled in the Stanislaus National Forest.
Individuals of this species had been previously reported only from White Fir, Abies concolor (Gord. & Glend.) Lindl. (W.W. Middlekauff, 1948; Pan-Pacific Entomol. 24(4):189- 190) and Douglas Fir, Pseudotsuga Menziesii (Mirb.) Franco (W.W. Middlekauff, 1960; Bull. of Cal. Insect Surv. 6(4)).
This is the first report of S. jongicauda from Incense Cedar (W.W. Middlekauff, personal communication), however, the Known host species A. concolor (300-8200 ft. elevation) and P. Menziesii (3000-5000 ft. elevation) occur sympatrically with L. decurrens over much of their ranges, including the Stanislaus National Forest. (P.A. Munz, 1959, 1968; A California Flora with Supplement; U. Cal. Press; Berkeley, California, pp. 49, 57, 59.) — S.F. COOK, L.W. TURLINGTON, Department of Biological Sciences, University of the Pacific, Stockton, California 952117.
The Pan-Pacific Entomologist 53:268 October 1977
Records and descriptions of some Mexican species of the genus Phaea Newman (Coleoptera:Cerambycidae)
John A. Chemsak University of California, Berkeley, 94720
The lamiine genus Phaea Newman is represented by about 30 described species occurring from the United States to Colombia. The group attains its greatest development in Mexico and Central Ameri- ca where a majority of the species are found.
Phaea is closely related to the genus Jetraopes Sch8herr and some
Phaea is closely related to the genus Jetraopes Schonherr and some of the species of each greatly resemble each other. Phaea may be characterized by the usually moderate to small size, cylindrical form, usual slight expansion of the elytra toward the apex, com- pletely divided eyes, convex front of the head, usually apically asper- ate scape of the antennae, ventral flying hairs of the antennal segments, usually well-developed thoracic umbone, short legs and appendiculate tarsal claws. The last abdominal sternite of females is inflated, somewhat elongate, and longitudinally, linearly impressed. The bifid tarsal claws of Tetraopes will immediately separate that genus.
This preliminary study is the result of many years of field collecting and accumulating specimens from most available institutions and collections. The basic problem in attempting to present a comprehensive review of the genus at this time is the lack of adequate material. Whereas some species are represented by large series, others are still Known only from the types. For various reasons, only a small portion of the Mexican species are treated at this time. In addition to the descriptions of new species, several of the less common previously described species are characterized and new records listed.
The National Science Foundation, through Grant GB-BM574 and previous grants, is gratefully acknowledged, as well as the authorities of the following institutions and individuals for the loan of speci- mens: American Museum of Natural History, New York; California Academy of Sciences, San Francisco; Canadian National Collection, Ottawa; Essig Museum of Entomology, Berkeley; Field Museum of Natural History, Chicago; Museum of Comparative Zooology, Cambridge; National Museum of Natural History, Washington, D.C.,; Texas A. & M. University, College Station; University of California, Davis; University of Kansas, Lawrence; Utah State University, Logan; and H. F. Howden and G. Nelson. Celeste Green prepared the illustra- tion.
The Pan-Pacific Entomologist 53:269-276 October 1977
270 THE PAN-PACIFIC ENTOMOLOGIST
Phaea biplagiata, new species
Male: Form small, subparallel; integument reddish to orange, femora yellowish, apices of mandibles, eyes, antennae and part of tibiae black, elytra usually with narrow black stripes extending from humeri to about middle. Head with front moderately convex, moderately densely, separately punctate, densely clothed with pale appressed pubescence; impression between antennal tubercles shallow, vertex sparsely punctate; antennae slender, slightly longer than body, scape moderate, apically asperate, densely punctate, third segment subequal in length to fourth, fourth longer than fifth, succeeding segments becoming progressively shorter, segments densely clothed with short, dark, subappressed pubescence, grayish beneath on basal segments, ciliae numerous beneath, becoming sparser toward apex. Pronotum slightly broader than long, sides sinuate; umbone large, rounded, distinctly delimited at anterior half, densely punctate at sides with a long, erect seta arising from each puncture; apical impression shallow, basal impression moderate; sides irregularly punctate, moderately pubescent; prosternum finely pubescent; meso- and metasternum finely punctate, moderately densely, finely pubescent. Elytra more than 2% times as long as broad, slightly expanding at apex; punctures moderately coarse, separated, linearly arranged, becoming obsolete toward apex; short, pale, appressed pubescence dense behind middle, long, erect hairs denser toward base; apices rounded. Legs short, rather slender, moderately pubescent. Abdomen finely punctate, moderately densely pubescent; last sternite elongate, narrowly rounded at apex. Length, 8-9 mm.
Female: Form more robust. Antennae shorter than body. Abdomen with last sternite linearly impressed. Length, 9-11 mm.
Holotype male, allotype (California Academy of Sciences) from Taxco, Guerrero, Mexico, 6000 ft, 24 June, 1937 (M.A. Embury). Paratypes as follows: 1d, Taxco, July, 1959 (Krauss); 1d, El Salto, Durango, Mexico, 9300 ft, 5 June, 1937 (Embury); 12, 38 miles E. Villa Union, Sinaloa, Mexico, 9 August, 1964 (Howden); 12, Apatzingan, Michoacan, Mexico, 1200 ft, 16 August, 1941 (Haag); 16, Santa Lucia, Sinaloa, Mexico, 4000 ft, 4 August, 1964 (Kelton); 12, San Blas, Nayarit, Mexico, 22 March, 1962 (Stange); 16, 6 miles E. Cuernavaca, Morelos, Mexico, 1 September, 1974 (Bohart, Hanson); 1d, 12, 7 miles SW Yautepec, Morelos, Mexico, 3500 ft, 2 July, 1961 (U. Kansas Mex. Exped.); 19, 30 miles NE Tehuantepec, Oaxaca, Mexico, 8 July, 1955 (Guiliani); 19, Isth. of Tehuantepec (Sumichrast); 1d, 12, Tuxtla Gutierrez, Chiapas, Mexico, August, 1959 (Krauss); 1d, 19, Suchiapa, Chiapas, Mexico, 17 July and 18 August, 1957 (Chemsak, Rannells, Hurd, Durham). 1¢', Champerico, Guatemala, 3 August, 1905 (Knab).
The small size, usual presence of the black longitudinal lines ex- tending back from the humeri and dense, pale pubescence over the apical one-half of the elytra separate this species.
Most specimens at hand possess the dark markings of the elytra. These are, however, reduced in some and absent in one. The color varies from bright reddish to orange.
Phaea flavovittata Bates
Phaea flavovittata Bates, 1881: 197; Chemsak, 1967: 79 (lectotype). Small, black, with front of head and broad line down vertex extending longitudinally down pronotum, orange. Antennae black, scape apically asperate, long, erect hairs beneath not abundant. Pronotum with umbone moderately elevated, rounded, distinctly delimited at sides of apical one-half. Scutellum pale. Elytra coarsely, linearly punctate at basal half; pubescence dense, short, depressed with numerous, long, erect hairs inter-
VOL. 53, NO. 4, OCTOBER 1977 271
spersed. Femora pale basally. Underside dark, moderately densely pubescent. Abdomen of female with last segment longitudinally impressed. Length, 6-9 mm. Type locality: Chacoj, Vera Paz, Guatemala.
New records: 2dd, 14 miles N. Tuxtla Gutierrez, Chiapas, 11 July, 1952 (Gilbert and MacNeill).
Phaea latifrons, new species
Male: Form small, parallel; integument reddish, apices of mandibles, eyes, antennae from apex of scape, umbone, basal spots at sides of pronotum, most of underside, apices of front and middle femora and apical one-fourth of elytra black. Head with front flattened, very broad, much broader across eyes than width of pronotum, rather sparsely punctate, very finely, densely pubescent; area between antennal tubercles barely impressed, vertex sparsely punctate; antennae slender, shorter than body, scape flattened, apically asperate, densely punctate, third segment equal to fourth, fourth slightly longer than fifth, remaining segments gradually decreasing in length, segments densely clothed with short, subdepressed pubescence, ciliae long, numerous. Pronotum about as long as broad, sides slightly sinuate; umbone broad, not prominently elevated, sides not dis- tinctly delimited, middle densely punctate, densely clothed with long, erect, black hairs; sides very sparsely punctate except for median lobes; pubescence fine, pale, appressed, moderately dense with long erect black setae moderately interspersed, base at middle thinly fringed; apical impression shallow, basal impression fairly deep; prosternum finely pubescent; meso- and metasternum finely punctate and pubescent. Elytra about 3 times as long as broad, sides parallel; punctures coarse, linearly arranged, separated; pubes- cence long, suberect, arising from punctures, underlying pubescence white, short, sub- depressed, moderately dense; apices rounded. Legs short, femora stout; pubescence fine, moderately dense. Abdomen finely punctate, moderately densely pubescent; last sternite subtruncate at apex. Length, 9mm.
Holotype male (National Museum of Natural History) and 1 male paratype from Puerto Angel, Oaxaca, Mexico, 31 July, 1965 (A.B. Lau).
This species may be readily recognized by the broadened head and low, densely pubescent umbone.
Phaea marthae, new species
Female: Form moderate sized, sides subparallel; integument black, head except man- dibles and eyes, pronotum dorsally except umbone, reddish, elytra reddish over basal two-thirds except for spots on humeri. Head with front mgoderately strongly convex, moderately coarsely, densely punctate, densely clothed with very fine, pale pubescence with longer, erect black setae numerously interspersed; area between antennal tubercles very shallowly impressed, vertex finely, very sparsely punctate; antennae slender, extend- ing to about second abdominal segment, scape dorsally flattened, strongly asperate apically, densely punctate, third segment slightly longer than fourth, fourth longer than fifth, remaining segments gradually decreasing in length, segments from fourth densely clothed with short appressed pubescence which is dark above and grayish below, long, erect setae sparse beneath. Pronotum broader than long, sides sinuate; umbone large, elevated, very sharply. delimited at sides, very finely, densely punctate; long erect hairs dense, brushlike; sides very sparsely, finely punctate, each puncture giving rise to along black seta; apical impression shallow, basal impression deep; prosternum transversely rugulose, sparsely pubescent; meso- and metasternum very finely, densely punctate, finely, densely pubescent. Elytra over 2% times as long as broad, sides shallowly impressed at middle; punctures moderately coarse, sparse, well separated, irregular; pubescence fine, short, pale, with numerous long, erect hairs interspersed, particularly toward base; apices rounded. Legs short, femora finely, densely punctate, finely, densely
272 THE PAN-PACIFIC ENTOMOLOGIST
pubescent. Abdomen finely, very densely punctate, densely pubescent; last sternite linearly impressed. Length, 13 mm.
Holotype female (California Academy of Sciences) from 9 miles N Mazatlan, Sinaloa, Mexico, 25-28 July, 1972 (J. and M.A. Chemsak, A. and M. Michelbacher).
This species may be readily recognized by the coloration, prominent thoracic umbone, and by the sparse punctation of the elytra.
P. marthae is dedicated to Mrs. Martha Michelbacher in recogni- tion of her field assistance.
Phaea maryannae, new species (Fig. 1)
Male: Form moderate sized, subparallel; integument reddish to orange, mandibles, eyes, appendages, underside, spots on pronotum, scutellum, humeral spots and apices of elytra black. Head with front strongly convex, moderately coarsely, shallowly punctate, pubescence very fine, depressed; impression between antennal tubercles very deep, vertex very sparsely punctate; antennae stout, longer than body, scape robust, apically asperate, densely punctate, third segment subequal to fourth, fourth longer than fifth, succeeding segments becoming gradually shorter, segments moderately densely clothed with short appressed pubescence which is black above and grayish beneath, basal segments sparsely ciliate beneath. Pronotum slightly broader than long, sides sinuate; umbone large, rounded, sides not delimited, punctures minute, very sparse; a large black spot present anteriorly on each side of umbone, two small spots usually at sides near base, an irregular spot also present at sides below median swelling; apex shallowly impressed, base deeply impressed; pubescence sparse, short with a few long, erect hairs interspersed; prosternum finely, moderately densely pubescent; mesosternum finely pubescent, intercoxal process almost plane; metasternum strongly convex, densely, minutely punctate, densely clothed with grayish subdepressed pubescence. Elytra more than twice as long as broad, sides slightly impressed at middle then expanding toward apices; humeri with black spots; apical one-fourth with anteriorly rounded black spots; basal punctures fine, sparse, punctures toward middle larger, irregular, well separated, becoming obsolete toward apex; pubescence very fine, yellowish, longer erect hairs numerous over basal one-half, pubescence on apical spots black; apices broadly rounded. Legs short, stout, gray pubescent. Abdomen minutely, densely punctate, mod- erately densely pubescent; last sternite narrowly rounded at apex. Length, 12-13 mm.
Female: Form more robust. Antennae shorter than body. Legs with femora more slender. Abdomen with pygidium exposed, last sternite inflated, linearly impressed at middle. Length, 16-17 mm.
Holotype male (California Academy of Sciences) and 4 male paratypes from 9 miles N. Mazatlan, Sinaloa, Mexico, 25-28 July, 1972 (J. and M.A. Chemsak, A. and M. Michelbacher). Additional paratypes as follows: 1d: 14 miles S. Cuernavaca, Morelos, Mexico, 3 August, 1954 ( U. Kansas Mex. Exped.); 19, Iguala, Guerrero, Mexico, September (Barrett); 12, Mexico, Mexico, 1950.
This species is distinctive by the large, non-delimited, rounded umbone with black spots at the corners and by the apical black spots of the elytra which curve out from the lateral margins back toward the suture.
The type series is relatively constant in coloration except for the occasional lack of the two basal spots of the pronotum.
This red-headed species is dedisated to Mary Ann Chemsak for the obvious reason in addition to her collecting efforts.
VOL. 53, NO. 4, OCTOBER 1977 273
Fig.1. Phaea maryannae, new species.
274 THE PAN-PACIFIC ENTOMOLOGIST Phaea miniata Pascoe
Phaea miniata Pascoe, 1858: 256. Phaea lineola Bates, 1872: 228; Bates, 1881: 198. New synonymy.
Form rather small, elytra expanding apically. Integument reddish except for tips of mandibles, eyes, apical half of antennal scape, antennae and tibiae and tarsi, which are black, elytra often with narrow black stripes extending from humeri to middle. Head of males with a prominent tubercle on front at middle. Antennae slender, about as long as body, scape apically asperate. Pronotum with umbone shallow, not delimited, rounded. Elytra sparsely, sublinearly punctate, punctures well separated; pubescence fine, moderately sparse, short and appressed and long and erect. Abdomen sparsely pubescent, last sternite linearly impressed in females. Length, 7-10 mm.
Type locality: of miniata, Venezuela; /ineo/a, Chontales, Nicaragua.
Known from Mexico to Venezuela.
New records: 7dd, 1499, Simojovel, Chiapas, 18-31 July, 1958 (J.A. Chemsak). This species was collected on fence posts during the day.
The elytra occasionally have black longitudianl stripes. In addition to the specimens described as /ineo/a from Nicaragua, two from Costa Rica are at hand.
The rounded, barely elevated thoracic umbone and prominent tubercle on the front of the head of males make this species distinc- tive.
Phaea saperda Newman
Phaea saperda Newman, 1840:14; Bates, 1881:195. Phoea saperda, Thomson, 1864:121. Lamprocleptes entomologorum Thomson, 1857:65.
Form moderate-sized, elytra expanding apically. Head, except eyes, pronotum, under- side and often elytra for varying widths at base, bright red to orange, antennae, except apex and legs, yellowish. Antennae with scape apically asperate, long erect hairs moderate. Elytra sparsely punctate at base, punctures becoming larger and denser to- ward middle along suture then obsolete at apex; pubescence fine, grayish, appressed, long erect hairs moderately numerous. Abdomen of female with a median line on last sternite. Length, 8-14 mm.
