P. A. Adams.
New Ant-lions from the Southwestern United States (Neuroptera: Myrmeleontidae).
Psyche 63:82-108, 1956.

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NEW ANT-LIONS FROM THE SOUTHWESTERN
UNITED STATES
(NEUROPTERA : MYRMELEONTIDAE)
BY PHILLIP A. ADAMS
Biological Laboratories, Harvard University In the course of identifying material from the South- west, the writer has encountered several new species and a new genus of Myrmeleontidae. Descriptions of these are given below, with a list and key to the species of the genus Eremoleon Banks. Sources of specimens are designated by the following abbreviations: CAS, California Academy of Sciences; CIlS, California Insect Survey, University of California, Berkeley; U'CD, University of California, Dav- is ; UCR, University of California, Riverside ; UCLA, Uni- versity of California at Los Angeles; LAM, Los Angeles County Museum; MCZ, Museum of Comparative Zoology, Harvard. The kindness of the staffs of these institutions in lending material is gratefully acknowledged. The terminology of the wing venation as used herein differs from the usual system (summarized by Markl)2 in several fundamental aspects. Markl's study is an excellent and invaluable treatment of the comparative morphology of the wing of the ant-lions, but unfortunately his scope, a tribal revision, was so large as to have discouraged detailed investigation of venation in other families. The best clues to the homologies of the myrmeleontoid wing are to be found in the primitive myrmeleonLoids -the Osrnylidae and the Myiodactylidae. A thorough account of the reasons for the adoption of the present system will be given in a forthcoming paper, dealing with the venation of the order as a whole.
In both wings, MA has become coalesced with Rs; the "basal piece" (Figure 8, "by') is to be seen at the base of the fore wing between R and MP. In the hind wing the basal Published with the aid of a grant from the Museum of Comparative Zoology, Harvard College.
ZMarkl, W., 1954, Verh. d. Naturforschenden Ges. Base1 65:178-263.



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19561 Adams - Myrmeleontidae 83
part of MA is retained in the Osmylidae as a sinuate vein running between MP and Rs shortly after the latter has di- verged from R; in the higher families this portion of the vein disappears. MA reappears, however, as the last branch from the "radial sector" (of previous authors, herein term- ed Rs+MA) , differing often in being strongly convex, and branching much more than do the true branches of Rs. Comstock has termed the triangular sector which results from this branching the "radial cuneate area". Since the "radial sector" is a complex vein, the cross veins which lie behind it, between the base of the wing and its divergence from R, cannot be termed "radial cross veins" ; they are in reality median cross veins. Because they have been used so much as a source of taxonomic characters, confusion might result in referring to them as median cross veins, hence the writer has adopted the term presectoral cross veins (Figure 8, "ps'?) .
There has been much controversy over the homology of the cubitus in the hind wing, Comstock holding that the con- vex forked vein which appears analogous tso the cubitus in the fore wing is actually M3-)-^. His system was based most- ly on evidence from the tracheation and from the absence of the "oblique vein" (MP3+4) from the hind wing. This forked vein is homologous to what has been termed MP3+4 in the other superfamilies; but it is the belief of the writer that it is actually composed primarily of cubital elements. There is evidence which indicates that CnAi has, in the hind wing of all the Neuroptera, coalesced with M, resulting in a condition similar to that found in the Meeoptera, so that CuAi appears as the posterior branch of the vein which has been called MP. In this paper, the veins which Mark1 terms "Ai" and "A2" are referred to as CuAa and CUP+A~, re- spectively. In the ant-lions, Ai is coalesced for a short dis- tance with 'CUP; the base of Ai although weak, is visible. Mark1 seems not to have noticed the double nature of this vein, although he shows the base of Ai correctly in his fig- ure 45. In the Osmylidae this coalescence has not taken place. The term inner cubital veinlets as used herein is equivalent to "anal veinlets" of previous authors. Few workers have made reference to the internal male



