Howard E. Evans.
Further Studies on Neotropical Epyrini (Hymenoptera, Bethylidae).
Psyche 72:265-278, 1965.

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PSYCHE
December, 1965 No. 4
FURTHER STUDIES ON NEOTROPICAL EPYRINI
(HYMENOPTERA, BETHYLIDAE)
BY HOWARD E, EVANS
Museum of Comparative Zoology
This paper is a -supplement to two recent papers published in the Bulletin of the Museum of Comparative Zoology.' Although these papers were several years in the making, as so often happens they had scarcely appeared when a considerable amount of new material came into my hands. The present paper is concerned with range extensions, notes on variation, and new species in the genera Rhabdepyris, Bakeri- ella, Calyozina, and Epyris. The abbreviations employed are the same as those used in my two previous papers, but for the sake of ready reference the abbreviations are listed again at the conclusion of this paper.
Rhabdepyris (Rhabdepyris) muesebecki Evans, 1965 This species was ~ecorded from the United States only from two specimens intercepted at quarantine in Texas. However, it is ap-
parently established in Texas, as I took a male while sweeping herba- ceous vegetation at the north edge of the city of Brownsville on June 29, 1965. I also have a female before me from Mera, Ecuador, collected January 26, 1923, by F. X. Williams [coll. Bishop Mus., Honolulu]. This is within the known geographic range of this species, but the specimen is of interest because of its large size (LFW 2.1 mm). It keys out well in my revision if one makes allowance for greater variation in size than previously indicated. 'Research and publication supported by a grant from the National Science Foundation, No. GB-1544. Manuscript received by the editor December 7, 1965.
'1964. A synopsis of the American Bethylidae (Hymenoptera, Aculeata). Bull. Mus. Comp. Zool., 132: 1-222.
1965.
A revision of the genus Rhabdepyris in the Americas (Hymenop- tera, Bethylidae). Bull. Mus. Comp. Zoo]., 133 :67-151.



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Psyche [December
Rhabdepyris (Rhabde~~ris) nigriscapus Evans, I 965 This species was described from a single female from the province of Jujuy, Argentina, unfortunately indicated as a ('d" in the orig- inal description, although stated to be a female two lines later. Mr. Fritz Plaumann has recently sent me a second female, this one from Nova Teutonia, Santa Catarina, Brazil, collected in March, 1965. This specimen agrees closely with the type in color and most struc- tural characters, but it is considerably larger (LFW 2.1 mm) and the front femora measure only 2.65 X as long as their maximum width. The head is also slightly wider (WH 1.02 X LH) and the front narrower (WF 1.02
X HE) ; the ocellar triangle is broad,
OOL only .77 X WOT. Thus this specimen bridges some of the gaps supposed to separate nigriscapus and minutulus. The latter species may still be separable on the basis of its extremely small size, more compact antenme, pale scape, and somewhat more robust front femora; but only the accumulation of much more material of this complex will permit final clarification of specific characters. Rhabdepyris (Rhabdepyris) multilineatus n. sp. This species is of unusual interest because the pronotum lacks a foveolate groove paralleling its posterior margin; actually such a groove is partially developed in one of the two available males, but absent in the other male and in all six females. I used the presence of such a groove as a subgeneric character in my 1965 revision, but evidently it will not hold. Otherwise multilineatus is a reasonably typical member of this subgenus, being related to those species in which the front angle of the ocellar triangle of the female is less than a right angle. In the male the ocellar triangle is somewhat broader, so that the males will run to gracilis in my key (as they also will with respect to head shape). Actually the males are readily separable from those of both gracilis and muesebecki on the basis of the pale and much more compact antennae.
This species is also of special interest since the type series has as- sociated with it the first host data for any member of this genus. Pinned with the type series is an adult and a larva of the dermestid Cryptorhopalum septemignatum Sharp [det. J. M. Kingsolver] ; the word "notes" is written on the label in the handwriting of the col- lector, F. X. Williams.
Dr. Williams writes that he can no longer find these notes, but he remembers collecting some empty cocoons under loose bark along the edge of a forest; he believes that the Rhabdepjv-is adults were reared from larvae which were feeding on