Type locality: of saperda, Mexico; entomologorum, Brasil?
Known from Mexico, Guatemala and Belize.
New records: San Lius Potosi: 16, El Pujal, 18 May, 1950 (M. Sanchez); 19, El Salto, 18 June, 1962 (J.M. Campbell); 12, 20 miles N Tamazunchale, 30 July 1960 (Howden); 10d, Tamazunchale, 21 June, 1962 (Campbell). Veracruz: 1d, 49, Cordoba, 8 July, 1965 (Lau), 12, 22 June, 1965 (Lau), 1 October, 1963 (Lau), 16 May, 1946 (J. and D- Pallister); 12, Catemaco, 8 June, 1965 (Burke, Meyer, Shaffner); 1d, 12, Puente Nacional, 21 June, 1962 (Janzen); 2992, Lake Catemaco, 8-16 August, 1960 (Howden); 1d, Sn. Rafeal Jicaltepec, 22 June, 1896; 19, SE Citlaltepetl, Fortin, 3 July, 1964 (Swan). Oaxaca: 1d, 16 miles N Matias Romero, 25 June, 1961 (Carney); 12, 25 miles N Matias Romero, 4 August, 1970 (Fisher, Sullivan).
The color of the elytra is subject to variation ranging from all-black to almost the basal one-half reddish.
VOL. 53, NO. 4, OCTOBER 1977 275
Phaea semirufa Bates
Phaea semirufa Bates, 1872:228; Bates, 1881:198. Phaea scapularis Bates, 1881:198, Chemsak, 1967:79 (lectotype). New synonymy.
Form small, expanding posteriorly, Head and pronotum yellow except for a broad, black line extending from behind eyes down sides of pronotum, antennae black, segments from fourth or fifth basally pale, elytra usually yellowish basally at sides from humeri to about middle, black band usually angled from sides to scutellum, basal half occasionally all yellow, femora yellow at basal half. Antennae stout, densely ciliate, ciliae much longer beneath. Pronotum longer than broad; umbone broad, sides margined, moderately elevated, sparsely punctate; pubescence long, bristling. Elytra coarsely, contiguously, linearly punctate, punctures arranged in six rows, becoming finer toward apex; pubes- cence long, bristling, with underlying short, appressed pubescence; apices subtruncate. Abdomen with last sternite with a median line in females. Length, 8-9 mm.
Type locality: of semirufa, mexico; scapularis, Zapote, Guatemala. Known only from Mexico and Guatemala.
New records: 1d, 7.3 miles SSW Yautepec, Morelos, 2 July, 1961, 3500 ft. (U. Kansas Exped.); 12, Tierra Colorado, Guerrero, 15 August, 1938 (L.J. Lipovski); 12, 49 miles S. Acayucan, Veracruz, 19 July, 1963 (W. Foster); 16, 16.2 miles N. Puente Nacional, Veracruz, 21 June, 1962 (Janzen); 12, Temescal, Oaxaca, 2 June, 1969 (Janzen); 1d, 4 miles NW Pueblo Nuevo, River Bajada, Chiapas, 30 July, 1965 (G.H. Nelson); 1d, 7 mi SE Chiapa de Corzo, Chiapas, 23 June, 1965 (Burke, Meyer, Schaffner).
The coloration and broad-appearing antennae suggest a lycid model for this species.
Phaea tricolor Bates
Phaea tricolor Bates, 1881: 199; Chemsak, 1967:79 (lectotype).
Form small, subparallel. Integument black, head and pronotum orange, umbone black, elytra yellowish at basal half except for area around black scutellum and spots on humeri, anterior edge of black band usually directed back from suture to Jateral margins. Head a little broader across eyes than pronotum, vertex with numerous, long, erect, black hairs; antennae moderately densely ciliate. Pronotum with umbone low, broad, extending from apex to base, sides not distinctly delimited, sides finely, densely punctate, erect ciliae numerous, middle glabrous, basal margin fringed at middle. Elytra coarsely, densely punctate, punctures contiguous, becoming obsolete at apex; pubescence dense, short and subdepressed and long and erect, longest at base. Underside finely, not densely pubescent. Abdomen with last sternite linearly impressed in females. Length, 9mm.
Type locality: Cordoba.
Known from Mexico and Guatemala. :
New records: 299, 20 miles S. Cuidad Victoria, Tamaulipas, 17 June, 1941 (H. Dybas); 12, Tamazunchale, San Luis Potosi, 4-5 July, 1964 (Fisher, Verity).
The coloration suggests a mimetic association although possible models are not yet known.
Literature Cited
Bates, H.W. 1872. On the longicorn Coleoptera of Chontales, Nicaragua. Trans. Entomol. Soc. London. 1872:163-238.
Bates, H. W. 1881. Biologia Centrali-Americana, Insecta, Coleoptera, Lamiidae, vol. 5, pp. 153-224, pls. xii-xv.
276 THE PAN-PACIFIC ENTOMOLOGIST
Chemsak, J. A. 1967. Lectotype designations of Cerambycidae in the British Museum (Natural History) (Coleoptera). J. Kansas Entomol. Soc. 40:73-81.
Newman, E. 1840. Entomological Notes. Entomolgist. no. 1:1-16.
Pascoe, F. P. 1858. On new genera and species of longicorn Coleoptera. Part Ill. Trans. Entomol. Soc. London., ser. 2, vol. 4, pp. 236-266, illus.
Thomson, J. 1857. Essai monographique sur le groupe des tetraophthalmites, de la famille des cerambycides (longicornes). Arch. Entomol., vol. 1, pp. 45-67.
Thomson, J. 1864. Systema cerambycidarum ou expose de tous les genres compris dans la famille des caambycides et familles limitrophes. M@én. Soc. Roy. Sci. Liege. vol. 19, pp. 1-540.
ZOOLOGICAL NOMENCLATURE
The following Opinions (listed by number) have been published recently by the International Commission on Zoological Nomencla- ture (see Bulletin Zoological Nomenclature Volume 34, part 3, 9 No- vember, 1977). ITZN 59.
Opinion No. 1093 (p. 143) Apis rotundata Fabricius, 1787 (Insecta: Hymenoptera): designation under the plenary powers of a neotype.
Opinion No. 1094 (p. 147) Refusal of request to correct the following on the Official List of Family-Group Names in Zoology:
No. 61 GYROPIDAE Kellogg, 1896 (Insecta: Mallophaga)
No. 139 PYRALIDAE Latreille, 1809 (In- secta: Lepidoptera)
No. 199 EPISEMIDAE Guénée, 1852 (In- secta: Lepidoptera)
No. 207 TRIOPSIDAE Keilhack, 1909 (Crustacea, Phyllopoda)
No. 213 TRETASPINAE Whittington, 1941 (Trilobita)
No. 324 TRINOTONIDAE Eichler, 1941 (In- secta: Mallophaga)
Opinion No. 1096 (p. 155) Psednura longicornis, Sjostedt, 1920 (In- secta: Orthoptera): designation under plenary powers.
Opinion No. 1099 (p. 164) Drosophila mercatorum Patterson & Wheeler, 1942 (Insecta, Diptera) conser- vation of;
Drosophila carinata Grimshaw, 1901, sup- pression of.
The Commission cannot supply separates of Opinions.
Prey, Predatory Behavior, and the Daily Cycle of Holopogon wilcoxi Martin
(Diptera: Asilidae)
Henry A. Hespenheide and Mark A. Rubke
Department of Biology University of California Los Angeles, 90024
The literature on the Asilidae has taken several points of view on robberflies as predators: The ethology of the act of predation (Lavigne and Holland 1969, Dennis et a/. 1975), diurnal activity pattern (Dennis and Lavigne 1975) seasonal and microhabitat selection (Scarbrough 1974), selection of prey by type (Linsley 1960, Powell and Stage 1962), and by size (Lavigne and Holland 1969, Dennis and Lavigne 1975, Hespenheide 1975.82). Studies of other organisms which prey on flying insects and the prey available to them have revealed that there is a strong diurnal periodicity in prey availability (Hespenheide 1975; Lewis and Taylor 1964). Because of the ease of observing predatory behavior in robberflies and the location of a locally large population of Ho/opogon wilcoxi, the present study was undertaken to determine the diurnal pattern of foraging in this species. Other aspects of the behavior of this species were also noted, and an analysis was made of the prey taken. Ho/lopogon wilcoxi perches on the tips of branches and sallies out after small insects which fly nearby. The overall behavior of Holopogon wilcoxi is similar to that recorded for species of Heteropogon (Lavigne 1970:270, Lavigne and Holland 1969) and Holopogon albipilosus (Dennis and Lavigne 1975).
Methods
Holopogon wilcoxi was censused hourly along a standard path in an area of approximately 20 by 35 meters (see description of study area, below). The day was divided into nine one-hour periods between 8 and 17 hrs, and each period was censused on three separate days during the period 25 to 30 May 1975. The total number of Ho/opogon seen was recorded as well as the number with prey. Those with prey were caught, when possible, in a small plastic vial, and then released after the prey was dropped, usually immediately. The prey item was then transferred to alcohol in the field. Because of the relatively small sample sizes, the prey specimens in alcohol were accumulated by two-hour time periods, and no distinction was made among the days of collection. Prey items were subsequently sorted to species and
The Pan-Pacific Entomologist 53:277-285 October 1977
278 THE PAN-PACIFIC ENTOMOLOGIST
AO 200
30
°/o
20 100
10
8 10 12 14 16
Time
Fig. 1. Diurnal changes in numbers of Holopogon wilcoxi and proportion of individuals with prey. Right ordinate and solid line show changes in number of flies, expressed as the sum of three censuses; left ordinate and dotted line, per cent of flies observed with prey; abcissa in hrs.
measured for body length (excluding wings) in the laboratory. Prey length was used as an index of prey size, and size data were analyzed statistically.
An attempt to measure flying insect density and diversity — i.e., insects available as prey to Holopogon — by means of sweep samples proved unsuccessful, as all samples were very small. Many general observations on behavior were also made in the field.
Study Site
The study was conducted along the north and west side of the road through the lower part of Cave Creek Canyon in the Coronado
VOL. 53, NO. 4, OCTOBER 1977 279
National Forest, Cochise County, Arizona, about 2 mi sw of Portal at about 5000 feet elevation. The site was located at the base of the steep southeast-facing wall of the canyon about 40 m from Cave Creek. The study area was relatively open, with Mimosa biuncifera Benth. as the dominant shrubby vegetation, 0.5 to 3 m tall, and with several species of oaks (Quercus spp.) roughly bounding the area. Mimosa was largely leafless because it is seasonally deciduous and was, as aresult, the most common plant used for perches, although exposed oak twigs and those of the few sumacs (Rhus sp.) present were also used.
Results and Discussion
Diurnal Feeding Periodicity: Mid-morning (8 to 10 hours) proved to be the preferred time of day for feeding activity, measured by the number of flies with prey (Figure 1). This was also the time of day when the greatest number of asilids were present in the study area, in turn probably a response to the greatest prey availability. The number of flies feeding is significantly correlated with the number present — the correlation coefficient r = 0.733, with a probability between .05 and .02 that the correlation is due to chance. Other studies (Lavigne and Holland 1969) have pointed out that the density of available prey was reduced when the air temperature was either very cool (early morning) or very hot (midday). Published data for flight periods of a variety of insects (Hespenheide 1975:82 Lewis and Taylor 1964) show insects are more often active during early to mid-morning. The importance of temperature is suggested by the observation that during one particularly cool mid-morning period (not censused) following a late-afternoon rain the preceding day, very few asilids were present in the area, and none of those had captured prey, this in strong contrast with atypical mid-morning period. (Fig. 1).
The diurnal cycle of feeding by Holopogon wilcoxi differs somewhat from that typical of mid-summer species studied by Lavigne and Holland (1969). There is a similar morning peak, but then a mid- instead of late-afternoon rise (Fig. 1). The reason for a mid-afternoon rise is unclear. It was still quite warm at 14 hrs and nearly as warm at 15 hrs; fewasilids were in the area, but nearly 25% had prey. The low density of Holopogon might indicate prey density was still low because of the warm temperatures, but for some reason theasilids were more efficient in foraging during this period, perhaps because of light conditions and the flies’ use of backlighting to detect prey (Lavigne, personal communication, and below).
One might expect arise in late-afternoon foraging activity when the temperature cools to that at which prey activity is once again high. That this was not observed was likely due to winds which developed each afternoon. The winds impaired prey flight activity as well as
280 THE PAN-PACIFIC ENTOMOLOGIST
Fi
Fig. 2. Holopogon wilcoxiin “sleeping” position, about 7:30 hrs.
efficiency of foraging by Holopogon, and few asilids were seen flying in the late afternoon hours. This has been found to be true of other species of asilids as well (Lavigne and Holland 1969). Prey become relatively scarce during windy weather, for only the stronger fliers are able to cope with it (Johnson 1950).
The phenomenon of moving into an area during the day and out again at dusk or earlier, as in the case of Holopogon, has been observed for other asilid species, even when present in dense popula- tions (Lavigne and Holland 1969). In the present study it was not determined where the majority of the asilids passed the night, and the reason for moving out of the area daily therefore remains unclear. Perhaps they move to an area where the sunlight will hit them earliest in the morning and then fly to areas better suited for foraging.
Holopogon wilcoxi always perches in the sun when active. If the sun is blocked by a hand that is even relatively far away, the fly will im- mediately move to another perch. This, too, has been shown for Heteropogon maculinervis and Backomyia (Lavigne 1970:270, 1971). This observation and the pattern of foraging behavior suggest the great importance of temperature on the diurnal cycle of Holopogon. Ambient temperature, combined with heat derived from direct insola- tion, appears to be the most important physical factor of the environ- ment to which the flies react.
Lavigne (1970:270) reports Heteropogon maculinervis pressing their bodies against the longitudinal axis of a branch during cooler, cloudy weather. Similar behavior was observed for Holopogon wilcoxi. Both in the evening and in the early morning the flies could be seen
VOL. 53, NO. 4, OCTOBER 1977 281
Fig. 3. Holopogon wilcoxi in typical foraging position, feeding on prey, probably a psyllid.
assuming various positions in which the head of the fly is pressed down close to the twig. These are believed to be “sleeping” positions of the flies and to be elicited by cool temperatures. The position has the effect of making the fly cryptic, in that the fly appears to be a thorn or an extension of the perch twig with the black abdomen characteristically protruding away from the twig. The crypticity could be beneficial, in that once a fly becomes cool it is essentially im- mobile and is vulnerable to predation. However, no predators of Holopogon were observed.
Prey Selection
Like most other asilid species, Holopogon wilcoxi captures prey on the wing by sallying out from a perch and then returning with the prey to a perch. (Fig. 3). Foraging flights were observed to be between 7-8 cm and a meter in length. The flies would often not capture an intended prey individual, but would almost always return to the same perch. The perches were from about .3 to 3 m above the ground, with most between 1 and 1.5 m. When another insect flies by, even a very large one or a conspecific, the flies turn their entire body to view it, similar to behavior exhibited by Holopogon albipilosus (Dennis and Lavigne 1975). Occasionally a fly was observed to move to another perch or to fly out of the immediate area altogether after along period of inactivity on its original perch. This suggests that flies will move to
282 THE PAN-PACIFIC ENTOMOLOGIST
a different perch or area if prey activity is insufficient in the im- mediate vicinity, an observation also made by Lavigne and Holland (1969).
