Charles C. Porter.
South American and Floridian disjuncts in the Sonoran genus Compsocryptus (Hymenoptera: Ichneumonidae).
Psyche 93:13-34, 1986.

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SOUTH AMERICAN AND FLORIDIAN DISJUNCTS IN THE SONORAN GENUS COMPSOCRYPTUS
(HY MENOPTERA: ICHNEUMONIDAE).
BY CHARLES C. PORTER!
Department of Biological Sciences, Fordham University Bronx, NY 10458
Taxonomy
Most Compsocryptus may be recognized at a glance by their elegantly yellow banded brown or black wings, large and anteriorly wide areolet, short and weak notauli, axillus intermediate in posi- tion between the anal margin of the hind wing and the submediella, strong ventro-lateral carina on postpetiole, and long, upcurved ovipositor.
My concept of this genus agrees, as to species included, with Townes' most recent definition (1969:203-4). Several of Townes' diagnostic features, however, do not apply to the Compsocryptus I have examined (C. fasciipennis, C. fuscofasciatus, C. melanostigma, C. texensis, and C. xanthostigma). All Compsocryptus I have seen possess a sharp and strong subvertical groove externo-ventrally near the base of the hind coxa, while Townes describes the hind coxa as "without a groove"(1969:203). All Compsocryptus examined by me have, in the female only, a prominent crescentic to subtriangular baso-lateral flange at the base of the petiole, while Townes main- tains that the petiole is "without a lateral tooth at the base" (1969:203). Compsocryptus forms a compact genus whose species, despite their far-flung and discontinuous distribution, seem unusu- ally homogeneous in color and structure. I thus suspect that all members of the genus will turn out to have a basal first g:istric projection in the female and a strong hind coxal groove. 'Research Associate, Florida State Collection of Arthropods, Florida Department of Agriculture and Consumer Services, Division of Plant Industry, P.O. Box 1269, Gainesville FL 32602.
Manuscript received by the editor May 28, I985 Pachi Ìö3:IJ- (1986). hup Wpsycht einclub nrgÌö3t91-01 html



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Relationships
Compsocryptus displays close affinity to Trachysphyrus, Aelio- potes, Joppidium, and Lanugo. Superficially, in color, size, habitus, and many details of geographic distribution, Lanugo especially parallels Compsocryptus. However, the phylogenetic connection seems distant, since Lanugo has only a weak groove at the base of the hind coxa, the axillus vein closer to the hind margin of the wing than to the submediella, and the ovipositor straight and shorter than in Compsocryptus. The South American Trachysphyrus [now re- garded as including only the Imperialis group, as defined by Porter (1967:275-319)] seems directly associated with Compso-
cryptus. Important characters that separate Trachysphyrus include: dark but never pale banded wings; body color usually metallic blue, green, or purple; female flagellum scarcely flattened below; notauli usually (not always) extending beyond the middle of the mesoscutum; surface of mesoscutum shining, never extensively mat; 1st gastric tergite without a baso-lateral expansion; and 2nd gastric tergite in many species smooth and polished (in numerous others mat). Aeliopotes paitensis (Porter, 1986) in some ways (especially the petiolar tooth) seems annectant between Compsocryptus and Trachysphyrus but in other features (epomial development) is aberrant and deserves generic status. Finally, Joppidium seems to be a direct offshoot of Compsocryptus. Joppidium is more slender than Compsocryptus (postpetiole at least 1.5 as long as wide), lacks a ventro-lateral carina on the 1st gastric tergite, and has the female flagellum more strongly flattened below toward apex. Some of its species have a baso-lateral tooth on the petiole and yellow banded wings, as in Compsocryptus. Joppidium also parallels its relative in distribution, with numerous Sonoran species, several which extend (but not disjunctly) into the southeastern United States, and with an isolated species group in subtropical Brasil and north Argentina (no representatives in the Peruvian Coastal Desert). Biogeography and Ecology
Compsocryptus belongs to the Sonoran Biogeographic category (Porter 1980:25-7). Possibly the genus evolved during the last half