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19651 Evans - Epyrini 267
dermestid larvae breeding in the refuse in the cocoons. Perhaps this should be considered a doubtful host record, since the notes are not extant. However, it seems a very logical association, since dermestids are the hosts of several species of Laelius. As I pointed out in 1965, the morphological gap between Laelius and Rhabdepyris sensu stricto is a very small one.
Holotype. - 9, ECUADOR: Milagso, December 1922 (F. X. Wil- Hams) [Bishop Mus., Honolulu].
Description of female type. -Length 3.1 mm; LFW 2.4 mm. Black; mandibles and antennae rufo-testaceous except flagellum weakly infuscated on upper surface; coxae and femora dark brown, legs otherwise bright rufo-testaceous; wings hyaline, veins and stigma amber. Clypeus sharply, obtusely angulate ; third antennal segment about as long as thick. Head slightly higher than wide, WH .97 X LH ; vertex produced above eye tops by a distance equal to about half the eye height; front of moderate breadth, WF .59 X WH, 1.17 X HE; front angle of ocellar triangle very slightly less than a right angle, OOL 1.25 X WOT. Front wholly alutaceous, with shallow, widely spaced punctures. Pronoturn alutaceous but somewhat shining, strongly punctate, its posterior margin depressed but not paralleled by a groove. Scutellas groove a slightly arcuate band of nearly equal width throughout. Propodeal disc 1.2 X as wide as long, with five strong discal carinae plus strong lateral and sublateral carinae, all nine carinae reaching the posterior margin or nearly so. Front femora 2.5 X as long as their maximum width.
Allotype. - d\ ECUADOR: same data as type [Bishop Mus., Honolulu].
Description of male allotype. - Length 2.8 mn1 ; LFW 1.9 mm. Coloration as described for female. First four antennal segments in a ratio of about 17 :7 :7 :8, segments three and eleven each about 1.2 X as long as thick. WH/LH = 1.0; WF .63 X WH, 1.36 X HE; ocellar triangle rather broad, front angle about a right angle; OOL 1.16 X WOT. Front alutaceous, somewhat shining, punc- tures inconspicuous. Pronoturn and scutellar groove as described for female; psopodeal disc 1.2 X as wide as long, carinate much as in the female; side-pieces of propodeum strongly beaded. Parawes. - 5 $29, I
$, ECUADOR: same data as type except 4
$9 dated December 4, 1922 [Bishop Mu., Honolulu; MCZ]. J'anatwn. -The females show only slight variation in size (LFW 2.2-2.4 mm) and only minor variation in color and sculpture : WH varies from .96 to 1.0 X LH, WF from 1.15 to 1.20 X HE,