The types, numbers, and sizes of prey taken are given in Table 1. Because sampling of insects available to Ho/lopogon as prey proved unfeasible, it is impossible to discuss prey selection directly. The predominant insect prey taken were Homoptera of the families Cicadellidae, Psyllidae and Aphididae. Diptera and Hemiptera each also accounted for more than 10% of the total prey items; Hymenoptera, Coleoptera, Thysanoptera and Lepidoptera accounted for less than 5% each. Among published records of prey taken by other North American Asilidae, only species of Backomyia (50 and 58%, Lavigne 1971) and Asi/us mesae (57%, Dennis and Lavigne 1975) show comparable proportions of Homoptera taken as prey. Homoptera, especially aphids, have been shown by some studies to predominate among samples of flying insects, although most studies show Diptera to be the most common order (review in Hespenheide 1975:82). Small Diptera were the second most frequent prey item, both in numbers of individuals and species. Hemiptera were next most common as individuals, but no more common in number of species than Hymenoptera or Coleoptera, which suggests that Hol/o- pogon preyed on locally abundant populations. Of the four orders taken only occasionally, thrips seem not to be recorded as prey items of asilids previous to this study.
That different taxa of insects have different preferred activity periods (Hespenheide 1975:82 and included references) is borne out by a comparison of prey taken by Ho/opogon at different time of the day. The ordinal composition of 76 prey items taken between 8 and 10 hrs is different grom the composition of 45 prey items taken by the flies during the rest of the day (10 to 17 hrs; although not significantly so [chi-square = 5.85, 0.5 > probability of chi-square > 0.1]). This demonstration of hour-to-hour differences in diet of a hunting pre- dator suggests the importance of taking time of day into account in quantitative studies of prey selection (discussed in Hespenheide 1975:82).
Although size has regularly been shown to be an important characteristic of prey to predators (review in Hespenheide 1973), comparison of mean sizes among the orders of prey reveals that Holopogon prefers significantly different sizes of prey from one order compared to those from another, an observation that has been made of other species of asilids, as well as birds (Hespenheide 1975.158). Prey size is almost certainly important, but not in a simple way. Minimum observed prey size was about 0.4 mm for an aphid, and many prey were less than 1 mm long. It is of particular interest to note that maximum prey size in Holopogon wilcoxi seems to be limited to that which still allows the fly to alight on its perch. Anumber of times
Table 1.
Taxon Homoptera Cicadellidae Psyllidae Aphididae Diptera Hemiptera Hymenoptera Coleoptera Thysanoptera
Lepidoptera
Total
‘log (size x 10)
*g, and g, are statistics testing significance (+) of skewness and kurtosis, respectively.
Prey of Holopogon wilcoxi —
Number total (%)
72 (59.5) 24
16
32
18 (14.8) 17 (14.0) 5 ( 4.1) 4 ( 3.3) 4 ( 3.3) 1 ( 0.9)
121
measured
65 23 16
113
species
18 6 7
Size — untransformed
x 1.16 1.09 1.32
se .06 15 .08 .06 15 15 .26 .23 17
.06
2 9,
SC O+F OO Oo oO + +
9, + 0 0 0) 0 0 + 0 0
x
1.02 + .02 0.96 + .03 1.10 + .03 1.03 + .03 1.12 + .04 1.35 + .03 1.18 + .06 1.28 + .05 1.03 + .06 1.34
1.11
— transformed + se g
o oO + oO Oo + + + «©
=
(> ee © © ee o Oo Oo
Significance of
difference in means?
rs a re - - = eo Ri
z aa ++
NN
do ph Ae ERS . & - & i ate 2 ss 7 u a -P -A D He
+ a - ++ + ++ - + + i, Px Hy
34 = significant at .05 level; + + = significant at .01 level; above the diagonal are comparisons of transformed means, below are comparisons of untransformed means.
2161 YHAEOLOO ‘v “ON ‘EG “OA
€8¢
284 THE PAN-PACIFIC ENTOMOLOGIST
an asilid was observed to capture a prey item two or three times its size, only to drop it finally because it could not land with it. The overall range of prey sizes is between 1/10 and 1/2 the length of the fly, similar to that of other species (Lavigne and Holland 1969).
For aerially-foraging birds it has been proposed that choice of prey is based partly on overall ease or difficulty of capture and partly on local prey density (Hespenheide 1975:82), and the same combina- tion has been proposed to explain prey preference in the asilid Stenopogon engelhardti by Powell and Stage (1962). For most organisms, flight speed and agility is an increasing function of body size, so that a predator will almost always have an advantage over a relatively smaller prey. The smaller Homoptera, particularly the aphids, and the Diptera are especially weak fliers and therefore more vulnerable as prey. The high frequency of certain species as prey of Holopogon suggests that differences in availability, expressed as local concentrations of individuals of these species, were important, although these could not be measured effectively for this study. Toughness of the prey integument may be an additional factor. Of the prey items most often caught, the Homopterans and Diptera are both characteristically soft-bodied.
Acknowledgments
Identification of Holopogon wilcoxi was made by Eric Fisher. The UCLA Campus Computing Network provided computational assistance. S.E. Jones and R. Lavigne made helpful comments on the manuscript.
Literature Cited
Dennis, D. S. and R. J. Lavigne. 1975. Comparative behavior of Wyoming robber flies Il (Diptera: Asilidae). Univ. Wyo. Agric. Exp. Sta. Sci. Monogr. 30:i-iv, 1-68.
Dennis, D. S., G. P. Roehrkasse, and R. J. Lavigne. 1975. Prey recognition by Efferia frewingi (Diptera: Asilidae). Ann. Entomol. Soc. Am. 68:404-408.
Hespenheide, H. A. 1973. Ecological inferences from morphological data. Ann. Rev. Ecol. Syst. 4:213-229.
Hespenheide, H.. A. 1975. Prey characteristics and predator niche width. In M. L. Cody and J. M. Diamond, eds., Ecology and Evolution of Communities, pp. 158-180. Cambridge, Mass.: Harvard Univ. Press.
Hespenheide, H. A. 1975. Selective predation by two swifts and a swallow in Central America. Ibis 117:82-99.
Johnson, C. G. 1950. The comparison of suction trap, sticky trap and tow-net for the quantitative sampling of small airborne insects. Ann. Appl. Biol. 37:268-285.
Lavigne, R. 1970. Courtship and predatory behavior of Cyrtopogon auratus and C. glarealis (Diptera: Asilidae). J. Kansas Entomol. Soc. 43:163-171.
Lavigne, R. 1970. Courtship and predation behavior of Heteropogon maculinervis (Diptera: Asilidae). J. Kansas Entomol. Soc. 43:270-273.
Lavigne, R. 1971. Backomyia seminoensis sp. nov. from Wyoming with ethological notes on B. limpidipennis (Wilcox) (Diptera: Asilidae). J. Kansas Entomol. Soc. 44: 337-342.
VOL. 53, NO. 4, OCTOBER 1977 285
Lavigne, R. J. and F. R. Holland. 1969. Comparative behavior of eleven species of Wyoming robber flies (Diptera: Asilidae). Univ. Wyo. Agr. Exp. Sta. Sci. Monogr. 18:1-61.
Lewis, T. and L. R. Taylor. 1964. Diurnal periodicity of flight by insects. Trans. R. Entomol. Soc. London 116:393-476.
Linsley, E. G. 1960. Ethology of, some bee- and wasp-killing robber flies of south- eastern Arizona and Western New Mexico (Diptera: Asilidae). Univ. Calif. Publ. Entomol. 16:357-392.
Powell, J. A. and G. I. Stage. 1962. Prey selection by robberflies of the genus Stenopogon, with particular observations on S. engelhardti Bromley (Diptera: Asilidae). Wasmann J. Biol. 20:139-157.
Scarbrough, A. G. 1974. A faunistic study of Asilidae (Diptera) at three locations in northern Baltimore County, Maryland: Incidence, relative abundance, and seasonal distribution. Proc. Entomol. Soc. Wash. 76:385-396.
SCIENTIFIC NOTE
Collection of a Larva of the Incense Cedar Woodwasp, Syntexis libocedrii (Rohwer, 1915) from Pioneer, Amador County, California. (Hymenoptera: Syntexidae) — Syntexis libocedrii, the monotype of the Family Syntexidae, heretofore has been reported from northern California in the Sierra, Siskiyou and Cascade ranges to southern Oregon (R.B. Benson, 1935; Ann. Mag. Nat. Hist.; 16:535-553; W. W. Middlekauff, 1974; Pan-Pacific Entomol. 50(3):288-290; B. E. Wickman, 1967; Ann. Entomol. Soc. Amer. 60(6):1291-1295; R. L. Wescott, 1971; Pan-Pacific Entomol. 47:310).
On 26 July 1977 a dessicated larva of S. /ibocedrii was recovered from approximately 2 cm into the sapwood of an 83 to 90 cm diameter Incense Cedar log (Libocedrus decurrens Torr.). The log was topmost in a deck of cedar logged from the Stanislaus National Forest in early 1977 and stored at the sawmill of California Mills, Inc. at Pioneer, California. Our tentative identification of the badly dessicated larva was confirmed by Dr. W.W. Middlekauff of the Dept. of Entomology at the University of California, Berkeley, who originally described the larvae of S. /ibocedrii (Middlekauff, 1974; Pan-Pacific Entomol. 50(3):288-290.)
Collection of this larva represents the first recorded occurrence of S. /ibocedrii this far south in California, although Wickman (1967; Ann. Entomol. Soc. Amer. 60(6):1291-1295) conceded that S. /ibocedrii probably occurs throughout the range of L. decurrens (southern Oregon to northern Lower California and east into western Nevada) (P.A. Munz, 1959; A California Flora with Supplement; U. Cal. Press; Berkeley, California, pp. 59). — L.W. TURLINGTON, S.F. COOK, Dept. of Biological Sciences, University of the Pacific, Stockton, California 95217.
The Pan-Pacific Entomologist 53:285 October 1977
New Species of Ephemerella (Ephemerella) from Alberta (Ephemeroptera: Ephemerellidae)'
Richard K. Allen California State University, Los Angeles 90032
A collection of Ephemerelia by Bonnie Moffat, Freshwater Institute, Manitoba, Canada, from central Alberta includes nymphs of an unde- scribed species.
Ephemerella moffatae, new species
Nymph. Length: body 9.0-10.0 mm; caudal filaments 5.5-6.5 mm. General color brown to dark brown. Head dark brown with irregular light brown markings; lateral margins pronotum pale; legs light brown with dark brown markings; femora light brown with basal and subapical dark brown transverse bands; tibiae light brown, dark brown apically; tarsi brown; tarsal claws with 9-10 denticles. Abdominal terga with paired submedian ridges segments 4-9, barely discernible tergum 4 (Fig. 1); terga dark brown; abdominal segment 2 without posterolateral projection, segment 3 with poorly developed projection (Fig. 1); abdominal sterna brown, sterna 1-8 with dark brown sublateral maculae. Caudal filaments brown.
Types. Holotype: mature nymph, Athabasca River, jct. Calling River (Station III), Alberta, Canada, 27-V-75, B. Moffat, in Canadian National Collection, Ottawa, Ontario. Para- topotypes: 3 nymphs, 1 collected 27-V-75 in collection California State University, Los Angeles, 2 collected 3-VI-75, in collection Freshwater Institute, Winnipeg, Manitoba, other data same as holotype.
This species is named in honor of Bonnie Moffat, collector of the type series.
The nymphs of E. moffatae are distinguished from all other western North American Ephemerella by paired dorsal abdominal ridges on segments 4-9 (Fig. 1).
Fig. 1, Ephemerella moffatae, abdomen, dorsal view.
‘The research on which this report is based was supported by National Science Foundation Grant No. BMS 75-17661.
The Pan-Pacific Entomologist 53:286 October 1977
A New Genus and Species of Stiletto-flies from Southwestern North America with Close Affinities to Chilean and Australian Genera
(Diptera: Therevidae: Therevinae)
Michael E. Irwin University of Illinois and Illinois Natural History Survey, Urbana, 61801
The Therevidae of the Americas are poorly known. Most genera and generic groups have discrete and continuous distribution patterns. Melanothereva Malloch known from two species in Chile and Peru, however, has a disjunct species in Mexico. Another case in point is Breviperna Irwin n. gen., mainly from north and central Mexico, which seems to have aclose phylogenetic relationship to the Chilean genus Peralia Malloch and, to a lesser degree, the Australian and New Zealand genus Anabarrhynchus Macquart. | have termed this generic grouping the Anabarrhynchus-group. The description of Breviperna and its two included species is the purpose of this paper. Elucidation of the genera and their phylogenetic relationships is included in another paper (Irwin and Lyneborg, Mss.).
Methods and Procedures
Each specimen has been assigned a unique number to facilitate the association of data. The number appears below the specimen on a separate yellow label bearing the following words in offset print: THEREVIDAE/M. E. IRWIN/SPECIMEN #. Numbers referring to specimens will be found in the text and figures in italics. These num- bers incorporate ecological and label data associated with the specimens into an automated data management system originally designed by Rauch (1970).
Descriptions and drawings follow the format used recently in describing therevid flies (Irwin 1973, 1977; Lyneborg 1972, 1976). Morphological definitions can be found in Lyneborg (1968, 1972), and Irwin (1973, 1976).
Acronyms for specimen depositories are expanded as follows: AMNH (American Museum of Natural History, New York, NY), CIS (California Insect Survey, University of California, Berkeley, CA), CU (Cornell University, Ithaca, NY), KSU (Kansas State University, Manhatten, KS), MEI (M. E. Irwin Collection, Urbana, IL), SWRS (Southwestern Research Station of AMNH, Portal, AZ), UA (University of Arizona, Tucson, AZ), USNM (National Museum of Natural History, Washington, D.C.), UZM (Universitetets Zoologiske Museum, Copenhagen, Denmark).
The curators of the above named depositories are gratefully acknowledged for the loan of specimens. | express my appreciation
The Pan-Pacific Entomologist 53:287-296 October 1977
288 THE PAN-PACIFIC ENTOMOLOGIST
to Drs. W. W. Wirth and L. Knutson, USDA/ARS and to Dr. W. Mathis, Smithsonian Institution for making the primary type of Psilocephala placida Coq. available for study. | also wish to express my gratitude to Lloyd LeMere for help on the layout of the illustrations, and to John K. Bouseman and Marcos Kogan for reviewing the manuscript. To Bonnie, my wife, | offer my heartfelt thanks for encouragement and her enduring patience.
Breviperna, new genus
Derivation of name: brevi = short; perna = femur; Gender: feminine
Type-species: B. placida (Coquillett), herein designated; holotype female in United States National Museum of Natural History, USNM Type no. 10420.