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19861 Porter-Sonoran genus Compsocryptus 15 of the Tertiary somewhere in southwestern North America. Cer- tainly, its xerophilous species would have adapted well to the increasingly drier climates of post-Oligocene times and to the microphyll and sclerophyll Madro-Tertiary Geoflora which then overspread the ever-rising Sierra Madre and Rocky Mountains. On the other hand, Compsocryptus may have originated in western and southern South America, where so many of its relatives are centered today. Here, the Miocene climate paralleled that of the Sonoran region. By the end of the Miocene the Argentine pampas had become well developed, composites and other dry-adapted plants of open habitats were radiating vigorously, and long dry seasons began to characterize the middle latitudes as a result of the reduced rainfall and "the ever-increasing rain-shadow effect of the rising Andes" (Solbrig 1976:22-3). Thus arose the Chaco, an austral Sonora. Whatever may have been its genesis, Compsocryptus today is centered in the western United States and northern Mexico (15 species). It also has 3 remarkable disjuncts: C. fasciipennis in tropi- cal Florida and Cuba, C. fuscofasciatus in the Peruvian Coastal Desert, and C. melanostigma in north Argentina and nearby areas in Paraguay and Brazil.
Even the most geographically remote species of Compsocryptus differ only in apparently minor features of color and sculpture. This fact may suggest that the disjunctions noted above arose in compar- atively recent times. Both the increasing aridity of the later Tertiary and xerothermic episodes within the Pleistocene probably allowed semiarid communities (Thorn Scrub, Subtropical Deciduous Forest, etc.) to range almost uninterruptedly from the southeastern United States to Argentina. Wet periods during the Pleistocene (glacial maxima at higher latitudes) would have favored the expansion of forests and probably caused the fragmented distribution that is observed among modern Compsocryptus species. For example, "during a past period of low rainfall a prairie type flora, such as is found today in Texas and Arizona, was able to extend its range into the eastern United States, and remnants of this flora reflecting dry conditions still exist on parts of the west coast of Florida and on Big Pine Key" and other lower Florida Keys (Spencer and Stegmaier 1973: 13). Such dry periods occurred both in the Pleistocene and in the climatically unsettled late Tertiary. They



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16 Psyche [vol. 93
allowed eastward expansion by a whole complex of Sonoran Biota, from Opuntia, Cereus, Acacia and other xerophytes, to insects like Compsocryptus fasciipennis, several species of Joppidium, Lanugo retentor, Derocentrus longicaudis, Eiphosoma dentator (all Ich- neumonidae), Eumenes srnithii (Eumenidae), Stictiella (Sphecidae) and even to vertebrates, such as the reptiles Crotalus, Sistrurus, Pit- uophis, Sceloporus, and Gopherus.
Concurrently, similar physio-climatic events could have produced the plausibly vicariant differentiation of Compsocryptus fuscofasci- atus and C. melanostigma in South America. As discussed under C. melanostigma, the common ancestor of these two species may have ranged in Chaco vegetation from Argentina to coastal Peru at a time before the Andes were high enough at this latitude to impede east-west exchange of lowland biota.
As intimated throughout the above discussion, Compsocryptus prefers semiarid or arid environments, but also may be abundant in open, degraded subtropical humid forests. Compsocryptus mela- nostigma, for example, has been cited from the very wet Selva Tucumano-Bolivians and Selva Misionera in Argentina. However, the majority of these forest records are from ecotones between forest and Chaco or from sites in the first stages of secondary succession (logging roads, clearings, windfalls, etc.). This fact demonstrates how precarious is the present-day equilibrium between forest and scrub (Selva and Chaco). Almost all modern forests are surrounded by drier environments, whose aggressive biota tends to encroach with the slightest ecological perturbation. Compsocryptus melano- stigma in Argentina and C. fasciipennis in south Florida are among the many indicator species of these often anthropogenic and usually disastrous environmental changes, from forest to scrub and finally to desert.
Hosts
Compsocryptus are among the most common, conspicuous, and frequently collected of New World Ichneumonidae. Nonetheless, practically nothing is known about their host relationships. Only Cornpsocryptus melanostigma has been reared. It parasitizes noc- tuid moths of the genera Alabama and Pseudaletia. Alabama larvae feed on cotton and pupate in rolled leaves. Pseudaletia larvae feed at night on many kinds of grains and grasses, hiding by day under