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2 68 Psyche [December
OOL from 1.2 to 1.3 X WOT. The male paratype is slightly smaller than the allotype (LFW I .8 mm) and is very similar except that the posterior margin of the pronotum is preceded by a row of shallow foveae.
R habdepyris (Rhabdepyis) grac'ilis Evans, I 965 I described this species from a female from Sacramento, California, and a male from near Durango, Mexico, remarking that the associa- tion seemed probable in spite of the distance between these two lo- calities. I have since seen a male from Danville, Contra Costa co., California (F. X. Williams, Aug. 8, 1948) [CAS] agreeing so closely with the allotype from Durango that I feel confident that my association of the sexes was correct. This specimen is identical to the allotype in size, color, and most standard measurements, but OOL is only 1 .I 5 X WOT. A female collected by F. X. Williams at Danville on May 29,
1952 [CAS] probably also belongs here,
but the groove paralleling the posterior margin of the pronotum is virtually absent and OOL is only 1.10 X WOT; in size, color, and other standard measurements it is similar to the Sacramento type. R habdepyris (Rhabdepyris) nigripilosus ( Ashmead), new combination
Mes'itius nigrip'ilosus Ashmead, 1895, Proc. Calif. Acad. Sci., (2) 5: 539. [Type: ?, MEXICO: NAYARIT: Tepic (CAS)]. Epyris nigripilosus Kieffer, 1914, Das Tierreicli, 41 : 345. - Evans, 1964, Bull. Mus. Comp. 2001. Harvard, 132: 108. Rhabdepyris (Rhabdepyris) huachucae Evans, 1965, Bull. Mus. Comp. 2001. Harvard, 133: 73. [Type: 9, ARIZONA: Ramsey Canyon, Huachuca Mts., March 22, 1956 (Werner & Butler) (MCZ)]. New synonymy Through the courtesy of Mr. Hugh B. Leech, I have recently had an opportunity to study the type of Ashmead's nigripilosus, and I find that Kieffer and I were incorrect in assuming it to be an Epyris, Beyond much question it is a second specimen of the species I de- scribed from southeastern Arizona as huachucae. The type of nigri- piloms differs from that of huachucae in the following particulars: slightly larger (LFW 2.6 mm) ; front femora with only a small brownish blotch toward base, otherwise testaceous, middle femora wholly testaceous; head more nearly circular, WH .98 X LH; front very narrow, WF and HE subequal, the punctures very slightly stronger and more closely spaced ; OOL 1.35 X WOT; propodeal disc somewhat broader, measuring 1.35 X as wide as its median length. This is a relatively large and distinctive species of the sub- genus.




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19651 Evans - Epyrini 269
Rhabdepyris (Trichotepyris) hirticulus new name Rliubd@yris (Tricbot~t.yris) nigropilosus Evans, 1965, Bull. Mus. Comp. 2001. Harvard, 133: 84. Preoccupied by Ashmead, 1895. Although Ashmead called his species nigripilosus (see preceding species), according to the International Code of Zoological Nomen- clature, article 58, compound words differing by only a connecting vowel are to be considered homonyms.
A new name is therefore
proposed for this species, which is known from several females from Panama and Brazil.
R hahdepyris (Trichotefiris) apache Evans, I 965 I described this large and striking species from three females from Arizona and a male from Sonora, Mexico. It now appears that it is widely distributed and locally not uncommon in western Mexico. I collected I I $9 and I
<^ from the tops of young Ambrosia plants, apparently containing honeydew, on the outskirts of Guadalajara, Jalisco, Mexico, July 17-28, 1965. I have also seen I <^ from 8 miles SE of Elota, Sinaloa, Mexico, collected May 19, 1962, by L. A. Stange [UCD] and I from Maria Madre Isl,, Tres Marias, Nayasit, Mexico, collected AIay 22, 1925 (H. H. Keifes) [CAS]. In the original description I noted that the pronotal disc of the male is sharply declivous anteriorly and laterally. In all three of the males listed above, the disc is actually subcarinate anteriorly and along the anterior half of the lateral margins. Also, in some of the females from Guadalajara, one can detect a faint, irregular carina along the anterior margin of the pronotal disc. I have already suggested that Anisepyris and T'richotepyris are closely related by way of the mega- rephalus species group of the latter subgenus. It w-ould appear that
apache provides an almost perfect intermediate between these two taxa. For the present I shall, however, leave it in Rhabdepyris (Trichotepyris), since on the whole it seems to fit best in the mega- cephalus group of that subgenus.
Rhabdepyris (Trichotepyris) fortunatus Evans, 1965 This species was based on a single female from Costa Rica. I have before me two females which resemble the type very closely. one from Bucay, Ecuador, Oct. 4, 1922, and one from Blairmont, British Guiana, Oct. 1923 (both collected by F. X. Williams) [Bishop Mus., Honolulu]. Both are slightly larger than the type (LFW 2.6-2.9 mm) and have the ocellar triangle slightly more re-