Length excluding antennae 8-13 mm; female slightly larger than male; rather thick- bodied flies. Head: Frons of male (fig. 1) at its narrowest about as wide as anterior ocellus; minimum distance between compound eyes of female at vertex; frons of female (figs. 2, 16) at level of anterior ocellus about twice as wide as ocellar tubercle; head 60-70% as high as wide; facial width at lower corner of compound eyes 60-75% of head height; eyes of uniform facet size, not divided; antennal insertion low, approximately 2/3 distance from vertex to gena; frons at antennal insertion 40-45% as wide as head; gena about 1/3 as deep as width of antennal segment 1; gena slightly flanged and darker tomentose than face; head somewhat prognathus (fig. 3); proboscis extending beyond base of antennae; palps one-segmented, about 60% as long as proboscis, thin and with a sharply incurved tip, sparsely covered with thin hairs; antennae 56-75% as long as head depth; antennal segment 1 70-90% as long as segment 3 (including style), sparsely covered with stiff hairs; dorsobasal surface of segment 3 with a few short, thin hairs, otherwise without pile; style terminal, one- or two-segmented, with a terminal bristle; 20-40 heavy occipital setae per side, in two hemispherical, irregular rows; pile on lower frons moderately thick, on upper frons relatively thin, evenly sparse on male, concentrated on lower frons with two irregular rows of shorter hairs per side on upper frons of female; tomentum fine, powdery, covering entire frons, face, gena and occiput; callosities completely lacking. Thorax: Width 70-80% of length; setae: notopleural (np) usually 4 (from 3-5), supra-alar (sa) always 2, postalar (pa) usually 1 (occasionally 2), dorsocentral (dc) 0 or 1, scutellar (sc usually2 (rarely 1 or 3); mesonotal pile sparse, short, thin, over entire surface except scutellum; mesonotal tomentum of two types: long, scale-like tomentum sparse, semi-appressed and pollinose tomentum dense and evenly covering surface; pile sparse to moderately dense on pleurotergite, anepimeron, anepisternum, prosternum and katepisternum; lacking on meropleurite, episternum and epimeron; pile present in central portion of prosternum; pollinose tomentum covering entire pleural region. Wing: Hyaline; cell M, open or closed; vein r, shallowly ‘“‘S” curved (fig. 4); rz longer than rs. Legs: Anterior surface of coxa 1 with two apical setae; posterior surface of coxae 1 and 2 tomentose, not pilose; femora with strong av setae, weak scattered pv setae; femur 1 short; pile on anterior and dorsal surfaces of femur 3 scale-like, appressed; on ventral surface short, sparse, thin; tibia 1 rather stout, straight, with 3-8 strong anterodorsal (ad), 3-8 usually strong posterodorsal (pd), 4-8 strong posteroventral (pv) setae; longest dorsal setae of tibia 1 1.2-1.4 (males) or 1.0-1.1 (females) times longer than greatest width of tibia 1; tibia 3 with 11-19 ad, 6-12 (males) or 9-21 (females) pv setae; setae on hind tarsus about 1/10 as long as basitarsus 3. Abdomen: Broad, nearly parallel-sided from segments 1 through 3, thereafter abdomen tapers sharply; segments 5-8 telescoped within segments 3-4 in males; female segments not telescoped; dorsum of abdomen flattened, more so in males; tergites 1-4 of males variably fasciate; pile moderately sparse, long to short, shortest and
VOL. 53, NO. 4, OCTOBER 1977 289
Figs. 1-6. Breviperna placida. Figs. 1, 3, male head (4040); Fig. 2, female head (4028). Fig 4. Tip of wing of female (4028). Figs. 5-6. Female terminalia (4028). Fig. 5, ventral view; Fig. 6, dorsal view. c = cercus; fur = furca (sternite 9); su = subanal plate; s8 = sternite 8;t9 = tergite 9;t10 = tergite 10.Scales = 1mm.
290 THE PAN-PACIFIC ENTOMOLOGIST
somewhat appressed dorsomesad, longest and erect anterolaterad; male dorsum densely clothed in silvery tomentum; female dorsum mostly lacking silvery tomentum. Female terminalia: Furca (fig. 6) elongate horseshoe-shaped, ends of horseshoe bridged by a sclerotized bar; subanal plate (fig. 5) with sparse, heavy spines over ventral surface; tergite 10 with nine heavy anterolateral setae and nine posterodorsal pairs of stout spines (figs. 5,6). Male terminalia: Sternite 8 (fig. 13) broadly bilobate; tergite 8 (fig. 14) narrowly con- stricted medially; epandrium (figs. 7, 10) large, covering most of terminalia, 1.4 times wider than long (bisected medially longitudinally and latitudinally), with outer posterior margins bidentate; cerci (fig. 7) slightly longer than paraproct; paraproct (fig. 11) not keeled; intersegmental membrane basad of paraproct (fig. 11) thinly sclerotized, inverted, shallowly bilobate, attached to base of paraproct and to inner tooth of posterior margin of epandrium; gonocoxites (fig. 12) large, solidly fused along inner ventral margin; an elongate, hindwardly directed projection twisted downward, inward and upwards and extending slightly beyond tip of stylus on lateral portion of each gonocoxite; hypandrium lacking or completely fused to gonocoxites, not identifiable; aedeagus (figs. 8, 9, 15) attached to ventral lobes and to basal portion of dorsal gonocoxal process; distal end of dorsal gonocoxal process (fig. 15) free, small very reduced, knob-like; midbasal portion of dorsal gonocoxal process attached by a thickened, wide, non-sclerotized membrane to the dorsal apodeme of the aedeagus; ventral lobes broadly attached by a non-sclerotized membrane to the center of the aedeagus; ventral lobes (fig. 12) fused basally, elongate; length of aedeagus basad of center three times length distad of center; dorsal apodeme as wide as base of phallus; phallus (figs. 8, 9, 15) curved downward, elongate, with an asymetrically twisted tip; pair of lobe-like, semicircular projections attached to basal, lateroventral portions of phallus; ventral apodeme narrow (less than half as wide as dorsal apodeme), elongate (1.5 x as long as doral apodeme), not forked; ring-like remnant of inner dorsal shield (figs. 9, 15) attached to midlateral section of ventral apodeme, circling above and anterior to dorsal apodeme, containing posterior portion of ejaculatory apodeme; ejaculatory apodeme (figs. 8, 9, 15) large, extending well beyond dorsal and ventral apodemes, its anterior end horizontally flattened on top, keeled below.
Diagnosis: Prognathus; compound eyes of male separated by about the width of anterior ocellus; antennae inserted low on head; proboscis reaching beyond base of antennae; mesonotal setae usually np 3 or 4, sa 2, pa 1, dc 0 or 1, sc 2; prosternum with hairs centrally; anterior surface of coxa 1 with two apical setae; posterior surface of coxa 1 tomentose, but not pilose; posterior surface of coxa 2 without pile; male abdominal segments 5-8 telescoped into segments 3-4; sternite 8 of male broadly bilobate; outer posterior margin of epandrium bidentate; gonocoxites fused; hypandrium notidentifiable; phallus with asymetrically twisted tip and with two lobe-like projections at lateroventral surfaces of base; ring-shaped inner dorsal shield attached to midlateral section of ventral apodeme and encompassing posterior portion of greatly enlarged ejaculatory apodeme.
The genus Breviperna is known from southeastern Arizona and the states of Nayarit, Oaxaca and Puebla, Mexico.
From the sparse information available on the two known species occuring in the genus, it seems likely that individual specimens of Breviperna inhabit the deciduous forest environment of mountainous zones. Larvae of B. milleri were collected from coarse sand in a narrow canyon above 2000 m elevation.
Included species are B. placida (Coq.), B. milleri Irwin. Material at hand indicates no undescribed species in genus.
Breviperna is clearly a member of the Anabarrhynchus-group and probably has a sister-group relationship with the genus Peralia Malloch. The male terminalia resemble in certain respects those of the genus Cyclote/us Walker, but | believe this resemblance may be convergent.
VOL. 53, NO. 4, OCTOBER 1977 291
Breviperna placida (Cogq.) 1894. Psilocephala placida Coquillett, J. N. Y. Entomol. Soc. 2:99.
Redescription of female holotype: The specimen is in very good condition; 12.2 mm in length, excluding antennae. Ground color brownish-yellow to black; pile on lateral and ventral surfaces moderately long, not dense, whitish on head and thorax, tan onabdomen, on dorsal surface short appressed and sparse; tomentum moderately dense, silver laterally, golden on mesonotum, golden and black, very sparse on abdomen. Head: Ground color brown. Head width 2.52 mm, head height 1.72 mm, head depth 1.36 mm. Hairs on frons (similar to fig. 2) strong, elongate, black, denser on lower than on upper portion, lacking on face and gena; pile moderately dense, white, thin on ventral half of occiput. Dorsal occipital setae strong, black, in two or three scattered, hemispherical rows, about 70 setae; approximately seven pairs of thin, short black ocellar and postocellar hairs. Tomentum on frons golden brown, on face golden buff, on gena golden brown, on ventral occiput silvery white (golden brown at margin of compound eye) grad- ing to golden brown dorsally, golden brown on ocellar tubercle. Distance between com- pound eyes at vertex 0.28 mm, at anterior ocellus 0.32 mm, at antennae 1.16 mm, at lower edge of compound eyes 1.08 mm. Compound eye depth 0.96 mm, height 1.52 mm. Antennal segments 1 and 2 golden yellow tomentose with stiff, black hairs (inner lateral surface with few hairs); segment 1 barrel-shaped, fat, cylindrical, 0.60 mm long, 0.24 mm wide (less than three times as long as wide); segment 3 rounded pear-shaped, 0.48 mm long, 0.20 mm wide; tomentum light golden yellow basally, darker golden brown medially and apically, with a few short, black hairs on dorsobasal surface. Basal segment of style stout, twice as wide as long and less than 1/2 as long as apical segment; terminal bristle slightly shorter than basal segment of style; entire style 0.13 mm long. Proboscis golden buff with a few golden hairs on theca. Palp pale yellow with moderately sparse, thin, elongate, white hairs. Thorax: Mesonotum dark brown to black, not patterned; length, excluding scutellum, 3.44 mm; widest width 2.68 mm. Ground color of mesonotum dark brown to black. Mesonotum densely covered with dark golden, powdery tomen- tum and sparsely covered with elongate, scale-like, semi-appressed, metallic-golden tomentum that becomes longer and denser posteriorly. Thin, dark brown pile sparse- ly covering mesonotum, shorter anteriorly, averaging 0.24 mm in length. Mesonotal setal pattern: np 4 (right side with 5), sa 2 (left side with 3), pa1, dc 0, sc 2 (left side witha third, smaller seta). Pleural area and coxae densely covered with grayish tomentum;; thin, white, elongate pile densely covering pleurotergite and dorsal half of anepimeron, more sparsely covering ventral half of anepimeron, anepisternum, prosternum and anterior half of katepisternum. Pile lacking on meropleurite, episternum, epimeron and posterior half of katepisternum. Wing (fig. 4): Wing length (humeral crossvein to apex) 8.56 mm; widest width 3.20 mm. Distance from fork r,,, to outrun of r, 2.68 mm; to outrun of rs 2.40 mm; distance between outruns r, and r, 1.32 mm. Hyaline with brown linear band along most wing veins; considerable amount of hyaline area remaining on wing. Veins yellowish- brown. Cell M; widely open. Haltere brownish-yellow, stem 0.76 mm long, knob 0.68 mm long. Legs: Brownish-yellow, becoming darker apically. Setae black; hairs mostly white on femora 2 and 3, some black on femur 1; setae on tibiae short, tan. Setal pattern: femur 1 (f,) [ad 0, anteroventral (av) 7, pd 0, pv 6], femur 2 (f,) [ad 0, av 4, pd 0, pv 4], femur 3 (f,) [ad 0, av9, pd 0, pv 21], tibia 1 (t,) [ad 5, av 0, pd 7, pv 5], tibia 2 (t,) [as 7, av 5, pd 7, pv 6] tibia 3 t, [ad 15, av 12, pd 12, pv 8]. Femur 1 relatively short, 2.32 mm long, about 1/2 as long as hind tarsus. Largest posterodorsal setae on foretarsus longer than width of foretarsus. Tibia 1, 2.32 mm long; tibia 3, 4.12 mm long; basitarsus 1, 1.08 mm long; basitarsus 3, 1.76 long. Abdomen: Ground color dard brown on anterior half of dorsum, yeallow to orange brown elsewhere; fasiciate on posterior margins of tergites 2 and 3. Pile thin, light tan on segment 1 and tergite 2, black elsewhere; elongate laterally, longest anterolaterad, shortest mid-dorsally, sparse but long ventrally. Tomentum elongate, golden, appressed on segment 1-3, very sparse on rest of abdomen; some gray polinose tomentum on segment 1. Female terminalia: As described in generic description.
292 THE PAN-PACIFIC ENTOMOLOGIST
Figs. 7-15. Male terminalia of Breviperna placida (4040). Fig. 7. Terminalia, lateral view. Fig. 8. Aedeagus, lateral view. Fig. 9. Aedeagus, ventral view. Fig. 10. Epandrium, dorsal view. Fig. 11. Epandrium, ventral view. Fig. 12. Gonocoxites, ventral view. Fig. 13. Sternite 8, ventral view. Fig. 14. Tergite 8, dorsal view. Fig. 15. Gonocoxites, with aedeagus, dorsal view. c = cercus; da = dorsal apodeme of aedeagus; dgp = dorsal gonocoxal process; e = epandrium; ea = ejaculatory apodeme; g = gonocoxite (basisty- lus); ids = inner dorsal shield of aedeagus; p = paraproct; ph = phallus; s = stylus (dististylus); va = ventral apodeme of aedeagus; vl = ventral lobe of gonocoxite. Scale = 1mm.
VOL. 53, NO. 4, OCTOBER 1977 293
Female variation: Antennal segment 1, 2/3 to 3/4 as wide as long; wing cell M, open or closed, if open distance between outruns of m, and m, 0.13 to 0.35 mm, if closed length of coalesced m,,, 0.00 to 0.05 mm. Most veins with wide band of opaque brown, leaving very little hyaline area. Femoral and tibial setal patterns vary as follows: f, [av 4-10, pv 4-8], f, [ av 2-6, pv 3-9], f; [av 8-10, pv 9-22], t, [ad 4-7, pd 6-8, pv 5-8], t, [ad 4-8, av 3-7, pd 4-7, pv 5-8], t, [ ad 11-19, av 10-14, pd 8-13, pv 8-21 though most specimens had a portion of a second row of pv setae].
Males similar to females except as follows: minimal distance between compound eyes situated at upper frons, from slightly greater to slightly less than width of anterior ocellus; pile on frons (fig. 1) moderately dense, elongate, black; frontal tomentum usually dark brown to brownish-gold, occasionally grayish (specimens 4039, 40417); specimens from the Chiricahua Mountains have a more abruptly raised frons than those from the Santa Rita Mountains. Mesonotal setal pattern: np usually 4 (3-5), sa always 2, pa usually 1(1-2), dc usually 1 (0-1), sc usually 2 (1-2). Wing cell M,; open or closed;if closed length of coalesced m;,,4 from 0.00 to 0.05 mm, if open distance between outruns of m, and m, 0.28 to 0.30 mm; femoral and tibial leg setae vary as follows: f, [av 5-9, pv 4-8],f. [ av 2-5, pv 1- 1-8], fs [av 7-10, pv 4-16], t, [ad 5-9, pd 4-8, pv 4-8], t, [ad 5-7, av 4-5, pd 5-7, pv 4-7], t, [ad 13-19, av 9-13, pd 10-16, pv 6-12 with specimen 4037 possessing a double row totalling 23 ]; wing hyaline with dark brown linear band around vein r,; abdominal segments 2-4 with a few short, black hairs medially; abdominal ground color black dorsally, brownish yellow ventrally; fasciate on posterior margins of tergites 1-2, slightly on 3; pile thin, white; tomentum silvery, dense dorsally, sparse ventrally, very sparse laterally. Male terminalia (figs. 7-15): As depicted under generic description. Ground color orangish- brown with black hairs. Sternite 8 with long, thin, white hairs along posterior margin; tergite 8 with long, mostly white (a few black) hairs along posterolateral margin; epandrium 1.30 mm wide and 0.93 mm long (measured medially longitudinally and latitudinally), with long, thick, black hairs over most of dorsum, the longest being posterolaterad; outermost tooth of posterior margin of epandrium much larger than inner- most, extending to level of apex of cercus; paraproct shallowly bilobate with very thin, brownish-yellow hairs apically, thinly sclerotized; cerci separated for most of their length, united only basally, sclerotized medially and apically with several hairs on dorsal surface, slightly longer than paraproct; gonocoxites large; stylus elongate, bladelike, straight, twisted inward and upward at its tip; basal portion of dorsal gonocoxal process short, squat, with a thickened, inwardly hooked base situated well within gonocoxal capsule. Aedeagus very large; ejaculatory apodeme enlarged, bar-bell shaped, extending from anterior portion of ventral apodeme 0.98 mm anteriorly into abdomen; tip of phallus asym- metrically twisted to the right when viewed from anterodorsal position (fig. 15).
Holotype data: Female, ‘Florida’, H. K. Morrison, collector, according to type notes entered by Coquillett in 1907; a USDA specimen housed in the Smithsonian Institution. Morphologically, this specimen fits well within the limits of the specimens collected from the mountains of Arizona and Mexico, listed below. It seems possible that this specimen originated in Arizona or mountainous Mexico; perhaps it was confiscated by quarantine personnel. The specimen now contains the following labels, from top to bottom: 1) ‘‘Fla.”’, 2) “Type/no. 10420/USNM” [red], 3) “HOLOTYPE/BREVIPERNA/placida/Coq. @/det. M. E. IRWIN, 1977”, 4) ‘‘Psilocephala placida Coq./Type ” [ in Coquillett’s hand printing ] , 5) “USNM”, 6) “THEREVIDAE/M.E. IRWIN/Specimen #4051”’.
