Charles C. Porter and Thomas J. O'Neill.
A Revision of the New World genus Chromocryptus (Hymenoptera: Ichneumonidae).
Psyche 92:407-446, 1985.

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A REVISION OF THE
NEW WORLD GENUS CHROMOCRYPTUS
(HYMENOPTERA: ICHNEUMONIDAE).
BY CHARLES C. PORTER' AND THOMAS J. ~'NEILL Department of Biological Sciences,
Fordham University,
Bronx, NY 10458
Taxonomy
Chromocryptus is a strictly New World representative of the cosmopolitan ichneumonid subtribe Ischnina (Gelinae: Meso- stenini), as defined by Townes (1969: 158-9). It may be distinguished from other Nearctic and Neotropic ischnines by the following con- cert of features: (1). Black, red, and white (sometimes mostly red); (2). Wings hyaline to moderately infumate; (3). Clypeus pyramidally raised, often nasute; (4). Mesoscutum largely with coarse punctures or puncto-reticulation and with notauli only faintly defined (fine but well impressed in some males); (5). Hind coxa with a strong subver- tical groove externo-ventrally near base; (6). Discocubitus gently arched to weakly angled, sometimes with a vestigial ramellus; (7). Axillus very close to hind margin of hind wing; (8). Propodeal spiracle 1.3-3.0 as long as wide; (9). Propodeal cristae large and well projecting, obtusely cuneate to subligulate, never spiniform; (10). Base of 1st gastric tergite with a lateral expansion that is low and rounded or sometimes prominently subtriangular; (1 1). Female gas- ter short and stout, short-fusiform to almost ovoid; (12). Second gastric tergite with abundant large punctures that usually are subad- jacent or denser; its setae short but mostly approaching the length of their interspaces; (13). Ovipositor slender to robust, straight, com- pressed; its sheathed portion 0.35-0.45 as long as fore wing. The foregoing diagnosis easily distinguishes Chromocryptus from its Research Associate, Florida State Collection of Arthropods, Florida Department of Agriculture and Consumer Services, P. 0. Box 1269, Gainesville, Fl32602. Manuscript received by the editor May 15, 1985.



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North and Middle American relatives, but must be used with care in considering the complex South American fauna. A complete des- cription is furnished later in this study antecedent to the key for Chromocryptus species.
My concept of Chromocryptus agrees with Townes earlier inter- pretations (1962: 256-7). Some recent authors (Porter 1967: 206- 1 1, Townes 1966: 68) have regarded Chromocryptus as a species group within the immense, almost exclusively South American genus Tra- chysphyrus Haliday. In 1969 (179-81), Townes restricted Chromo- cryptus to those "Trachysphyrus" in which the axillus vein runs close to the posterior margin of the hind wing. This arrangement still leaves under Chromocryptus a heterogeneous and perhaps poly- phyletic complex of more than 45 species. I thus prefer to return to Townes' 1962 perception of the genus and to regard it as comprising only those species intimately related to the North American C. planosae. This viewpoint has been strengthened by discovery of 5 previously unknown Argentine and Peruvian taxa that agree in most points with C. planosae.
Chromocryptus probably is related to at least some of the stocks formerly included under Trachysphyrus (Porter 1967). It may be particularly close to the central Argentine Xiphidium group of Tra- chysphyrus (Porter 1967: 21 1- 15). In both groups the 2nd gastric tergite has large, dense punctures and the base of the petiole shows an at least moderately well developed lateral flange or blunt tooth. The Xiphidium group differs from Chromocryptus in having the axillus vein far from the hind margin of its wing. Species of the Xiphidium group thus belong to Trachysphyrus, as most recently defined (Townes 1969: 181-3). However, displacement anteriad of the axillus vein (a derived feature in ichneumonids) probably has occurred independently several times within various evolutionary lines associated with Trachysphyrus. Further studies, consequently, might allow expansion of Chromocryptus to include those species currently grouped with Trachysphyrus xiphidium. Chromocryptus ranges throughout most of the New World from the northeastern United States and California to Mexico, Peru,