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270 Psyche [December
mote from the eye margins (OOL 1.3-1.4 X WOT). The very short third antenna1 segment is characteristic of this species. Rhabdepyris (Chlorepyris) virescens Evans, 1965 This large, well characterized species was known to range through- out tropical South America and into Panama. Apparently it also ranges throughout the remainder of tropical Central America and Mexico, for I took a very typical female of this species while sweep- ing vegetation just behind the beach at Playa Matanchen, near San Bias, Nayarit, Mexico, on August 7, 1965. As might be expected, this specimen is more similar to material from Panama than to the type
(from southern Brazil) ; the apical third of the abdomen is rufous, the femora arc wholly rufous, and the front is relatively nar- row (WF 1.10 X HE). LFW measures 4.0 mm; WH/LH = 1.05; the front femora measure 2.05
X as long as wide.
Bakeriella inconspicua Evans, 1964
On July 6, 1965, I collected two males of this species on banana leaves at Tamazunchale, San Luis Potosi, Mexico, thus extending the known range of the species several hundred miles northwestward. F 7
1 hese males are small
(LFW 2.1-2.3 mm) and agree with males
from Tabasco in having the scape slightly infuscated and the front and thoracic dorsum rather strongly shining. The female of this species is still unknown.
Bakeriella brasiliana Evans, I 964
This species was described from a single female from Minas Gerais, Brazil. I have seen one additional female from a place far distant from the type locality but so similar to the type that its conspecificity can scarcely be questioned.
This specimen was taken on July 14,
1918, at the Limon Plantation, Chagres R., Panama (sweeping around cornfield, H. Dietz & J. Zetck) [USNAI]. This specimen is of the same size as the type and of similar coloration except that the apex of the abdomen is dark brown rather than ferruginous; standard measurements are very much like those of the type except that OOL is 1.7 X WOT, the propodeal disc 1.1 X as wide as long. It is conceivable that brasiliana represents the otherwise un- known female sex of depressa Kieffer or flavicornis Kieffer, though by analogy with olmeca Evans and with subcarinata n. sp. I would expect the females of depressa and flavicornis to have five-toothed



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19651 Evans - Epyrini
271
mandibles and at least some evidence of a median cai-ina on the prono- turn.
Bakeriella subcarinata n. sp.
This is the fourth known species in which the male possesses a vertical carina on the temples and a median carina on the pronotum; it is the second of these four species in which the female is known (or at least tentatively associated with the male). In this case the female lacks a carina on the temples (as in olmeca Evans) but possesses a weak median carina on the pronotum (in contrast to olmeca). The female will run to inca Evans in my 1964 key, but it differs in several respects, most conspicuously in having 5-toothed mandibles and in having the median pronotal carina very weak and not set in a depression. The male runs to olnzeca in my key, but differs in having well formed lateral ridges on the pronotum and in having the third antennal segment only I .3 X as long as thick. klotype. - $2, PERU : Quincemil, Dept. Cuzco, 750 meters, Nov. 5-16, 1962 (L. Pefia) [MCZ].
Description of female allotype. - Length 5.0 mm ; LFW 4.3 mm. Black; mandibles testaceous except black at base, the teeth rufous; antenna pale ferruginous, scape and outer flagellar segments strongly infuscated on upper surface; legs black except tibiae and tarsi dull fei-ruginous; wings lightly tinged with yellowish brown. Mandibles with a broad, oblique apical margin which bears five teeth, the basal three teeth very much smaller than the two large apical teeth. Clypeus very short, broadly subangulate. Eyes weakly hairy; antennal scrobes not margined ; third antennal segment about 1.5 X as long as thick. WH .95
X LH; eyes converging below, WF 1.20 X HE; OOL 1.5 X WOT. Vertex broadly rounded off a considerable distance above eye tops; temples fairly broad but ecarinate. Front shining, weakly alutaceous, with strong punctures which are separated by 1-3 X their own diameters. Pronotal disc somewhat more strongly alutaceous, also with strong punctures; anterior, transverse carina straight; median carina weak, alutaceous, not set in a depression; posterior margin with a polished ridge; sides of disc subcarinate in front; side-pieces rugose, especially below. Mesoscutum polished anteriorly, posterior half alutaceous punctate, slightly depressed ; scutellar pits large, transverse, separated by a thin septum. Propodeal disc 1.15
X 2s wide as long, transversely striate between the three discal carinae, otherwise with only weak sculpturing. Front femora
measuring 2.4 X as long as wide.