Other material examined: 12 males, 12 females. ARIZONA, Santa Cruz Co. ; Santa Rita Mts., Madera Canyon, J. G. Franclemont, 1 d (4025 CU) 1700 m, May 8, 1963; 1 d (4026 CU) and 1 2 (4028 MEI) 1490 m, May 8, 1963; 1 d (4027 MEl) and 1 9 (4029 CU) 1490 m, May 3, 1963; 2 2 (4030 CU, 4037 CU) and 2 d (4042 UZM, 4043 MEI) 1490 m, May 14, 1963; 1 2 (4032 CU) 1490 m, May 9, 1963. Same locality, 2 9 (4033 UA, 4034 UA) May 15, 1960, Laubscher; 1 6 (4041 UA) May 8, 1957, G. Butler & F. Werner; 1 9 (4049 CIS) 1400 m, June 14, 1965, R. D. Sage; 1 2 (4048 MEl) Santa Rita Mts., May 15, 1940, Bryant (Lot 19). Pima Co.; Santa Catalina Mts., 1 2 (4035 UZM) Marshall Gulch, June 20, 1960, F. Werner. Cochise Co., Chiricahua Mts., 1 d (4036 KSU) Cave Creek, 1590 m, May 18, 1961, R. H. & E. M. Painter; Southwestern Research Station of the American Museum of Natural History (N.Y.), 8 km
294 THE PAN-PACIFIC ENTOMOLOGIST
west of Portal, 1650 m, V. Roth, malaise trap, 2 d (4039 SWRS, 4040 MEI) May 28, 1965; 1d (4038 SWRS) June 5, 1965; 1 d (4037 AMNH) June 7, 1965. Gila Co., Globe, 1 2 (4050 USNM), May 14, 1948, F. H. Parker. MEXICO, Nayarit; 15 km NW Santa Isabela, 1d and 19 (mating) (4052 USNM, 4053 USNM), March 10, 1972, F. D. Parker and D. R. Miller.
To date specimens of B. placida have been collected between 1350 and 1700 m above sea level in the mountain ranges of southeastern Arizona and in the Sierra Madre Occidental of Nayarit, Mexico. The species probably occurs throughout the range of the Sierra Madre Occidental and its northern limit is probably southern Arizona.
Breviperna milleri, new species
Female holotype (4055), 9.7 mm in length, excluding antennae. Two other females, paratypes (4054, 4056), 9.3 and 9.1 mm long respectively. Ground color black; pile on lateral surfaces generally whitish-gray and moderately long, that on dorsum short, appres- sed and sparse; tomentum sparse, whitish-gray laterad and ventrad, predominently brown on mesonotum and head, very sparse on dorsum of abdomen. Head: Width 1.92 mm, height 1.20 mm, depth 1.06 mm. Frons not patterned except for a pair of velvety-brown semicircular spots midway between antennae and ocellar tubercle adjacent to compound eyes; silvery-white tomentose area just below each velvety spot (these barely distinguish- able on holotype because face is greasy). Hairs on frons (fig. 16) black, short, sparse on upper frons, rather dense, longer on lower frons, continuing along lateral areas of face; pile white, elongate on face and lower occiput, shorter and black on gena. Dorsal occi- pital setae thick, black, in two uneven hemispherical rows, about 40 setae; approximately seven pairs of thin, short, black ocellar and post-ocellar hairs. Tomentum on frons sparse, brown, on face whitish, on gena black, on occiput sparse and whitish, on ocellar tubercle brown. Distance between compound eyes at vertex 0.40 mm, at anterior ocellus 0.46 mm, at antennae 1.04 mm, at lower edge of compound eye 1.20 mm. Compound eye depth 0.66 mm, height 1.08 mm. Antennal segments 1 and 2 brown, thinly whitish and golden tomentose with stiff black hairs (inner lateral surface of segment 1 without hairs); segment 1 elongate, thin, curved outward in center, 0.44 mm long, 0.14 mm wide (more than three times as long as wide); segment 3 elongate pear-shaped, 0.38 mm long, 0.14 mm wide; tomentum dark brown basally, more golden apically, with several short, black hairs on dorsobasal surface. Style apparently one segmented with an elongate terminal bristle. Proboscis and palp black with sparse brown tomentum. Thorax: Mesonotum patterned, brown tomentose with a pair of silvery-gray tomentose stripes and a pair of silvery-gray patches covering area of sa and pa setae, also covering scutellum; mesonotum sparsely covered with metalic golden, appressed, scale-like tomentum and sparse, black pile. Mesonotal setal pattern: np 3, sa 2, pa 1, dc 0, sc 2. Pleural area and coxae densely covered with grayish tomentum; thin, white, elongate pile densely cover- ing same areas as on B. placida. Wing: Wing length (humeral crossvein to apex) 5.34 mm, widest width 2.30 mm. Distance from fork r,,, to outrun of r, 1.64 mm, to outrun of r, 1.56 mm; distance between outruns r, and r, 0.76 mm. Hyaline with light brown patch near outrun of r,. Veins medium brown. Cell M,; closed with a coalesced m,,, 0.12 mm long. Haltere dark brown, stem 0.52 mm long, knob 0.54 mm long. Legs: Femora black, tibiae yellowish-brown. Setae black; femora with whitish pile and appressed scale-like whitish hairs; setulae on tibiae short, black. Setal pattern: f, [ad 0, av 1-2, pd 0, pv 0], f, [ad 0, av 0, pd 0, pv 0-1], f; [ad 0, av 4-5, pd 0, pv 5-6], t, [ad 3-5, av 0, pd 3-6, pv 4-6], t, [ad 4-5, av 2-3, pd 4-5, pv 4-5], t, [ad 12-14, av 7-10, pd 7-8, pv 9-11]. Femur 1 relatively short, 0.77-0.81 mm long, about 2/3 as long as femur 3 and about 1/2 as long as hind tarsus. Longest postero- dorsal seta on foretarsus longer than width of foretarsus. Tibia 1, 1.66 mm long; tibia 3,
VOL. 53, NO. 4, OCTOBER 1977 295
HX
ey
yy
Fig. 16. Head of Breviperna milleri, female (4056), frontal view. Scale = 0.5mm.
3.06 mm long; basitarsus 1, 0.74 mm long; basitarsus 3, 1.16 mm long. Abdomen: Ground color black; fasciate on posterior margins of tergites 2 and 3. Pile relatively long, dense on lateral margins of tergite 1, sparse elsewhere, whitish on segments 1 and 2, black else- where. Thin silvery tomentum on segment 1, thinner on 2. A pair of silvery tomentose patches laterally on tergites 4 and 5. Female terminalia: See generic description.
Male: Unknown.
Material examined: 3 females. MEXICO, Puebla, 8 km S. Tecomachalco, 2100 m, August 10, 1967, collected as larva in narrow canyon, M. E. Irwin (with last larval exuvium and pupal exuvium) (holotype female, 4055 MEI; paratype female, 4056 temporarily kept in the collection of MEl). Oaxaca, 17 1/2 km N of Miltipec, March 3, 1972, F. D. Parker and D. R. Miller (paratype female, 4054 USNM).
Derivation of name: Named in honor of Douglass R. Miller, lifelong friend and col- league.
Diagnosis: This species is smaller than B. placida, darker in color and can easily be dis- tinguished by the following combination of characters: frons with a dark velvety brown spot contiguous with each compound eye (fig. 16), lateral portion of face with elongate whitish (and some black) pile, antennal segment 1 much thinner (3 times as long as wide), and
296 THE PAN-PACIFIC ENTOMOLOGIST
antennal style with only one apparent segment plus terminal bristle.
To date specimens of B. milleri have been collected from the states of Puebla and Oaxaca, Mexico, at an elevation of over 2000 m.
Two larvae of B. milleri were sieved from a coarse sandy substrate along the side of a narrow canyon on August 10, 1967. They were reared in the laboratory at the University of California, Riverside, CA on a diet of Tribolium confusum larvae. They pupated on January 14, 1967 (4056) and January 16, 1967 (4055). Dates of eclosion were not recorded, but on the average this subfamily of therevids emerges 10 to 14 days after pupal initiation. One specimen (4055) was kept alive as an adult for several days. She was fed sugar water and maintained in a glass-sided terrarium with a sand substrate. During the time she was alive and active she layed 39 eggs, even though she did not have the opportunity to mate. These eggs are preserved in glycerin in a microvile attached to the pin of specimen 4055.
Literature Cited
Coquillet, D. W. 1894. Revision of the dipterous family Therevidae. /. N. Y. Entomol. Soc. 2(3):97-101.
Irwin, M. E. 1973. A new genus of the Xestomyza-group from the western coast of South Africa, based on two new species with flightless females (Diptera: There- vidae). Ann. Natal Mus. 21(3):533-556.
Irwin, M. E. 1976. Morphology of the terminalia and the known oviposition behaviour in female Therevidae (Diptera: Asiloidea) with an account of correlated adapta- tions and comments on phylogenetic relationships. Ann. Natal Mus. 22(3): 913-935.
Irwin, M. E. 1977. Two new genera and four new species of the Pherocera-group from western North America, with observations on habitats and behavior (Diptera: Therevidae: Phycinae). Proc. Entomol. Soc. Wash. (in press).
Irwin, M. E., and L. Lyneborg. Mss. The genera of Therevidae of America north of Mexico.
Lyneborg, L. 1968. A comparative description of the male terminalia in Thereva Latr., Dialineura Rond., and Psilocephala Zett. (Diptera, Therevidae). Entomo/l. Medd. 36:546-559.
Lyneborg, L. 1972. A revision of the Xestomyza-group of Therevidae (Diptera). Ann. Natal Mus. 21(2):297-376.
Lyneborg, L. 1976. A revision of the Therevine stiletto-flies (Diptera: Therevidae) of the Ethiopian Region. Bull. Br. Mus. nat. Hist. Entomol. Suppl. 26, 157 p.
Rauch, P. A. 1970. Electronic data processing for entomological museums, an economical! approach to an expensive problem. Ph.D. Dissertation in Entomology, Univ. of Calif., Riverside, 78 p.
Exodontiellini, a New Tribe of Opiinae with Exodont Mandibles
(Hymenoptera: Braconidae)
Robert Wharton
Somerset, California, 95684
In the Ichneumonoidea, the term exodont refers to a type of mandible in which the teeth point in an outwardly direction. In addition, such mandibles are usually apically broadened and com- paratively massive. Unlike the mandibles of the vast majority of the ichneumonoid species, those of the exodont form cannot be used in normal chewing and biting. Griffiths (1964) has discussed the func- tion of these remarkable structures with respect to the Alysiinae.
Previously described exodont ichneumonoids are now placed in 3 different tribes: the Idiogrammatini of the ichneumonid subfamily Tryphoninae; and the Alysiini and Dacnusini of the braconid sub- family Alysiinae. The 8 species of the Idiogrammatini all belong in the genus /diogramma Foerster (Townes, 1969). The Alysiini and Dacnusini contain over 600 and 500 species respectively; and these are currently placed in about 60 genera (Shenefelt, 1974; Fischer, 1975). Vanhornia Crawford, of the Proctotrupoidea, also has exodont mandibles (Crawford, 1909). The exodont condition is thus not unique; and has evolved independently at least three times. A recent examination of material from the Canadian National Collection (CNC), the U. S. National Museum (USNM), and the California Insect Survey has shown that even within the Braconidae, this feature has ap- parently evolved several times.
In material from South America, specimens representing a new genus near Aspilodemon Fischer were found to possess overlapping mandibles on which the spine-like teeth (Fig. 1) were pointed in an outward direction. The specimens otherwise resemble members of the Hormiini; and, unlike the Alysiinae, have well-developed occipital and prepectal carinae. The mandibles, though exodont, are unlike those found in any of the Alysiinae; and are undoubtedly of a different origin.
Several specimens representing an unusual exodont braconid have also been collected from various localities in Western North America. Unlike members of the Alysiinae, these do not have a well-developed median longitudinal sulcus (the posterior-median sulcus) extending from a depression in the ocellar triangle back to the foramen magnum. In addition, the wing venation and facial features are quite different from those of previously described alysiines. Two species, representing a new genus, are described below. Because of their
The Pan-Pacific Entomologist 53:297-303 October 1977
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22
0.25mm
Fig. 1. Left mandible of exodont hormiine. Fig. 2. Left mandible of Exodontiella muesebeckin.sp. Fig.3. Right mandible of Exodontiella sp., male.
overall similarity to some of the smaller opiines, | prefer to place them in the Opiinae rather than the Alysiinae. In either case, this new group should be treated as adistinct tribe.
Terminology and measurements used in the descriptions have been defined elsewhere (Wharton, 1977).
Exodontiellini, new tribe.
Mandibles very broad, exodont, with well-defined teeth. Clypeal margin convex, open- ing between clypeus and mandible absent when mandibles closed; epistomal sulcus narrow, very weakly impressed; paraclypeal pits small, not noticeably impressed. Occipital carina absent; posterior-median sulcus very weak to absent. Prepectal carina absent. Wing venation reduced: fore wing without anal cross-veins; radial cell reduced to nearly absent, radius terminating far from wing tip; postnervellus absent. Abdomen sessile, short, depressed; first tergite heavily sclerotized throughout.
Exodontiella, new genus.
Head transverse; concave posterior-medially; narrowed ventrally. Eyes small, malar space large. Mandibles with 3 well-defined teeth; median teeth touching or slightly over- lapping when mandible closed. Clypeus with outer surface weakly convex; epistomal sulcus narrow but distinct. Subocular suture present. Maxillary and labial palps reduced; apparently 4-5 and 2-3 segmented respectively. Antennae shorter than body; with few
VOL. 53, NO. 4, OCTOBER 1977 299
0.25mm
Figs. 4-5. Frontal view, head of Exodontiellaspp. Fig. 4. &. deserticolan.sp.; Fig. 5. E. sp., male.
(less than 20) segments. Thorax broad, deep. Notauli weakly impressed; sternauli absent. Prescutellar pit narrow, deep. Metanotum with posterior-median plate nearly reaching anterior border. Propodeal carinae completely absent. Posterior (or inner) spur of hind tibia slightly longer than anterior spur; femora short, relatively broad. Fore wing with 3 cubital cells; the second small, triangular. Radial cell extremely narrow, nearly absent; radius strongly upcurved and terminating immediately behind enlarged stigma. Cubital segment forming lower border of second cubital cell distinctly thickened distally. Brachial cell opened distally; third discoideal segment and distal portion of brachius absent. Nervulus postfurcal. Recurrent vein antefurcal. Submediellan cell closed distally by well-developed nervellus. Cubitella stronger than radiella; both nearly absent. Three hamuli. Abdomen moderately depressed. Petiole short, very broad. Suture between tergites 2 and 3 distinctly impressed; tergite 2 and at least part of tergite 3 sculptured. Type: Exodontiella deserticola new species.
Exodontiella deserticola, new species. (Fig. 4, 6, 7)
Head: 1.67 times broader than long; 1.33 times broader than mesonotum. Head in dorsal view wider at eyes than at temples. Eyes small, converging ventrally; roughly twice length of temples dorsally, about half length of temples ventrally. Eyes bare; occiput densely hairy medially, more sparsely so laterally and on temples. Malar space large; about one-third eye height. Ocellar triangle large; distance between posterior ocelli nearly as great as ocellar-ocular line. Face moderately convex; matt; roughly twice wider than high. Clypeus nearly ellipsoidal; approximately 3 times broader than high. Mandibles hairy, about 1.20 times longer than apical width; upper and lower borders slightly diverg- ing; tooth 2 broadly triangular, moderately protruding; teeth slightly overlapping when mandible closed; tooth 1 and 3 broad, separated from tooth 2 by indented borders; tooth 1 extending distad of tooth 3; tooth 3 distinctly larger than tooth 1, and with slightly sinuous outer margin. Antennae 12 segmented, roughly 1.50 times longer than thorax; first flagellomere 1.27 times longer than second; flagellomeres distinctly broader apically; all flagellomeres bearing long subapical hairs. Maxillary palps shorter than head.