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19861 Porter-Sonoran genus Compsocryptus 17 clods of earth or in other slightly subsurface shelters. Pseudaletia pupae are made in the ground. These data explain why females of C. melanostigma most often are collected on the ground or from low vegetation. Other Compsocryptus species occur in similar micro- habitats and probably parasitize comparable hosts. Listed below in alphabetic order are the collections in which material from this study has been or is to be deposited. Institutional collections are designated by the name of the city where they are located. Individual collections are referred to by the surnames of their owners.
CAMBRIDGE. Museum of Comparative Zoology, Harvard Univer- sity, Cambridge, MA 02138.
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 SW 34th Street, Gainesville, FL 32602.
LAWRENCE. Department of Entomology, Snow Entomological Museum, The University of Kansas, Lawrence, KS 66045. 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, FL 32602.
Genus COMPSOCR YPTUS
Cryptoideus Ashmead, 1900. Proc. U. S. Natl. Mus. 23: 42. Type: Cryptuspurpuri- pennis Cresson.
Compsocryptus Ashmead, 1900. Proc. U. S. Natl. Mus. 23: 43. Type: Cryptus calip- terus Say.
Callicryptus Ashmead, 1900. Proc. U. S. Natl. Mus. 23: 43. Type: (Cryptus "fascia- tus" Brullk) = fasciipennis Brullk.
Stictocryptus Cameron, 1908. Trans. Amer. Ent. Soc. 34: 243. Type: (Cryptus "fascia- tipennis"Brull6) = fasciipennis Brullk.
Sophocryptus Mallo, 1961. Idia 165: 17. Nomen nudum.



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Fore wing 7.2-13.0 mm long. Wings usually dark with yellow transverse bands. Female flagellum somewhat widened and flat- tened below on apical 0.3. Male flagellum with linear tyloids on many intermediate segments. Mandible usually moderately broad with lower tooth slightly shorter than upper tooth (rarely long and slender with lower tooth much shorter than upper). Clypeus rather large, gently convex in profile; its apical margin always edentate and straight to weakly convex. Occipital carina sharp and narrow. Malar space about 1.0 as long as basal width of mandible. Prono- turn with epomia well defined but not extending much dorsad or ventrad of scrobe. Mesoscutum with notaulus faint, traceable less than half its length; surface mat, dully shining, or sometimes pol- ished and with numerous, small to medium sized, crowded to well separated punctures (punctures sparser in males). Mesopleuron without a ridge on prepectus below. Hind coxa with a sharp and strong subvertical groove externo-ventrally near base. Wing vena- tion: areolet large, symmetrically to asymmetrically pentagonal, intercubiti slightly to definitely convergent dorsad, front side of areolet (2nd abscissa of radius) 0.9 to more than 1.0 as long as 1st intercubitus (mesa1 side of areolet); discocubitus gently arched, without a ramellus; mediella nearly straight; axillus long, diverging from anal margin of wing, as close to submediella as to anal margin. Propodeum: spiracle elongate; apical trans-carina varying from strong throughout, to strong laterad but obsolete mesad, to almost completely absent; cristae usually defined and subcrescentic to bluntly triangular (cristae often obsolete in males). First gastric ter- gite in female with a prominent crescentic to subtriangular baso- lateral flange and with the postpetiole strongly expanded but in male without a baso-lateral expansion and with the postpetiole slender; ventral longitudinal carina usually sharp throughout, dorso-lateral and dorsal carinae less well developed, weakest in males. Second gastric tergite mat with very fine and dense punctures (sparser in males) which sometimes become more widely spaced mesad and with short, recumbant, mostly overlapping setae. Ovi- positor: long, sheathed portion 0.5-1.4 as long as fore wing, gently upcurved, cylindro-compressed, its tip elongate (0.10-0.25 as high at nodus as long from nodus to apex), nodus low and without a notch, ventral valve on tip with sharp and inclivously oblique ridges.