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272 Psyche [December
Holotype. - d, PERU : Tingo Maria, Dept. Huanuco, 620 meters, Oct. 5-12, 1964 (C. C. Porter) [MCZ].
Description of male type. - Length 4.0 mm ; LFW 2.6 mm. Black; mandibles testaceous, except basal third black, teeth rufous; scape black except apical fourth, as well as all of following three segments, testaceous, remainder of antennae mostly fuscous ; legs black except tarsi straw-colored, tibiae only partially infuscated; wings lightly tinged with yellowish brown. Mandibles with five sharp teeth; clypeus obtusely angulate; scrobes carinate; eyes weakly hairy. First four antenna1 segments in a ratio of about 20 :8 :g :IO, segment three about 1.3 X as long as thick, segment eleven 1.6 X as long as thick. WH 1.08 X LH; eyes converging strongly below, WF 1.15 X HE; OOL 1.4 X WOT. Temples with a carina which starts at the vertex but extends only to about the middle of the eyes. Front alutaceous, moderately shining, obscurely punctate. Pronotal disc alutaceous, even over the crest of the median carina, which is stronger than in the female; anterior, transverse carina angled backward only very weakly at the midline; sides of disc sub- carinate anteriorly, posterior margin with a polished elevation pre- ceded by a row of small foveae. Mesoscutum strongly alutaceous and slightly depressed on the posterior half; scutellar pits large, separated by a thin septum rnedially. Propodeal disc 1.1 X as wide as long, disc transversely striate between the three discal carinae; posterior angles dentate in front of the strongly developed foveae. Paratypes. - 3 d" 8, PERU : Monson Valley, Tingo Maria, Oct. 10, Nov. 29, 1954 (E. I. Schlinger and E. S. Ross) [CAS]. Variation. - The three male paratypes vary slightly in size (LFW 24-2.8 n11n) but resemble the type very closely in color and standard measurements. In all three specimens the carina on the temples ter- minates near the middle of the eyes; in all three the median carina of the pronoturn is slightly weaker than in the type, although more distinct than in the female allotype.
Bakeriella inca Evans, I 964
This species was described from a single female from Puerto Ber- mudez, Rio Pichis, Peru. I have recently studied I 9 and 2 cj' cj' of this species collected by E. I. Schlinger and E. S. Ross in the Monson Valley, Tingo Maria, Peru, Sept.-Dec. 1954 [CASJ. The female is slightly larger than the type (LFW 3.3 mm) and the head is as wide as high, but there are no other differences worthy of note. The males lack a carina on the temples, have a strong median carina on



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19651 Evans - Epyrini 273
the pronotum which is set in a depression, and have hairy eyes; thus they key out readily in my key. In these males LFW is 2.1-2.6 mm; WH is 1.02-1.05 X LH, 1.25-1.30 X HE, OOL about 1.5 X WOT. There are no important differences in color or sculpturing as compared to the female except that the front is only obscurely punctate and the antenna1 scrobes are carinate (as is usual in males of this genus).
Bakeriella rossi Evans, I 964
The collecting of Schlinger and Ross in the Monson Valley, Tingo Maria, Peru, in September and October 1954 also turned up I $? and 3 c? c? of this species, previously known from 3 c? from Co-
lombia. The female will run to couplet 8 of my key; the 5-toothed mandibles at once separate it from brasil'iana, the very weakly arched, non-angulate transverse pronotal carina from olmeca, the fuscous coxae and femora and much larger size from floridana. The female
is 5.5 mm long, LFW 3.9 mm; the body is black except that the apical fourth of the abdomen is suffused with dull ferruginous. The clypeus is sharply, obtusely angular; the eyes are hairy; the vertex is broadly rounded off far above the eye tops, WH being only .9 X LH ; WF is 1.22 X HE, OOL 1.8 X WOT, the propodeal disc 1.2 X as wide as long and bearing the usual three carinae. Bakeriella depressa Kieff er
Since treating this species in 1964, I have seen one additional male, from Tena, Ecuador, collected February 28, 1923, by F. X. Williams [Bishop Mus., Honolulu]. It is the largest specimen I have seen (LFW 3.0 mm) but presents no unusual structural features. The antennae of this species are consistently more elongate than in flavi- cornis Kieffer, segments three and eleven being I .35-I .5 X as long as thick as compared to 1.1-1.3 X as long as thick in flavicornis. Also, the antennae of most specimens are decidedly dusky apically, while in flavicornis they are wholly light yellowish brown. In addition to the type of the latter species I have seen four topotypic paratypes in the collection of Cornell University.
Calyozina mexicana n. sp.
This remarkable species seems to be closer to the type species, ramicornis Enderlein, from Formosa, than to the two neotropical species I assigned here in 1964. In particular, there are ten long branches on the antennae as opposed to only seven or eight short