Thorax: 1.56 times longer than broad; 1.24 times longer than high. Notauli parallel,
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narrow, weakly impressed; covering about anterior two-thirds of mesonotum. Mesonotal midpit absent. Mesonotum sparsely hairy anteriorly. Prescutellar pit nearly smooth; roughly 5.40 times broader than long. Propodeum and metapleuron densely covered with short, white hairs; hairs on posterior half of propodeum dorsally-directed, others ventrally-directed; propodeum granular. Stigma of fore wing short, very broad, roughly twice longer than wide. Recurrent vein distinctly shorter than first discoideal segment. Second cubital cell petiolate, distinct second radial segment absent. First cubital cross- vein (cuqul) about 1.30 times longer than second (cuqu2); the latter quite weak in speci- men from Palm Springs. Second mediellan segment about 0.65 times length of first; about 1.80 times longer than basella.
Abdomen: petiole 0.73 times as long as apical width, apex distinctly more than twice wider than base; surface matt and densely hairy except at extreme apex. Tergite 2 and basal half to two-thirds of tergite 3 irregularly longitudinally aciculate. Ovipositor roughly two-thirds length of thorax; not, or only slightly extending beyond tip of abdomen.
Color: dark reddith-brown; petiole and tergites 2+3 of holotype orange. Specimen from Palm Springs lighter reddish-brown.
Length: 1.5mm.
Holotype female: USA, California, San Bernardino Co., Helendale, 16/V/1955, W. R. M. Mason, Collector. Deposited in CNC. Additional material: Cal., Riverside Co., 6.7 km. S. Palm Springs, 11/VII/1954 (12). Deposited in USNM. Males unknown.
Diagnosis: differs from the following species in its lighter coloration, less granular abdominal sculpture, shorter first cubital cross-vein, shorter second flagellomere, and longer mandible with shorter, broader tooth 2.
Discussion: The female from Palm Springs appears to belong here despite its slightly lighter coloration. In addition, the first mediellan segment is distinctly shorter than the second. Additional material is needed to determine whether or not differences in the relative lengths of the mediellan segments are significant in this group.
Fig.6. Exodontiella deserticola n. sp. lateral view.
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I Q.5 mm
Fig. 7. Fore and hind wing of Exodontiella deserticola n. sp. from Palm Springs.
Exodontiella muesebecki, new species. (Fig. 2)
Head: 1.61 times broader than long; 1.36 times broader than mesonotum. Head in dorsal view as broad at temples as at eyes. Eyes bare; temple and occipital pubescence as in deserticola. Malar space about one-third eye height. Ocellar triangle moderately large; distance between posterior ocelli about 0.57 times length of ocellar- ocular line. Face moderately convex, with weak, median ridge; matt; roughly twice wider than high. Clypeus more nearly hemispherical; about 2.50 times broader than high. Mandibles hairy, distinctly shorter than apical width; upper and lower borders slightly diverging; tooth 2 narrowly triangular, slightly protruding, the teeth barely touching when mandible closed; tooth 1 angular, separated from tooth 2 by distinct cleft, tooth 1 protrud- ing distinctly distad of tooth 3, the latter very broad, with distinctly sinuate outer margin. Antennae 13 segmented, roughly 1.85 times longer than thorax; first flagellomere equal in length to second; shape and pubescence as in deserticola. Maxillary palps shorter than head.
Thorax: 1.67 times longer than broad; 1.25 times longer than high. Notauli weakly impressed; covering anterior two-thirds of mesonotum; gradually weakening and slightly converging posteriorly. Mesonotal midpit absent. Mesonotum sparsely hairy anteriorly. Prescutellar pit sculptured; roughly 6.50 times wider than long. Propodeum as in deserticola; metapleuron slightly less densely hairy. Stigma of fore wing short, very broad, a little more than twice longer than wide. Recurrent vein much shorter than first discoideal segment. Second cubital cell petiolate as in deserticola, but with cuqul 1.79 times longer than cuqu2. Second mediellan segment about 1.40 times longer than first, about 2.85 times longer than basella.
Abdomen: petiole roughly 0.75 times as long as apical width; about twice as wide at
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= 0.5mm —
Fig.8. Fore wing of Exodontiella sp., male.
apex as at base; sculpture strongly granular, pubescence as in deserticola. Tergite 2 with fine reticulate sculpture, basal three-fourths of tergite 3 granular. Ovipositor as in deserticola.
Color: black;abdominal sternites, tegulae, and coxae dark brown; scape, pedicel, first flagellomere and basal half of second yellow; femora brown, with yellowish apices;
anterior femora noticeably lighter; mandibles, tibiae, and basal 2-3 tarsomeres yellow- brown.
Length: 1.75mm.
Holotype female: Canada, Alberta, Cypress Hills Provincial Park, Elkwater Lake, 14/ VIII/1973, L. Masner, Collector. Deposited in CNC. Males unknown.
This species is named in honor of Mr. C. F. W. Muesebeck in recog- nition of his pioneering work on the Nearctic Braconidae.
Diagnosis: distinguished by the shape of the mandible and the length and color of the basal flagellomeres.
In addition to the above, a single male from northwestern Nevada has also been examined. It is strikingly different in the shape of the head — having a broader face and much smaller eyes (Fig. 5). A second radial segment is also present (Fig. 8); and the outer margin of the third mandibular tooth is denticulate (Fig. 3). Some or all of these may be sexual characteristics, however, and the lack of additional material precludes description at this time.
General discussion: The shape of the head (reduced eyes, face narrowed ventrally), the small paraclypeal pits, the narrow, weakly impressed epistomal sulcus, the lack of a well-developed posterior- median sulcus, the reduced facial pubescence, the short, very broad mandible, and the agathidine-like fore wing venation all serve to distinguish Exodontiella from any of the previously described mem- bers of the Alysiinae. The unusual shape of the mandibles, and the fact that they touch or overlap when closed, indicate a probable independent origin of these structures from the alysiine evolutionary
VOL. 53, NO. 4, OCTOBER 1977 303
line. The discovery of Exodontiella and the other exodont braconids mentioned above will necessitate a more critical definition not only of the Alysiinae, but also of the Opiinae and probably other braconid subfamilies as well.
The presence of exodont mandibles in both Exodontiella and the various members of the Alysiinae is apparently a result of parallel evolution. In the Alysiinae, the primary function of these mandibles is in escape from their hosts’ puparium. It will be interesting to learn if the mandibles in Exodontiella serve a similar function, or whether they are perhaps used somehow in host finding.
Acknowledgements
| wish to thank Drs. Paul Marsh and W. R. M. Mason for the loan of specimens used in this study. | am particularly grateful to Dr. Mason for bringing the hormiines to my attention, for suggesting improve- ments in the original manuscript, and for helpful discussions. | also wish to thank Mr. C. F. W. Muesebeck and Dr. C. van Achterberg for their comments on braconid classification.
Literature Cited
Crawford, J. C. 1909. A new family of parasitic Hymenoptera. Proc. Entomol. Soc. Wash., 11:63-64.
Fischer, M. 1975. Taxonomische untersuchungen an_ neotropischen § Alysiini besonders solchen aus dem British Museum (Nat. Hist.) in London (Hymenoptera, Braconidae, Alysiinae). Stud. Entomol., 18:127-152.
Griffiths, G.C.D. 1964. The Alysiinae (Hym. Braconidae) parasites of the Agromyzidae (Diptera) |. General questions of taxonomy, biology and evolution. Beitr. Entomol., 14:823-914.
Shenefelt, R. D. 1974. Pars 11, Braconidae 7, Alysiinae. /n: van der Vecht, J. and R. D. Shenefelt, eds. Hymenopterorum Catalogus (nova editio). Dr. W. Junk B. V.’s-Gravenhage.
Townes, H. 1969. The genera of Ichneumonidae, part 1. Mem. Amer. Entomol. Inst., 11:1-300.
Wharton, R.A. 1977. New World Aphaereta species (Hymenoptera: Braconidae), with a discussion of Terminology used in the tribe Alysiini. Ann. Entomol. Soc. Am., 70:782-803.
SCIENTIFIC NOTE
Notes on the Host Plants and Distribution of Acanthoscelides pauperculus (LeConte) (Coleoptera: Bruchidae).—The two bruchids that are usually taken most often by sweep- ing in the Sierra Nevada and westward in California and are most abundant in collections from the Pacific States are Acanthoscelides aureolus (Horn) and A. pauperculus (LeConte). The ubiquitous A. aureo/us has a variety of hosts, but the hosts of A. pauperculus have remained unknown although a concerted effort was made to find them (Johnson, 1970, Univ. Calif. Publ. Entomol., 59:1-116). | recently reared A. pauperculus from seeds of a native clover, Trifolium obtusiflorum Hooker, collected at 1100’, 24 mi NE Sanger, Fresno County, CA, on 26 June 1975. This is the first record of a bruchid feeding in the seeds ofa
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New World clover although Kingsolver (1968, Proc. Entomol. Soc. Wash., 70:318-322) reported Acanthoscelides pyrrhomelas (Philippi), a bruchid endemic to the New World, feeding in seeds of the Eurasian species Trifolium pratense Linnaeus in Chile. Zacher (1952, Zeit. angew. Entomol., 33:460-480) reported several species of the Old World genus Bruchidius feeding in Old World clovers.
Acanthoscelides pauperculus is most often collected by sweeping flowers of Achillea sp., Asclepias sp. and Lotus sp. In the foothill woodland plant community where the seeds of T. obtusiflorum were collected, adults of A. paupercu/us were swept in abundance from the flowers of Rorippa nasturtium-aquaticum (Linnaeus), Veronica anagallis-aquatica Linnaeus, Boisduvalia stricta (A. Gray) Greene, Asclepias sp., Lupinus sp., Melilotus sp., the host plant, and from the leaves and fruits of Ribes sp., and Rumex sp. All of the plants except Asclepias were growing either in a small stream or along its banks. Asclepias plants were scattered in an open, grassy field near the stream. Because most adult bruchids feed mostly in the flowers of their larval hosts, it is unusual to find adult bruchids with such catholic tastes as A. pauperculus. Its abundance in flowers of such a wide variety of plants would seem to indicate that it is necessary for the adults to feed prior to mating - and oviposition, again a somewhat unusual trait for bruchids.
Two very close relatives of A. pauperculus, A. inquisitus (Fall) and A. napensis Johnson, will probably be found to infest seeds of species of Trifolium when more rearings from different species are made. The distribution of 7. obtusiflorum and the wider distribution of A. pauperculus suggest that this bruchid feeds on more than one species of clover. Because of its apparent specificity to clover seeds and its abundant numbers, there is a distinct possibility that A. pauperculus could become a pest of economic clover seed crops, especially if some of our native clovers are used as crops in the future.
According to Turner (1959, The Legumes of Texas) and Munz and Keck (1959, A California Flora), the genus Trifolium has about 300 species and is most abundant in north temperate areas but also occurs in Asia, Africa and South America. One center of distri- bution seems to be western North America (Turner, 1959). Of the 49 species of Trifolium listed in Munz and Keck, about two-thirds are considered endemic to California or the Pacific States and their distribution is so limited. About one-third are introduced European or Eurasian species. According to Johnson (1970), A. pauperculus has a distribution from southern California to British Columbia and Idaho. The distribution of this bruchid, then, generally matches that of the native species of the genus Trifolium. Adults of A. pauperculus are most often collected along streams or other moist areas, the habitat of most clovers. A distribution confined to the Pacific States can be explained to some extent on the abundance of its host. Because native species of Trifolium also occur east of the Sierra Nevada, a more plausible explanation for the limited distribution of A. pauperculus are the barriers to their distribution of our high western mountain ranges and arid deserts. A similar pattern of distribution is found in another bruchid, Gibbobruchus mimus (Say). This species is widespread in the central United States where it breeds in the seeds of Cercis canadensis Linnaeus. It has never been reared from seeds of redbud, Cercis occidentalis Torrey ex Gray, from Tulare County, CA, although | have collected many lots of seeds from there. Recently | reared G. mimus from C. occidentalis from the Charleston Mountains, Nevada, and the Grand Canyon, Arizona. Apparently the barriers of mountains and deserts have not allowed this bruchid to reach California although the redbud has successfully traversed these barriers. The bruchids, Acanthoscelides pullus (Fall) and A. aureo/us have a much wider distribution in the West, most likely because their hosts (mostly Astraga/us spp.) occur in both high high mountains and deserts.
The lack of any reported damage to introduced clover seeds by A. pauperculus, A. napensis, and A. inquisitus, and their relatively wide distribution, indicate that they are most likely not introduced species.
| thank M. Johnson for assisting with the field work for this research, J. Rominger for plant identifications, and the Agricultural Research Service, U.S. Department of Agriculture, for partial support under Grant #12-14-100-9970 (33). — CLARENCE DAN JOHNSON, Department of Biological Sciences, Northern Arizona University, Flagstaff, Arizona 860717.
Courtship Behavior of Scaptomyza (Bunostoma) australis Malloch (Diptera: Drosophilidae)
Robert F. Rockwell and Joe Grossfield
Department of Biology The City College of New York N.Y., New York 10031
There is general agreement that courtship in the Drosophilidae involves an exchange of information between the males and females and that during this exchange the female is released from inhibitions by the male (Grossfield, 1972). Differences in opinion do exist, how- ever, concerning the actual intersexual interactions and the stimulus modalities involved in the exchange of information (Grossfield, 1972; Averhoff and Richardson, 1974; Petit and Nouaud, 1975; Bennet-clark et al., 1976, Averhoff and Richardson, 1976; Ehrman and Parsons, 1976). Research to date has centered primarily on the examination of species representing one subgenus (Sophophora) of the genus Drosophila. The members of that subgenus possess a somewhat elaborate courtship pattern and, hence, a complex array of potential information exchange interactions even during the early portion of courtship. It is not surprising then that attempts to pinpoint the stimulus modalities involved in the intersexual information exchange during initiation and early courtship have led to disagreement. This is especially true since the two species involved in that research (Drosophila melanogaster Meigen and D. pseudoobscura Frolova) are both capable of integrating stimuli from different sensory inputs (Grossfield, 1971).
The genus Scaptomyza, which is closely related to Drosophila (Throckmorton, 1975), displays a generally simple courtship pattern when viewed in relation to members of the genus Drosophila (Sturtevant, 1921; Carson et. a/., 1970). In the present work we describe the courtship of one member of this genus, subgenus Bunostoma, (S. australis). We also present evidence on the stimulus modalities involved in intersexual information exchange and demonstrate that the use of a simple system to generate a baseline model can provide valuable insight on the mechanisms and stimulus modalities involved in early components of courtship of the Drosophilidae. The description here is the first detailed account of courtship in the genus Scaptomyza.
Materials and Methods
Scaptomyza (Bunostoma) australis Malloch was collected in August 1974 by aspirating adults from the interstices of fallen, partly rotted
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apples in an orchard 20 km. from Melbourne in Victoria, Australia. Stocks were established and maintained on a cornmeal-agar medium at 20°C on a 12:12 light:dark cycle. In addition to observations on field caught specimens, virgin females and males were collected and stored separately in 8 dram food vials. These individuals were aged for 3 to 15 days before courtship observations were made.