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Porter-Sonoran genus Compsocryptus
19
Fig. I .
Campsocryptus melanosligma. 9. Photograph of whole insect in lateral view. Fig. 2. Compsocryptus fuscofasciatus. Q. Photograph of whole insect in lateral view.




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KEY TO THE SOUTH AMERICAN SPECIES OF COMPSOCRYPTUS 1. Fore wing yellow with 2 narrow brown cross bands and with brown on apex; 2nd abscissa of radius 0.9-1.0 as long as 1st intercubitus; female mesopleuron usually with some strong longitudinal wrinkling ................................ ....................... 1. C. fuscofasciatus (Brullk) V. Fore wing dark brown with a broad median yellow cross band and a large subapical yellow blotch; 2nd abscissa of radius 1.2- 1.5 as long as 1 st intercubitus; female mesopleuron with- out any longitudinal wrinkling, almost uniformly puncto- reticulate .................. 2. C. melanostigma (Brullk) 1. Compsocryptus fuscofasciatus (Brullk) (Fig. 2, 3,4)
Cryptus fusco-fasciatus Brullk, 1846. In Lepeletier: Histoire naturelle des insectes. Hymknoptkres 4:194. Holotype Q: Peru, Lima (lost). Callicryptus ornatipennis Cameron, 1902. Trans. Amer. Ent. SOC. 28:372. Holotype Q: Peru, Callao (London).
FEMALE. Color: antenna ferruginous on scape and pedicel, yel- lowish ferruginous on 1st (sometimes also 2nd and 3rd) flagello- mere, mostly yellow on flagellomeres 2,3 or 4-9, brown and yellow on flagellomeres 10-1 1 or 12, and black or brownish black beyond 12th or 13th flagellomere; head ferruginous with black on apex of mandible; palpi dull ferruginous; mesosoma ferruginous with inconspicuous dusky staining on some margins and sutures or some- times with rather extensive black markings (as described for male); gaster dull ferruginous with faint dusky staining on 2nd and follow- ing tergites or occasionally with better defined black areas toward base on 2nd and 3rd tergites; legs ferruginous, duller on tarsi, with dusky staining on apical tarsomeres, narrowly on apex of hind tro- chantellus and base of hind femur, and with blackish on much of hind tibia except toward its paler (sometimes contrastingly flavo- ferruginous) base; fore wing light yellow with three brown areas as follows: a broad transverse band on most of apical 0.3 of median cell, on base of discocubital cell, on apical 0.3 of submedian cell, on basal 0.3 of 1st brachial cell, and on adjoining region of anal cell; a second brown cross-band covering basal 0.3 of radial cell, apical 0.3