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Argentina, and Brasil. It has not been recorded from the Neantarc- tic Biotic Province of South America (Chile and adjoining south- west Argentina) nor from any of the West Indian islands. Of the known species, 6 occur in northern Argentina and/or Brasil, 1 is reported from the Peruvian Coastal Desert, 1 from Mkx- ico and the southern United States, 1 from Florida, 1 from the northeastern United States, and 1 from central California. Habitats occupied by Chromocryptus include Temperate Deciduous Forest (C. planosae), Floridian Subtropical Evergreen Forest (C. weemsi and eastern populations of C. mesorufus), Tamaulipan Subtropical Thorn Scrub (western populations of C. mesorufus), Chaco-like Tropical Thorn Scrub on the north Peruvian coast (C. teres), the northwest Argentine Andean ecotone between Subtropical Cloud Forest, highland Chaco, and Montane Temperate Forests (C. alva- radoi and some populations of C. tomsici), the northwest Argentine Andean Desert or Prepuna (some populations of C. tomsici), the north-central Argentine Dry Chaco (C. prosopis, C. golbachi), and very wet subtropical to tropical forests in the south Brasilian biotic zone (C. huebrichi).
Wherever it occurs, Chromocryptus almost always seems extra- ordinarily scarce. Specimens which I have observed usually were crawling or flying within the shelter of spiny shrubs. I suspect they may be common in such microhabitats, from which they rarely emerge to enter Malaise Traps and in which it is difficult for the collector to detect and net them.
Chromocryptus have been reared most often from tough lepidop- terous cocoons attached to bark. Lasiocampidae are the usual hosts, with C. planosae recorded from Epicnaptera and Tolype in the eastern United States and C. huebrichi cited from Titya near Buenos Aires, Argentina. There is also 1 rearing for C. planosae from the arctiid genus Halisidota.
Parasitism by Chromocryptus is gregarious, with up to 22 speci- mens emerging from a single host cocoon (Porter 1967: 33-4). This habit also has been recorded for the Chilean Trachysphyrus horsti and the Argentine T. chacorurn, but has not been noted among such commonly reared Holarctic bbtrachysphyroid" genera as Itamoplex and Buathra. Pratt (1945: 551) believes C. planosae to be polyem- bryonic.




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INFRAGENERIC AFFINITIES
Chromocryptus forms a compact genus in which establishment of subgenera or species groups seems as yet unnecessary. Little has been discovered about the ancestral or derived nature of the states of features used to separate these species, so that phylogenetic anal- ysis within Chromocryptus would be premature. Nonetheless, some idea of relationships within the genus may be derived from the following hierarchically indented schema: I. Ovipositor stout, its tip not sagittate; fore tibia swollen; clypeus approximately symmetric, convex to bluntly pyra- .................................
midal C. alvaradoi.
11. Ovipositor slender and delicate, its tip sagittate; fore tibia not inflated; clypeus asymmetrically and nasutely pyramidal. 1. Mesopleuron with conspicuous, long, rather regular wrin- kles that radiate extensively from speculum; mesoscutum with some smooth areas between puncto-reticulation on .......................
both median and lateral lobes
............................. C. poecilma, C. teres. 2. Mesopleuron without or with only short wrinkles radiating from speculum; mesoscutum usually without smooth areas among the coarse sculpture ......... C. planosae, C. mesorufus, C. weemsi, C. vandykei, C. huebrichi, C. prosopis, C. golbachi, and C. tomsici. Plausibly, Chromocryptus alvaradoi diverged early from the Chromocryptus evolutional line. Specializations, possibly asso- ciated with parasitism of small hosts and with a newly acquired polyembryonic life style, then led to the origin of a cluster of smaller species with more delicate ovipositors. More recently, this latter category has subdivided into the weakly differentiated Poecilma and Planosae complexes.
Listed below in alphabetic order are the collections which fur- nished material for this research and/ or in which type specimens are to be deposited. I refer to institutional collections by the name of the city in which they are located and to individual collections by the surname of their owner.