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274 Psyche [December
branches in neotropica and amazonica.
This species also differs from
the latter two species in having 5-toothed mandibles, dentate claws, a groove connecting the two scutellar pits, and some unusual sculptur- ing on the sides of the thorax and propodeum. It is possible that
mexicana should not be regarded as congeneric with neotropica and amazonica, but considering the fact that all three species are known from only one specimen (in each case a male), as well as the many problems of generic classification in this section of the Epysini, it seems best to be conservative.
Holotype. - d, MEXICO: 23 mi. S Matias Romero, Oaxaca, Aug. 14, 1963 (F. D. Parker & L. A. Stange) [MCZ] . Description of male type. - Length 6.2 mm; LFW 3.8 mm. Black, head and thoracic dorsum with a faint bluish cast, last two abdominal segments rufous; mandibles rufo-testaceous; scape black, flagellum dull ferruginous, the branches dusky; coxae black, legs otherwise wholly bright rufo-testaceous; wings lightly tinged with brown. Mandibles with a large apical tooth and four small teeth in a slightly oblique series; clypeus broadly angulate, its median carina nearly straight in profile. Antennae strongly pectinate ; segment three con- siderably shorter than two, much wider than long, bearing a rather short process; segments 4-12 each bearing a slender process slightly longer than the segment proper (segments 4 and 12) up to about 3
X as long as the segment proper (segments 7-10) ; segment 13 elongate, clavate; segments 2-5 each bearing long, erect setae, fol- lowing segments with shorter setae but with long setae on the proc- esses, these setae much longer than the width of the processes. Head unusually broad, WH 1.25 X LH; front broad, WF .70 X WH, T .50 X HE; vertex broadly rounded off a short distance above eye tops; ocelli in a broad triangle, WOT slightly exceeding OOL. Front polished, non-alutaceous, with small, widely spaced punctures. Thoracic dorsum slightly alutaceous, duller than front; pronotal disc sloping gradually toward the front and its sides approaching to a rounded anterior margin, the disc not at all prominent antero-lat- el-ally; scutellar pits large, subcircular, separated by slightly less than their own diameters, connected by a very slender but fairly deep groove. Propodeal disc 1.15 X as wide as its median length, with three strong discal casinae between which it is transversely ridged; laterad of the discal casinae the surface is smoother, but there is a low longitudinal ridge between the lateral discal and strong sub- lateral carinae ; propodeal side-pieces polished, with several prominent longitudinal ridges which are continuous with similar ridges on the



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19651 Evans - Epyrini 275
mesopleura and which form (more especially on the mesopleura) two concentric ellipses. Middle and hind tibiae somewhat spinose; claws with a short, erect tooth. Thoracic venter rather flat, the middle and hind coxae rather widely separated on the venter. Epyris cubanus n. sp.
This is the first Epyris known from Cuba, and only the third species reported from the West Indies. It is one of few species of Epyris of metallic coloration.
Holotype. -9, CUBA: Rio Cauto, Oriente Prov., April 1928