Courtship was observed in food vials, beeswax chambers and plastic cells (18mm diameter by 6mm deep; Cook, 1973) at 20°C. Densities from two to twelve flies were observed at sex ratios of 1:1 and 2:3 (females:males). Neither of these variables altered the court- ship pattern. A total of over 80 flies were observed in 16 observation periods ranging from 30 minutes to 4 hours. In addition, four sets of 3 males with 2 “headless” females were studies (see Grossfield, 1972 for procedural details).
Results
Courtship: The male positions himself behind another individual, with his long body axis in line with that of another fly. This behavior is initiated by the male detecting the movement of another individual and he achieves his oriented position by a series of stereotyped movements termed tracking. The precise pattern of the tracking behavior depends on the relative position and direction of movement of the tracked individual.
Tracking involves a set of coordinated, step-wise two-component body movements by the male. His head, pivoting at the neck, follows the moving individual through an arc. After the head passes through the maximum possible excursion of the arc, delineated by one side of the head contacting the anterior portion of the thorax, the body is realigned so that the head and body are on the same long axis. In some cases, depending on the direction and speed of movement of the tracked individual, there is a pause between the attainment of the maximum excursion and the realignment of the body. The logical inference from such pauses is that the detection of movement is being integrated across the surface of the eye (Gotz and Wenking, 1973). The two-stage tracking, with the aforementioned pauses, is continued, with some lateral displacement of the entire body, until the male is positioned behind the tracked individual with his long body axis in line with that of the other fly. The tracking is sometimes (15%, N = 34) preceded by a slow waving of one or both forelegs.
When the proper orientation is achieved, the male rushes to the rear of the tracked individual. If that individual continues moving the male follows and maintains the same orientation. If the individual stops, without turning, the male mounts and attempts to intromit. (Receptive females do not spread their wings prior to being mounted.) No attempt to mount a moving individual was observed
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and non-receptive females may continue moving for varying periods of time (12 minutes in one case). Nonreceptive females being pursued were observed to depress the rear of the abdomen, extrude the ovipositor plates, or turn and fend (see below). Mounted non- receptive females were observed to depress and to dislodge the male with violent kicking. Males being pursued were observed to turn and fend (see below) and if mounted immediately dislodged the other male with violent kicking.
During copulation, the male’s head is under the female’s wings and the female is still. Usually, his forelegs are on the second tergum his midlegs are on the fourth tergum and his hindlegs are held around the venter. Alternately, the midlegs may be held around the venter and his hindlegs are on her hindlegs. His hindlegs may slip onto the sub- strate and he immediately repositions them. This may be repeated several times. At the end of copulation, the male withdraws and steps off without turning. The female immediately runs away. Twelve copulations were timed to yield a mean duration of 1 minute 58.9 seconds + 5.1 seconds. Males and females were observed to copulate more than once during a 30 minute observation period.
Fending. Males, when face to face, rear up on their midlegs and hindlegs and fend with their forelegs. Males and females display a similar behavior when face to face and may even get the forelegs entangled for several seconds at the tibia-femur joint. Females, when face to face, rear up slightly and display a crab-like motion with the forelegs. In all situations the foreleg movements are slow and exaggerated.
Headless Female Studies: The use.of headless females in court- ship studies has been shown to provide data on intersexual information exchange and attendent stimulus modalities (Grossfield, 1972). Thus, observations of courtship involving headless females as well as observations on the general behavior of such females have been included in this study. Within minutes of decapitation, S. australis females twitch their legs and right themselves. They do not display any spontaneous locomotion nor do they locomote in response to external stimuli. They do respond to tactile stimulation with a slow crablike fending. They respond to directed air currents and chalk dust with preening. All parts of the body are preened in- cluding the ‘head’. Some preening was observed which was apparently spontaneous in origin. Eighteen of 20 headless females were still living after 24 hours and 9 were still living after 48 hours.
In the observation cells with males, headless females again dis- played no locomotion. The males approached these females and often tapped them but no tracking or mounting of these females was observed. The females respond to male contact with fending and preening. This tapping may bear no relationship to courtship in this species in that males were observed to tap numerous non-moving
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objects which they encountered. Further, tapping was never ob- served to precede tracking or successful copulation.
While in the observation cells with headless females, males were observed to track, pursue and attempt to mount each other in response to movement. Demonstration that the males were heterosexually active in this situation was achieved by introducing a normal female. Within minutes, the moving, normal female was tracked pursued and mounted. All of the individuals from the observa- tion cells were placed in food vials after the observation period. After 24 hours, the females were dissected; all of the normal females con- tained sperm but none of the headless females had been insemi- nated. The above observations are all consistent with the postulate that movement (from locomotion) is necessary to elicit male tracking behavior and the subsequent components of courtship.
Discussion and Conclusions
The courtship behavior of Scaptomyza australis is clearly initiated by movement. The ensuing courtship pattern consists of a two-stage tracking process followed by pursuit. When movement ceases, the male mounts and attempts to intromit. The head movement followed by body alignment to the tracked individual is an invariant component of this courtship.
Previous reports of courtship behavior in Scaptomyza include studies of North American and Hawaiian species (Sturtevant, 1921; Spieth, 1966). The North American species (S. (Mesoscaptomyza) adjusta (Loew) and S. (Scaptomyza) graminum (Fallen)) have been described as showing “‘very little courtship” all of which is directed at the female from the rear and includes some wing vibration and scissoring by the male. Mounting in these species occurs after genital contact (Sturtevant, 1921). Most of the Hawaiian species also incorporate some wing movement but, in contrast to the North American species, males mount before making genital contact (Spieth, 1966). If the genus Scaptomyza evolved as an offshoot of the Drosophila in Hawaii as hypothesized by Throckmorton (1975), then S. auStralis would represent a line in the evolution of this genus which has lost wing movement completely in courtship but retains the mounting-preceding-intromission sequence of the Hawaiian stem population. Thus, S. australis represents a lineage whose expressed courtship is the simplest yet known in Scaptomyza.
Relative to members of the genus Drosophila, the courtship pattern of S. australis from initiation through mounting is comparatively simple. That is, none of the complex components so common in Drosophila (i.e. licking, wing displays, and movements involving the proboscis, antennae and foretarsi) occur during this period of court- ship in S. australis. There could be components of Scaptomyza court-
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ship and intersexual information exchange which are not present in Drosophila. These might be related to the more complex genital structure of male Scaptomyza (Throckmorton, 1975). Such structural complexity could provide intersexual information and afford the female post-mounting discrimination capabilities. It is. clear, how- ever, that such capabilities are possible only after mounting occurs. It should be recalled that tracking and pursuit did not always result in mounting in this species. Thus, the period of courtship preceding attempted intromission remains of prime importance for considera- tions of intersexual information exchange. During that period the courtship pattern of S. australis is simpler than that of the Drosophila species used to date to study intersexual information exchange.
Averhoff and Richardson (1974) postulated that in Drosophila melanogaster intersexual information transfer is necessary for male “turn-on” and the initiation of courtship. Further, they suggested that the information is mediated through pheromonal channels. The evidence presented here for a simple drosophilid pattern indicates that movement is a necessary component of the information transfer involved in the initiation of courtship. The most obvious inference is that the stimulus modality involved is vision. However, a peromonal channel can not be immediately ruled out as the stimulus modality for the initiation of courtship since it is possible that pheromones are produced in this species. Movement could enhance the production, emission or direction of diffusion of a pheromone. If a pheromonal channel is involved in the initiation of courtship directed at an indi- vidual then, for males, intrasexual information is as effective as inter- sexual information.
Two lines of evidence are inconsistent with peromonal mediation of male courtship initiation. Tracking behavior, which is elicited only by movement, is immediate, rigidly stereotyped and precise. Given these constraints on the attendent stimulus modality and the small size of the observation cell (which would become rapidly permeated with pheromones), it appears that a visual channel mediates the initiation of courtship and the tracking behavior (cf. Wehrhahn and Poggio, 1976).
The second line of evidence is also consistent with a visual channel mediating the initiation of courtship. A single male was placed in a meticulously cleaned observation cell with a small ball of clean cotton. Undisturbed for 30 minutes, the male walked about the cell and occasionally preened. When the cell was rocked so that the cotton ball moved, the male immediately began stereotyped tracking behavior and, when the appropriate orientation had been achieved, pursuit. When the rocking ceased, so did tracking. This was in- variantly repeatable. No mounting with the cotton ball was observed nor was tracking observed when a Cell containing only a male was rocked.
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It would appear, then, that the initiation of courtship in males is activated by movement and mediated through visual channels in this drosophilid. The visual tracking ability and the involvement of vision in the very early stages of courtship is not surprising in light of the results obtained in measurement of the visual orientation capability and control mechanisms demonstrated in laboratory tests of Diptera (Wehrhahn and Poggio, 1976). This is, however, the first report of this fine a degree of control of visual orientation obtaining during sexual behavior. It must be pointed out that the visual mediation demon- strated here is reflective of sighted flies tested in the light. Whether this species is capable of switching to a different stimulus modality when deprived of vision (heteromodal behavioral plasticity) is not known but is under investigation. Several species of the Drosophilidae are known to possess such capabilities (Grossfield, 1971).
The study of courtship in the Drosophilidae has included analysis of behavioral mechanisms (Connolly and Cook, 1973) as well as com- parative studies with a view towards evolutionary relationships (Spieth, 1966). The examination of courtship of a species in this family with asimple courtship repetoire provides a baseline model for intersexual interaction which may provide a more precise evaluation of the various information transfer mechanisms involved in the more complex courtship patterns of other members of this family.
Acknowledgements
We thank Drs. Patrick Colgan and David Miller for their comments. Financial support was provided by USPHS S507 RR 07132 to RFR and FRAP 10576 and NIH GM21630 to JG.
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Averhoff, W. and R. Richardson. 1974. Pheromonal contol of mating patterns in Drosophila melanogaster. Behav. Genet., 4:207-225.
1976. Reply: Pheromones vs. wings in Drosophila courtship. Behav. Genet., 6:97-98.
Bennet-Clark, H., M. Dow, A. Ewing, A. Manning and F. von Schilcher. 1976. Courtship stimuli in Drosophila melanogaster. Behav. Genet., 6:93-95.
Carson, H., D. Hardy, H. Spieth and W. Stone. 1970. The evolutionary biology of the Hawaiian Drosophilidae In: Essays in Evolution and Genetics in Honor of Theodosius Dobzhansky. M. Hecht and W. Steere (eds.) Appleton-Century Crofts, NY, pp. 437-543.
Connolly, K. and R. Cook. 1973. Rejection responses by female Drosophila melano- gaster: their ontogeny, causality and effects upon the behavior of the court- ing male. Behav., 44:142-166.
Cook, R. 1973. Courtship processing in Drosophila melanogaster |. selection for receptivity to wingless males. Amin. Behav., 21:338-348.
Ehrman, L. and P. Parsons. 1976. The Genetics of Behavior. Sinauer, Sundeland, Mass.
Gotz, K. and H. Wenking. 1973. Visual control of locomotion in the walking fruitfly Drosophila. J. Comp. Physiol., 85:235-266.
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Grossfield, J. 1971. Georgaphic distribution and light-dependent behavior in Drosophila. Proc. Nat. Acad. (US), 68:2669-2673. 1972. Decapitated females as a tool in the analysis of Drosophila behavior. Anim. Benav., 20:243-251.
Petit, C. and D. Nouaud. 1975. Ecological competition and the advantage of the rare type in Drosophila melanogaster. Evol., 29:763-776.
Spieth, H. 1966. Courtship behavior of endemic Hawaiian Drosophila. Univ. Texas Publ., 6615:245-313.
Sturtevant, A. 1921. The North American Species of Drosophila. Carnegie !nst. Publ., Washington. ;
Throckmorton, L. 1975. The phylogeny, ecology and geography of Drosophila. In: Handbook of Genetics (Vol. 3). R.C. King (ed). Plenum Press, NY.
Wehrhahn, C. and T. Poggio. 1976. Real time delayed tracking in flies. Nature 261:43-44.
RECENT LITERATURE
Fundamentals of Entomology and Plant Pathology. L. L. Pyenson. Avi Publishing Co., Westport, CT. 1977. 344 pp. $16.00.
Laboratory Manual for Entomology and Plant Pathology. L.L. Pyenson and H.E. Barke. Avi Publishing Co., Westport, Conn. 1977. 122 pp. Softcover $8.00.
Thesarus of Entomology. R. Foote. Entomological Society of America. College Park, MD. $9.00.
A Catalog of the Mosquitos of the World. Entomological Society of America. College Park, MD. Clothbound. $12.50.
“A new, revised, and expanded catalog with over 3,000 valid species names, distribution, type localities, and location of types. ... cutoff date: Dec. 31, 1973.”
Verhandlungen des Sechsten Internationalen Symposiums uber Entomofaunistik in Mitteleuropa Lunz am See (Osterreich). Edited by H. Malicky. W. Junk, The Hague. 1977. XIV + 375 pp. 90 figs, 18 tables. Paperbound. Dutch Guilders, 95.