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CAMBRIDGE.
Museum of Comparative Zoology, Harvard Univer- sity, Cambridge, MASS 02 138.
COLLEGE STATION. Department of Entomology, Texas A & M University, College Station, TX 77843.
GAINESVILLE.
Florida State Collection of Arthropods, Bureau of Entomology, Division of Plant Industry, Florida Department of Agriculture and Consumer Services, P. 0. Box 1269, 191 1 S W 34th Street, Gainesville Fl 32602.
LAWRENCE. Department of Entomology, Snow Entomological Museum, The University of Kansas, Lawrence, KS 66045. OTTAWA. Canadian National Collection, Biosystematics Research Institute, Agriculture Canada, Ottawa KIA 06C, CANADA. PORTER. Collection of Charles C. Porter, 301 North 39th Street, McAllen, TX 78501.
TOWNES. American Entomological Institute, c/o Dr. Virendra Gupta, Bureau of Entomology, Division of Plant Industry, Florida Department of Agriculture and Consumer Services, Gainesville, Fl32602.
Genus CHROMOCRYPTUS
Chromocryptus Ashmead, 1900. Proc. U.S. Natl. Mus. 23: 41. Type: (Chromocryp- tus albopictus Ashmead) = planosae (Fitch). Mesostenimorpha Viereck, 1913. Proc. U.S. Natl. Mus. 44: 566. Type: (Cryptus nebraskensis Ashmead) =planosae (Fitch). Fore wing 3.4-9.1 mm long. Coloration usually black, red, and white (occasionally mostly red) with hyaline to moderately dusky wings. Flagellum moderately long and slender, a little flattened below toward apex in female, in male with some segments bearing sharp and linear longitudinal tyloids, its 1st segment 3.7-6.2 as long as deep at apex. Mandible rather short and broad with lower tooth often a little shorter than the upper. Clypeus in profile asymmetri- cally to symmetrically pyramidal, often nasute; its apical margin edentate. Occipital carina sharp and narrow, usually sinuate later- ally, reaching the weakly to moderately raised hypostomal carina below. Malar space 0.50-1.0 as long as basal width of mandible. Pronotum a little swollen on dorsal margin and with the sub- marginal groove vague or absent to shallowly defined; epomia sharp



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in scrobe and sometimes a little prolonged dorsad, where it occa- sionally terminates in a weak swelling. Mesoscutum with notaulus weak (sometimes rather strong in male) but traceable 0.5 or more the length of mesoscutum; surface at least in part coarsely and densely punctate, reticulo-punctate, or reticulately wrinkled. Meso- pleuron with speculum largely smooth and shining or sometimes much invaded by coarse wrinkling; its surface otherwise wrinkled and/ or puncto-reticulate; no ridge on prepectus below. Fore tibia scarcely to palpably inflated. Hind coxa with a sharp and strong subvertical groove externo-ventrally near base. Wing venation: radial cell 3.3-4.4 as long as wide; areolet large, moderately broad to rather high and narrow, its intercubiti slightly to definitely conver- gent above; 2nd recurrent vertical or reclivous, straight or slightly outcurved dorsad; discocubitus gently arched to broadly angled, sometimes with a vestigial ramellus; mediella beyond base usually weakly arched but fairly strongly arched in some males; axillus very close to and paralleling hind margin of wing. Propodeum: spiracle 1.3-3.0 as long as wide; profile short and high; basal trans-carina traceable and often sharp throughout; apical trans-carina irregular but traceable between cristae, sometimes strong; cristae large and obtusely cuneate to subligulate, often strongly projecting; median longitudinal carinae often traceable but weak in female but stronger in males, where they often define both an areola and a median apical area. First gastric tergite: petiole at base with a low and rounded to often prominent and subtriangular lamella; postpetiole usually strongly expanded, in females 1.7-2.3 as wide apically as long from spiracle to apex and in males 1.2-1.9 as wide; surface of postpetiole at most with vague dorsal carinae and with a few to very many medium sized to large and frequently conspicuous punctures; ventral longitudinal carina defined throughout. Second gastric tergite with abundant, medium sized to large, shallow to (often) sharp punctures that are mostly subadjacent to (often) adjacent; its setae abundant but short and often (but not always) mostly equal to the length of their interspaces. Ovipositor: sheathed portion 0.34-0.45 as long as fore wing; straight, slender to robust, compressed; nodus distinct to obsolete; notch vague to rather large but shallow; dorsal valve with a steep to very gentle taper between notch and apex; tip 0.14-0.21 as high at notch as long from notch to apex; ventral valve on tip with ridges weak and inclivously oblique to sharp and almost vertical.