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19861 Porter-Sonoran genus Compsocryptus 2 1 of discocubital cell, areolet, apical 0.5 of 2nd discoidal cell, and expanding below to cover all but basal 0.3 (or less) of 2nd brachial cell; as well as with a third brown area on apical 0.5 of 3rd cubital cell and apical 0.5 of 3rd discoidal cell and confluent below with dark area of 2nd brachial cell; hind wing pale yellow with apical 0.3 dusky and with dusky staining prolonged more narrowly far basad on its hind margin, as well as sometimes with an irregular transverse dusky area at level of nervellus.
Length of fore wing: 10.5-12.5 mm. Flagellum: 1st segment 3.2-3.5 as long as deep at apex; apical segments averaging 0.7-0.8 as long as wide. Malar space: 1.0 as long as basal width of mandible. Mesoscutum: dully shining with abundant, dense, sharp, tiny punc- tures which emit inconspicuous, short and mostly close-packed setae. Mesopleuron: surface with delicate to strong, trans-biased puncto-reticulation and, at least ventrad, usually with some strong longitudinal wrinkling. Wing venation: radial cell 3.6-4.1 as long as wide; areolet about as high as broad, symmetrically pentagonal, intercubiti weakly to moderately convergent above, 2nd abscissa of radius 0.9-1.0 as long as 1st intercubitus. Hind femur: 5.6-6.0 as long as deep. Hind tibia: below and laterally on apical 0.5 with a few, scattered enlarged setae. Propodeum: apical face discrete from the gently arched basal face and almost vertical; basal trans-carina traceable throughout, uniformly fine and sharp or sometimes partly weak and irregular. First gastric tergite: dorso-lateral carinae per- current but faint; dorsal carinae well defined, but not sharp, on apex of petiole and basal 0.5 of postpetiole; surface of postpetiole mat, sometimes dully shining toward apex, with fine micro-reticulation and with tiny, sparse, shallow punctures that are best developed apico-laterad (where their short setae partially overlap). Ovipositor: sheathed portion 0.69-0.81 as long as fore wing; tip 0.15-0.17 as high at nodus as long from nodus to apex. MALE.
Differs from female as follows: Color: pedicel sometimes marked with black; flagellomeres 1-5 (sometimes up to 9) ferrugi- nous to yellowish with dusky staining, mostly above; at least flagel- lomeres 10-13 yellow with some ferruginous staining; rest of flagellum black; head ferruginous with yellowish on base of mandi- ble, much of face laterally, and narrowly on lower 0.6 of frontal orbit as well as with black on apex of mandible, broadly above and between antenna1 sockets (sometimes reaching and including stem-



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22 Psyche
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Compsocryptus en ~udamgrica
fuscofasciatus
melanos tigma (
Fig. 3.
Map showing geographical distribution of Compsocryptus fuscofasciatus and C. melanostigma.




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19861 Porter-Sonoran genus Compsocryptus 23 maticum) and on most of postocciput; mesosoma ferruginous with black markings usually better developed than in female and includ- ing areas on propleuron anteriorly, pronotal collar, spot on epornia (sometimes contiguous with black on collar), broad band on hind margin of pronotum, prescutellar groove, much of meso and metanotal axillary troughs, groove at base of propodeum, all of prepectus, mesosternal sulcus, hind face of mesosternum, broad band on hind margin of mesopleuron-prolonged dorsad along upper mesopleural margin to subalarum, broadly on all but dorsal margin of lower metapleuron, and irregularly on hind margin of propodeum; gaster dull to bright ferruginous with a little black at base of 1st tergite and with succeeding tergites sometimes only with irregular dusky staining and sometimes with well defined black areas toward base of tergites 2 and 3; fore and mid tibiae and tarsi more yellowish than in female; mid femur with some blackish stain- ing dorsad and apicad; hind tibia black with basal 0.15 contrastingly pale yellow; hind tarsus blackish with light yellow at least near base of 1st segment and sometimes almost throughout on both segments 1 and 2.
Length of fore wing: 8.5-9.6 mm. Flagellum: linear, largely per- current tyloids present on segments 1 1 or 12 to 19 or 20; 1 st segment 2.5-2.7 as long as deep. Malar space: 0.77-0.90 as long as basal width of mandible. Mesoscutum: shining with abundant, moder- ately small, sharp punctures that are separated by 1.0-2.0 their diameters and which emit dense, erect, moderately long setae. Mesopleuron: more shining than in female, with medium sized, sharp, dense, subadjacent to reticulately confluent punctures and some longitudinally biased reticulation. Hindfemur: 6.3-7.4 as long as deep. Hind tibia: with enlarged setae more abundant and con- spicuous than in female. Propodeurn: rather elongately convex in profile; apical face not discrete from basal; apical trans-carina weaker than in female, forming low and subcrescentic cristae or sometimes with cristae obsolete. First gastric tergite: ventro-lateral carina obsolete on petiole but sometimes becoming sharp toward apex of postpetiole; dorso-lateral and dorsal carinae in great part obsolete; surface of postpetiole smooth and shining with abundant but well separated tiny punctures that emit long and uniformly overlapping setae.