312 THE PAN-PACIFIC ENTOMOLOGIST
Acanthoscelides bisignatus 68 compressicornis 70 derifieldi 61 desmanthi 64 mundulus 161 pauperculus 303 pectoralis 72 Allen-Mayfly Records 210 Review of Ephemerella 215 A New Ephemerella 286 Allobrox stephani 219 Alloscopus 241 Ametropus ammophilus 210 Anisolabis maritima 114 Annecke & Prinsloo-A New Cheiloneurus 233 Anthocopa abjecta 52 copelandica 52 elongata 50 enceliae 47 Anomala 34, 129 Aphilanthops hispidus 123 Aporthopleura 32 Aranea 46 Arnaud-Robert Grant Wind 74 Trogonoptera trojana 75
Arnaud & Davies-Flight Activity of
Autographa 77 Ashmeadiella parkinsoniae 193 Autographa californica 77 Baetisca bajkovi 222 Barnard-Skeletal-Muscular
Mechanisms 223
Barr-A New Genus of Cleridae 32 Beal and Seeno-Dermestid Synonym 33
Bezark-A New Asilid 76
Bionomics Acanthoscelides 60, 161, 303 Ametropus 210 Anthocopa 47 Aphilanthops 123 Autographa 77 Baetisca 222 Carmenta 175 Centris 191 Ceratina 233 Cheiloneurus 233 Chionodes 258 Choroterpes 210
INDEX TO VOLUME 53
Clypeadon 144 Coelopa 81 Dermaptera 114 Dianthidium 189 Epicauta 137, 139 Evergestis 113 Formica 250 Hermatobates 87 Holopogon 277 Hydropsyche 169 Hypothyris 104 Idiomelissodes 103 Lucilia 223 Melitoma 191 Meloe 101 Nectopsyche 253 Orgyia 46 Oroperla 126 Phyciodes 167 Pleuropompha 135 Scaphinotus 27 Scaptomyza 305 Sirex 268 Stator 72 Stichopogon 76 Syntexis 285 Tipula 118 Vanessa 257 Blapylis 19 Bolbodimyia bermudezi 98 Book Reviews 73, 180, 187, 214 Bouseman - /diomelissodes Hosts & Distribution 103 Brennus 27 Breviperna milleri 288, 293 Callirhytis cedros 43 cedrosensis 43 protobalanus 45 Carmenta welchelorum 175 Centris totonaca 191 Chandler & Hagen-Notoxus synonymy 230 Cheiloneurus leptulus 233 Chemsak-New Jetranodus 124 Mexican Phaea 269 Cheng-The Seabug Hermatobates 87 Chionodes psiloptera 258 Choroterpes inornata 210 Chrysothamnus 237 Clypeadon 144 Coelopa vanduzeei 81
VOL. 53, NO. 4,
Coleoptera Anthicidae 230 Bruchidae 60, 161, 303 Carabidae 27 Ceramycidae 124, 195, 269 Cleridae 32 Dermestidae 33 Hydrophilidae 142 Meloidae 101, 135 Pselaphidae 219 Scarabaeidae 34, 129 Tenebrionidae 1, 19 Collembola 241 Coniontini 1 Coniontis 1 Corrections 97 Cryptopleurum tenue 142 Culicoides leechi 53 Dailey-Elevation of Loxaulus atrior 145 Dailey and Sprenger-Three New Callirhytis 43 Dermaptera 114 Dermestes medialis 33 tristis 33 Desmanthus 66 Dianthidium macrurum 189 Dickson-Book review 73 Dicranocentrus platensis 248 Diptera Asilidae 76, 277 Calliphoridae 223 Cratopogonidae 53 Coelopidae 81 Drosophilidae 305 Tabanidae 98 Tephritidae 237 Therevidae 287 Tipulidae 118 Doru taeniatum 115 Dos Passos-A Note on Polygonia 179 Doyen-Synonymy in Coniontini 1 Duckworth & Eichlin-A New Clearwing 175 Editorial Notice 146, 232, 236 Edmunds-Baetisca in Wyoming 222 Edwards-Book Review 187 Eleodes aristatus 22 orophilus 24 panamintensis 20 volcanensis 23 wakelandi 19 Ephemerella bartoni 217 moffatae 286 Ephemeroptera 210, 215, 222, 286
OCTOBER 1977 313
Epicauta insignis 137 corvina 139 Euberellia annulipes 144 cincticollis 114 Exodontiella deserticola 298, 299 Exodontiellini 298 Eufriesia lucifera 18 Euplusia anisochlora 8 boharti 10 corusca 10 dressleri 12 eburneocincta 12 fragrocara 13 lucida 13 pallida 15 rufocauda 15 tectora 16 xantha 17 Eusattus 1 Evans-Aphilanthops as a predator 123 Prey Specificity in Clypeadon 144 Evergestis angustalis 113 Fields-A New Megaleuctra 211 Forficula auricularia 116 Formica fusca 250 Giuliani-Migration of Vanessa 257 Gordh-A New Scotolinx 205 Grigarik & Schuster-A New Allobrox 219 Haddock-Stream Velocity vs. Hydropsyche 169 Nectopsyche Caddisflies 253 Hartman and Hynes-Biology of Tipula 118 Hemiptera 87 Hermatobates 87 Hespenheide and Rubke-Biology of Holopogon 277 Heteromurus tenuicornis 245 tetracanthus 243 Heterosarus dakotensis 56 setiger 57 Historical Notes 117 Hoffmanseggia 66 Holopogon wilcoxi 277 Hydropsyche oslari 169 Hymenoptera Anthophcricae 189, 233 Apidae 1 Apoidea 57, 103, 123 Braconidae 297 Cynipidae 43, 145 Encyrtidae 233
314
Eulophidae 205, 237 Formicidae 181 Megachilidae 47, 189, 193 Siricidae 268 Sphecidae 123, 144 Syntexidae 285 Hypothyris enclea 104 Idiomelissodes duplocincta 103 Indoscopus strebeli 248
Irwin-New genus and species of Therevidae 287
Johnson-New Acanthoscelides and Host Records 60 Acanthoscelides 161 Distribution of Acanthoscelides 303
Kavanaugh-Aggregation in Scaphinotus 2°
Kimsey-New Euplusia and Eufriesia 8 Kono-Book review 214 Labia curvicauda 116 Labidura riparia 115 Landa-Ant Nest Morphology 250 Langston and Miller-Distribution of Earwigs 114 Lepidoptera Butterflies 78 Gelechiidae 258 Lymantriidae 46 Noctuidae 77 Nymphalidae 104, 167, 179, 257 Pyralidae 113 Sesiidae 175 Lewis-The Sternidius of Arizona 195 Linsley-Correction 97 Loxaulus brunneus atrior 145 Lucilia sericata 223 Mari Mutt-A//oscopus and its species 241 Mayer and Johansen-Cantharidin from Meloe 101 Megaleuctra sierra 211 Melitoma euglossoides 191 Meloe niger 101 Nectopsyche albida 253 diarina 253 Nissolia schottii 161 Notoxus 230 Odonata 147 Oetting-Biology of Chionodes 258 Orgyia pseudotsuga 46 Oroperla barbara 126 Paracosmia toltecus 116 Parker-Nests of Anthocopa 47 Mexican Bees 189 A New Ashmeadiella 193 Paulson & Garrison-Pacific Coast Odonata 147
THE PAN-PACIFIC ENTOMOLOGIST
Phaea
biplagiata 270
flavovittata 270
latifrons 271
marthae 271
maryannae 272
minuta 274
saperda 274
semirufa 275
tricolor 275 Phyciodes mylitta 167 Pinto-Larvae of Blister Beetles 135 Plecoptera 126, 211 Pleuropompha costata 135 Poinar-Observations on the Kelp fly 81 Polygonia faunus arcticus 179 Potts-Anomala checklist 34
Key to Anomala 129 Powell-Butterflies at Mud Puddle 78 Data for Evergestis 113
Proceedings 74 Quercus cedrosensis 43 Recent literature 7, 42, 55, 174, 229, 311 Rockwell and Grossfield-Courtship of
Scaptomyza 305 scaphinotus
interruptus 27
ventricosus 27
straitopunctatus 27 Scaptomyza australis 305 Scientific Notes 33, 46, 103, 113, 123, 144,
167, 204, 210, 222, 257, 268, 285, 303 Scotolinx
gallicola 206
california 207 Shepard-Immigration of Phyciodes 167 Siegfried et al.-The Adults of Oroperla 126 Sirex longicauda 268 Skalistes
inopinatus 116
vara 116 Smetana-A New Cryptopleurum 142 Somerby-New Eleodes 19 Stator pruininus 72 Sternidius
chemsaki 196
centralis 197
imitans 198
decorus 200
incognitus 201 Stichopogon 76 Tetranodus
tropipennis 124
xanthocollis 125 Tetrastichus
cecidophagus 237
VOL. 53, NO. 4, OCTOBER 1977 315
Tidwell and Philip-A New Bolbodimyia 98 | Wangberg-A New Tetrastichus 237
Timberlake-Two new Heterosarus 56 Wharton-New Tribe Exodontiellini 297 Tipula simplex 118 Wickman-Spider predation on Orgyia 46 Trichoptera 169, 253 Wind, Robert Grant, Obit. 74 74 Trogonoptera trojana 75 Wirth-A new Culicoides from California 53 Tumlinson-Book review 180 Yensen et al-ldaho Ants 181
Turlington and Cook-Host for Sirex 268 Young-Biology of Hypothyris 104
Larva of Syntexis 285 Zoological Nomenclature 31, 86, 192, 203, Vanessa cardui 257 218, 252, 276
July issue mailed November 28, 1977.
Published by the
Pacific Coast Entomological Society
in cooperation with
The California Academy of Sciences
VOLUME FIFTY-THREE 1977
EDITORIAL BOARD
T.D. EICHLIN and A.R. HARDY, Co-Editors C.M. WALBY, Editorial Assistant E.G. LINSLEY HUGH B. LEECH E.S. ROSS H.V. DALY P.H. ARNAUD, JR., Treasurer
PUBLICATION COMMITTEE
1977 1978 1979 J.A. Powell, Chairman T.D. Eichlin R.M. Bohart R.W. Thorp J.D. Pinto J.G. Edwards
San Francisco, California 1977
CONTENTS FOR VOLUME 53
Allen R.K.
New records of mayflies from Californiaand Mexico......... 210 Allen, R.K.
Review of Ephemerella and description of new species....... 215 Allen, R.K.
New species of Ephemerella from Alberta .................. 286
Annecke, D.P. and Prinsloo, G.L. New species of Cheiloneurus parasitic in Ceratina from
HEARS © chess SBP a cheeks oy ncsRyOe" Geb 7. SGLE ays len nema ele wale aha 233 Barnard, D.R.
Skeletal-muscular mechanisms of larval Lucilia sericata ...... 223 Barr, W.F.
New genus of European Cleridae ............... 0c cee eee oe Beal, R.S., Jr. and Seeno, T.N.
Dermestes medialis asynonym of Dermestes tristis ........... 33
ES GN are NN So ee Toh heats ap ick “g Pee A eee i h osoes ey eeaee 73, 180, 187, 214 Bouseman, J.K.
Distribution and host relationship of /diomelissodes duplocincta
RT ODAN SCT Sg cat ele yuh areata ocean eueelin, woatigcx ose Minti Lee ‘soos <a ane oa 8 103 Chandler, D.S. and Hagen, K.S.
New synonymy of North American Notoxus...............08. 230 Chemsak, J.A.
New Neotropical Tillomorphiniin Tetranodus ............... 124 Chemsak, J.A.
Some Mexican species of Pha@a........ eee eee 269 Cheng, L.
The elusive seabug Hermatobat@s ...... 2.2.00. 87 Cook, S.F. and Turlington, L.W.
New host species for long-tailed sawfly, Sirex Jongicauda ..... 268 ORE ON estrecho Leet SMa te an ETD sree Mur ales, ods SR Seg ne eee 5. ated 97 Dailey, D.C.
Elevation of Loxaulus brunneus variety atrior to species Dailey, D.C. and Sprenger, C.M.
Three new gall-inducing Ca/lirhytis from Quercus cedrosensis .. 48 dos Passos, C.F.
Taxonomic note on Polygonia faunuSs arctiCuS...........0005 179 Doyen, J.T.
SS VLTVTTV SLEPT TRUITE We 5 5S etive cen crn hehe a gee ay ae a 1 Duckworth, W.D. and Eichlin, T.D.
New species of clearwing moth from southcentral Texas..... 175 EeLOneINGMICE Ss. 4 cdo 25 ie eRe ee ee OS a he 146, 249 Edmunds, G.F., Jr.
Bacusca oa kovi In-VWVY¥OMING. « 1 6 siak an mele wae oa 4 bills va ene 222 Evans, H.E.
Aphilanthops hispidus aS a predator on bees................. 123
Evans, H.E.
Prey specitict ty Intype adorn oes e vane ee cee bes Se ee 144 Fields, W.C., Jr.
Anew Megaleuctra from California ..... 0... 0... eee eee 211 Giuliani, D.
Notes on the.1973 migration of Vanessa cardui .............. 257 Gordh, G.
A new Species of Scotolinx with notes onthe genus.......... 205 Grigarick, A.A. and Schuster, R.O.
Anewepecios of Alloerox i. sad. Pc le Fe ky ew a ees ale 219
Haddock, J.D. Effect of stream current velocity on habitat preference of PAGO GT STI SALT ela ys be 5 fe RR ee asp ee eens glee 169 Haddock, J.D. Larval morphology, behavior and distribution of two
Nectopsvene Caddistiee «4... 5.c2veG ocx e eke du codename eae 253 Hartman, M.J. and Hynes, C.D. Biology of range crane fly, Tipula simplex... 6... - es 118
Hespenheide, H.A. and Rubke, M.A. Prey, predatory behavior, and daily cycle of Holopogon wilcoxi . 277
ac Rad eyc (iL) (20 oa en awa ee Mae ARE a ele RNa Rag h i MeAar en ecg es 117 Fay eh eoe ial Mg (ee Mh, Me rb oe rma dar poe” Dies fore mngn Nets | Mlk fs Meee eh 312 Irwin, M.E.
A new genus and species of stiletto-flies from southwestern
INP EIAUTTa 5 eae hte he ey OO Wtl sw oan xen OU ALS: we eee Gs ate 287 Johnson, C.D.
Two new species of Acanthoscelides andnewhostrecords.... 60
Johnson, C.D. Ecology and behavior of Acanthoscelides mundulus in seeds of
BS ST OCH ty tay ana etek We. ol kath ahtatieks oleae eh edd Peele 161 Johnson, C.D.
Notes on host plants and distribution of Acanthoscelides
DAU OT CEOS ts aah Gale ate Sale ee a eee eS ey Be dug ha aes ees 5 303
Kavanaugh, D.H. Example of aggregation in the Scaphinotus subgenus Brennus . 27 Kimsey, L.S.
New species of bees in Eup/usia and Eufriesia ............4.. 8 Landa, E.R. 7
Unusual ant nest aE Oy for Formica fusca in western
32,515]. Eee Tee Oo eae Aer ac tO a ne Ae ed ota eS ene 250 Langston, R.L. and Miller, S.E.
Expanded distribution of earwigs in California.............. 114 Lewis, A.E.
The Stennefgs Of ARZONal o.oo. ee eve ee ea) To Rate 195 Mari Mutt, J.A.
Status of A/iocopus and redescriptions of its two species..... 241
Mayer, D.F. and Johansen, C.A. Canthanai tint MACS THOR. os ee aev ve wskes «eee ae ees 101
Gl St re eS tiie id ora dae ee a Hew. ese ee tea neat. reas 232, 236 Oetting, R.D.
Immature stages and biology of Chionodes psiloptera in
DIREC EAS SAS CO CTC ES 4. siete wha FTG ee « we, a ces SY © Wllv ps 258 Parker, F.D.
Nests of Anthocopa enceliae andA. elongata ..............+.. 47 Parker, F.D.
Biological notes on some Mexican bees ................-... 189 Parker, F.D.
A new Ashmeadiella from Arizona Paulson, D.R. and Garrison, R.W.
List and new distributional records of Pacific Coast Odonata . 147 Pinto, J.D.
Descriptions of first instar larvae of three species of
Epicautine blister bettas... ss oy ves del S eb he ee bed 135 Pinto, J.D.
Unusual seasonal occurrence of blister beetles in the
voter: tote a Cfo) 2 aie nen ene eee en Meee SC Oe ecm ne gC, a eta 204
Poinar, G.O., Jr. Observations on the kelp fly, Coelopa vanduzeei in southern SOUT y as ARRON a et ee ew SPRL NOM Oy RRepC Ae | RRO dy werrces 81 Potts, R.W.L.
Anomalinae 2. An annotated checklist of Anomal/a for United States and Canada
ee ee ieee oe, Sls Peet a eee ee 34 Potts, R.W.L.
Anomalinae 3. Key to the species of Anomala of America north
CNT IDL UN, oh arn MRE SED oo weeded ects erecenek ween aie Teer AR 129 Powell, J.A.
Biological and distributional data for Evergestis angustalis .... 113 Proceedings, Pacific Coast Entomological Society ............ 74 Recent Literature. soak ne ee ee cae ek eee ee 7, 42, 55, 174, 229, 311 Rockwell, R.F. and Grossfield, J.
Courtship behavior of Scaptomyza (Bunostoma) australis ...... 305
Shepard, J.H.
Immigration of Phyciodes mylitta to Vancouver Island......... 167 Siegfried, C.A., et al.
Adults of Oroperla barbara Smetana, A.
Anew species of Cryptop/eurum from Central America........ 142 Somerby, R.E.
New species of E/eodes (Blapylis) from western United States.. 19 Tidwell, M.A. and Philip, C.B.
Anew Bolbodimyia from Mexico’s Central Plateau ........... 98 Timberlake, P.H.
Description of two new species of Heterosarus
Turlington, L.W. and Cook, S.F.
Collection of larval Syntexis libocedrii from Amador County,
EEUU 9 <Ptatct ta d eal te? ys Ph cE Ke Sot Leelee ce does Pal gehen Mey settee 285 Wangberg, J.K.
A new Tetrastichus parasitizing tephritid gall-formers on
Ghrvsoeihanesus in ldan@ .ajs45 <3 ks be eee cae Hee ee 237 Wharton, R.
Exodontiellini, anew tribe of Opiinae with exodont mandibles 297 Wickman, B.
Spider predation on early instar larval Douglas fir tussock
PYRITE nn Pee cee hale sleet oe lay ed Oe ee ee hee ee 46 Wirth, W.W.
A new Culicoides biting midge from California............... 53 Yensen, N.P., et al.
CRneckiShe Male anter, .bia hed p< exw le tells wpe ee oe 181 Young, A.M.
Notes on the biology of Hypothyris eucleain Costa Rica ...... 104 Zoological Nomenclature ............ 31, 86, 192, 203, 218, 252, 276
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