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Porter & O'Neil- Genus Chromocryptus
FEMALES
(Female of C. golbachi unknown.)
1. South American species (Peru, Argentina, Brasil) ........... 5 1'. Mexican and North American species ..................... 2 2. Propodeum and mesopleuron largely red; 2nd and 3rd gastric tergites with apical white bands obsolete or absent ....... 3 2'. Propodeum and mesopleuron black and white, without con- spicuous red areas; 2nd and 3rd gastric tergites with promi- nent white apical bands ............................. - 4 3. Ground color of head, pronotum, and mesoscutum black; hind face of propodeum with a pair of white bars that reach from cristae to its apical margin; gastric tergites 4-7 with white apical bands and otherwise mostly blackish; punctures on upper 0.3 of mesopleuron mostly adjacent to subadjacent; sheathed portion of ovipositor 0.30 as long as fore wing. . - . ............................... 1. C. planosae (Fitcin) 3'. Ground color of head, pronotum, and mesoscutum red; propo- deum red with only a trace of whitish on cristae and area immediately apicad; gastric tergites 4-7 red with white apic a1 bands on 5-7; punctures on upper 0.3 of mesopleuron mostly reticulately confluent; ovipositor 0.40 as long as fore wing .............................. 2. C. weemsi (Porter) 4. Second and succeeding gastric tergites black with apical band; broad white stripe in anterior 0.6 of sternaulus; hind femur and tibia dusky on apex; hind tarsus partly white ....... - . .......................... 3 . C. mesorufus Cushman 4'. Second and succeeding gastric tergites red with a white apical band; only a trace of white in sternaulus; hind femur and tibia not dusky on apex; no white on hind tarsus. ....... - . ............................ 4. C. vandykei Townes 5. No white on propodeum; fore wing palpably darkened toward apex, 8.6-9.1 mm long; speculum invaded by coarse wrin- kles; fore tibia stout and inflated; propodeal spiracle 2.3-3 -0 as long as wide; ovipositor robust, its notch and nodus obs o- lete, its profile on tip weakly tapering and not sagittate, its ventral valve on tip with strong, almost vertical ridges ... - . ............................... -5. C. alvaradoi n. s p.