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24 Psyche [vo~. 93
SPECIMENS EXAMINED.
15 $ and 47 6: PERU, Lambayeque
Province, 33 km E. Olmos, Ruta a Jah, 23-VII-1975, C. Porter, L. Stange; 1 km S. Lambayeque, 24-27-VII-1975, C. Porter, L. Stange; La Libertad Province, Laredo nr. Trujillo, 7-8-VII-1974, C. Porter, L. Stange; Simbal nr. Trujillo, 4-7-VII-1974, C. Porter, L. Stange; Lima Province, Cupiche, 10 km E. Chosica, 25-VI-2-VII-1974, C. Porter, L. Stange; Palle nr. Chosica, 17-VII-1974, C. Porter, L. Stange; San Geronimo nr. Chosica, 28-VI-5-VII-1976, C. Porter, C. Calmbacher; nr. Surco on Carretera Central at km 59, 30-VI- 1976, C. Porter, C. Calmbacher.
RELATIONSHIPS. This Peruvian Coastal Desert endemic differs only in minor chromatic and structural features from the other South and North American Compsocryptus. It seems closely related to the Argentine C. melanostigma (Brullk) but may be distin- guished by the following characters: (1). Fore wing yellow with two narrow brown cross bands and with brown on apex (vs. dark brown with a broad median yellow cross band and a large subapical yellow blotch), 2. Second abscissa of radius 0.9- 1.0 as long as 1 st intercubi- tus (vs. 1.2-1.5 as long), 3. Female mesopleuron usually with some strong longitudinal wrinkling (vs, puncto-reticulate), 4. Male mesoscutum with punctures mostly separated by 1.0-2.0 their diameters (vs. 2.0 or more their diameters) and 5. Male flagellum with tyloids extending to segments 19-20 (vs. 21-23). FIELD NOTES.
Compsocryptus fuscofasciatus has been reported only from the northern and central Peruvian Coastal Desert between Lima and Piura. Here, it frequents most well watered habi- tats between sealevel and 1500 m. I have collected it along rivers and irrigation ditches in arid country as well as in orchards and degraded cloud forest. Like other Compsocryptus, this species most often occurs near or on the ground in exposed, disturbed, weedy or grassy places.
2. Compsocryptus melanostigma (Brullk)
(Fig. 1, 3, 5)
Cryptus melanostigma Brullk, 1846. In Lepeletier: Histoire naturelle des insectes. Hymknopthres 4:191. Lectotype Q: Brasil: "Prov. de Misiones" (Paris Museum).
Cryptus opaco-rufus Taschenberg, 1876. Ztschr. f. die Gesam. Natuw. Halle 4864. Lectotype 9: (Brasil): Paran6 (Halle).




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Porter-Sonoran genus Compsocryptus
Fig. 4.
Compsocryptus fuscofasciatus, Q. Fore wing, showing color pattern. Fig. 5. Compsocryptus melanostigma, Q. Fore wing, showing color pattern. Fig. 6. Compsocrypt us fasciipennis, Q. Fore wing, showing color pattern. Cryptus lateritus Taschenberg, 1876. Ztschr. f. die Gesam. Naturw. Halle 48:65. Lectotype 8: (Brad): Parana (Halle).
Callicryptus pulchrifasciatus Cameron, 1909. Trans. Amer. Ent. SOC. 35:437. Lecto- type Q: Argentina: Mendoza (London).
Sophocryptus bisulcatus Mallo, 196 1. Idia l65:17. Nomen nudum. FEMALE. Color: antenna with scape brownish ferruginous, pedicel dusky ferruginous with apex paler, and flagellum black with



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