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4 14 Psyche [VOI. 92
5'. Hind face of propodeum with a pair of conspicuous white lat- eral bars between cristae and apical margin; fore wing nearly hyaline, 3.4-6.7 mm long; speculum largely smooth; fore tibia slender and scarcely inflated; propodeal spiracle 1.3-2.1 as long as wide; ovipositor slender and delicate, its notch and nodus usually detectable, its profile on tip more strongly tapering and sagittate, its ventral valve on tip with delicate, inclivously oblique ridges. ........................... .6 6. Mesoscutum not uniformly reticulo-punctate; mesopleuron with long wrinkles that radiate from speculum; gaster red with variably developed white apical bands on all but 3rd tergite 6'. Mesoscutum throughout coarsely reticulo-punctate; mesopleu- ron with fine reticulo-punctation and wrinkling but without long rugae that radiate extensively from speculum; gaster dull to bright red, sometimes with considerable blackish suf- fusion, and often with white absent or obsolete on apices of 1st and/ or 2nd tergites .............................. .8 7. Face mostly white; large white blotch in upper hind corner of mesopleuron; postpetiole with scattered large and strong punctures that become subadjacent laterad and apicad; 2nd gastric tergite with mostly subadjacent to adjacent large and sharp punctures. ................ .6. C. poecilma (Porter) 7'. No white on face or in upper hind corner of mesopleuron; postpetiole with only a few small to medium sized and scat- tered punctures; 2nd gastric tergite with abundant medium sized and shallow punctures that are separated by 1-3X their diameters or more. .................... .7. C. teres n. sp. 8. Pair of white stripes on mesoscutum; large anterio-median white blotch on mesopleuron; large, ellipsoid white area on apical 0.5 of lower metapleuron; no white on 1st and 2nd gastric tergites ....................... 8. C. huebrichi (Brkthes) 8'. No white on mesoscutum and on mesopleural disc; lower meta- pleuron at most with a tiny white area at apex; 1st (and ..
sometimes 2nd) gastric tergite with a white apical band. .9
9. White flagellar band begins on 6th segment; basal 0.3-0.6 of scutellum white; white apical bands on both 1st and 2nd gastric tergites; 4th gastric tergite with its broad white apical



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Porter & O'Neil- Genus Chromocryptus
415
band abruptly truncate laterally; areolet nearly parallel- sided, the intercubiti scarcely convergent above; propodeal cristae asymmetric, cuneo-ligulate; 2nd gastric tergite with its large punctures mostly subadjacent and surface between punctures with only faint micro-aciculation .............. ................................ 9. C. prosopis n. sp. 9'. White flagellar band begins on 7th segment; basal 0.3 of scutel- lum white; no white apical band on 2nd gastric tergite; white apical band of 4th gastric tergite nearly complete, not abruptly cut off laterally; areolet not parallel-sided, the intercubiti definitely convergent above; propodeal cristae symmetrically short-ligulate; 2nd gastric tergite with its large punctures in great part practically adjacent and surface between punctures with palpable micro-aciculation. ....... ................................ 10. C. tomsici n. sp. MALES
(Males of C. alvaradoi, C. mesorufus, C. poecilma, C. teres, and C. vandykei unknown).
................
1. South American species (Argentina, Brasil) .3
1'. North American species ................................ .2 2. Ground color of head, pronotum, and mesoscutum black; hind face of propodeurn with a pair of white blotches from cristae to hind margin; gastric tergites 4-7 with white apical bands and otherwise mostly blackish; punctures on upper 0.3 of mesopleuron at least partly discrete ..................... ................................ 1. C. planosae (Fitch) 2'. Ground color of head, pronotum, and mesoscutum red; propo- deum red with only a trace of white on cristae and area immediately apicad; gastric tergites 4-7 uniformly red; punc- tures on upper 0.3 of mesopleuron reticulately confluent. .. ........................... 2. C. weemsi (Porter). 3. Pair of white stripes on mesoscutum; white blotch on clasper; no white on 1st and 2nd gastric tergites, the 4th and following tergites blackish with white apical bands; notauli faint, extending 0.5-0.6 the length of mesoscutum .............. ............................ 8. C. huebrichi (Brkthes)



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Psyche
[Vol. 92
3'. No white stripes on mesoscutum (occasionally a median white blotch); no white on clasper; ground color of gaster usually in large part red, 1st gastric tergite with at least a vague white apical band, 2nd tergite with or without a white band, 3rd tergite with or without a white band, 4th and 5th tergites often with an apical white band; notauli fine but impressed, ............. reaching 0.7-0.8 the length of mesoscutum .4 4. Large white spot on mesoscutum between ends of notauli; post- scutellum mostly white; gaster bright red with broad white apical bands on tergites 1-5; mesopleural disc on upper 0.5 with rather fine and longitudinally biased variably reticulate wrinkling and on lower 0.5 uniformly puncto-reticulate; postpetiole 1.8-1.9 as wide apically as long from spiracle to apex ............................ 1 1. C. golbachi n. sp. 4'. No white on mesoscutum; no white on postscutellurn; gaster duller red with white bands not as described above; meso- pleural disc uniformly with rather fine and granular puncto- reticulation; postpetiole 1.2- 1.7 as wide apically as long from ..................................... spiracletoapex 5 5. Mesopleuron extensively white; gaster bright to dull red with some dusky staining, with a broad white apical band on tergites 1 and 2, a narrow white band on tergite 3, a broad but laterally aborted white band on 4, and no white on 5th and following tergites; malar space 0.53-0.66 as long as basal width of mandible; postpetiole 1.6 as wide apically as long from spiracle to apex; large punctures on postpetiole uni- formly subadjacent to reticulately confluent .............. ................................ 9. C. prosopis n. sp. 5'. Mesopleuron with only a small white area in lower hind corner; gaster red with some dusky suffusion, with white vaguely on apex of 1st tergite, no white apical bands on tergites 2 and 3, with a broad but laterally aborted band on tergite 4, and with white apical bands on 5th and following tergites; malar space 0.80 as long as basal width of mandible; postpetiole 1.2 as wide apically as long from spiracle to apex; large punctures on postpetiole irregularly spaced, not uniformly subadjacent or denser, sometimes mostly sparse ..................... ................................. 10. C. tomsici n. sp.



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1. Chromocryptus planosae (Fitch)
(Figs. 1, 2, 6).
Phygadeuonplanosae Fitch, 1856. Trans. New York State Agr. 'Soc. 15: 501. Holo- type $: New York (Washington).
Cryptus nebraskensis Ashmead, 1890. Proc. U.S. Natl. Mus. 12: 412. Type 9: West Point, Nebr. (Washington).
Chromocryptus albopictus Ashmead, 1900. Proc. U.S. Natl. Mus. 23: 41. Type $: Urbana, 111. (Washington).
Agrothereutes (Itamoplex) cressonii Viereck, 1917. Bull. Connecticut Geol. Natl. Hist. Surv. 22: 1332. Type 9: Connecticut (Philadelphia). Chromocryptus planosaeplanosae Townes, 1962. Bull. U.S. Natl. Mus. 216(3): 257. -- -
Trachysphyrusplanosae Porter, 1967. Mem. Amer. Ent. Inst. 10: 207. ............................
1. C. alvaradoi (Jujuy Province, Argentina) .................................
2. C. poecilma (Minas Gerais, Brad).
A
................................
3. C. zeres (Simbal near Trujillo, Peru) 4. C. huebrichi (Curitiba, Brasil ... also south to eastern Argentina) ........ V 5. C. prosopis (Santiago del Estero, Argentina) ......................... 0 ..............................
6. C. tomsici (Jujuy Province, Argentina) å
.................
7. C. golbachi (Santiago del Estero Province, Argentina) V
Fig. 1.
Distribution map for Chromocryptus in tropical and subtropical South America.




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Townes' revisionary study of this taxon (1962: 256-8) summarizes the characters that separate C. planosae from other North and Mid- dle American Chromocryptus. It defines the species biogeographi- cally as a member of the Temperate Deciduous Forest Biome in eastern North America from Massachusetts south to Pennsylvania west to Michigan, Indiana, Illinois, and Nebraska. It also cites numerous host records for C. planosae as a "gregarious parasite'' in pupae of the bombycoid genera Tolype and Epicnaptera (Lasio- campidae) and one rearing from the noctuoid Halisidota (Arctiidae). No additional data have accumulated on C. planosae, since the publication of Townes' monograph.
2. Chromocryptus weemsi (Porter)
(Figs. 1, 5)
Trachysphyrus weemsi Porter, 1974. Fla. Ent. 57: 331. Holotype Q: Sarasota, Florida (Gainesville).
Chromocryptus weemsi Carlson, 1979. Catalog of Hymenoptera in America North