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84 Psyche
[September
genitalia when doing taxonomic work on the Mymeleonti- dae. The writer has found that the structure of the gon- arcus, and especially of the parameres, furnishes most use- ful characters; accordingly these are illustrated for all the species treated below, when male material was available. Within a species, there seems to be little variation in their size and structure ; unfortunately, it will sometimes be found that c'osely related species have almost identical in- ternal genitalia. Tjeder's terminology3 is used. Dorsally, there is present an arched sclerite, the gonar- cus, which bears at its apex a small hood-like structure, the mediuncus. Loosely articulated with the mediuncus are the parameres, which often are produced to f,orm hooks. Or- dinarily, these structures will be found to lie in a pouch 'between the bases of the ectoprocts ("male appendages") ; occasionally, this pouch will be found everted (Figure 32). The genital opening is on the ventral side of this sac, its position being marked by a small sclerite (Figure 32, dotted lines). As eversion takes place, the spatial relationships of the sclerites change markedly. In some species (e. g. Hesperoleon and relatives) the parameres have a thin re- gion near their middle, the hinge, where they may f'old when the genital sac is inverted. Upon eversion, they straighten out, and the points of the hooks become more divergent. The reader is cautioned that, because of this folding, it was not possible to draw all the figures from the same view- point, and that, as a result, several of the figures are dis- torted. The side view gives a much better impression of the shape of the parameres than does the rear view, for this reason. In Eremoleon the parts are fairly rigid. For critical examination, it is necessary to remove the gonarcus and parameres from the abdomen; these can easily be teased out after boiling the tip of the abdomen in KOH. The genitalia are stored in glass vials containing glycerine, pinned beneath the specimen. If the corks are boiled in paraffine, there is less danger of the glycerine soaking through to corrode the pin. This can also be mini- mized by piercing the cork at a 45 degree angle, so that 3Tjeder Bo, 1954, Ent. Medd. 27 :23-40.




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the glycerine will remain at the bottom of the vial whether the specimen is stored in a flat tray, or in a box placed vertically on a shelf.
Among the measurements given are the lengths of the pteroth'orax and the third abdominal tergite. These lengths can be measured easily and accurately; their ratio gives a value useful in describing the amount of abdominal elonga- tion.
Although the size of the labial palpi varies considerably within a given species, the size and shape of the terminal segment tends to be quite characteristic. In this segment is a sense-organY4 to accomodate which it is swollen to a great- er or lesser degree. In the male, the terminal segment is often smaller than in the female.
Genus Eremoleon Banks.
Banks, 1901, Trans. Am. Ent. Soc. 27 :365; 1928, Bull. Mus. Comp. Zool. 68: 69-71; Proc. Calif. Acad. Sci. 4th Ser. 24: 143-144.
This genus is still too poorly represented in collections to enable a full-fledged revision; the following is simply a list of the described species, with a key and such figures as may be found helpful in identification.
Key to the Species of the Genus Eremoleon. 1.
A very slender species, the antennae about 1% times length of head and thorax together; third abdominal tergite 1.4 times length of pterothorax
...... longior
- More robust, the antennae at most equal to length of head and thorax together; third abdominal tergite at most equal to length of pterothorax .............. 2 2.
A deep black mark on base of fore wing, mesepimeral wing process black ...................... nigribasis - No such mark, mesepimeral wing process brown or yellow ........................................ 3 3. CuP+Ai in fore wing curved forward, margin curved outward, so that the space between is wider in middle than at ends; prothorax wider than long . . . mexicam 4Eisner) T.) 1953, J. Morph. 93 : 109-122.



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Psyche [September
CuP+Ai in fore wing straight or sinuate, the space between wider at base than middle; prothorax longer than wide .................................. 4 Mesepimeral wing process dark umbraceous ; 12-16 . . . . . . . . . branches of CuP+Ai in fore wing macer
Mesepimeral wing pr'ocess yellow; 11 or less, usually . . . . . . . 8-9, branches of CuP+A, in the fore wing 5
Antennal segments longer than wide; antennae about as long as head and thorax together, the pedicel and .......... scape black; wings heavily spotted gracile
Antennal segments wider than long; antennae shorter than head and thorax together, pedicel and scape pale; wings only weakly marked ...................... 6 Large species (fore wing 30 mm., or longer) ; fore ti- bial spurs as long as 2 tarsomeres; all setae light brown . . . . . . . . . . . . . . . insipidus Smaller species (fore wing 28 mm., or less) ; fore ti- bial spurs as long as 3V2 tarsomeres; most setae on legs, and many on thorax, black . . . . . . . . . . . pollens Eremoleon macer (Hagen)
Figures 21, 30
Myrmeleon macer Hagen, 1861, Synopsis of N. Amer. Neur. :236
Eremoleon macer Banks, 1901, Trans. Amer. Ent. Soc. 27 :365 ; 1928, Bull. Mus. Comp. Zool. 68 :70 ; 1938, Car- negie Inst. Wash. Pub. 491 :235
Segura vitreus Navas, 1914, Ent. Zeitung 28:18 In the Museum of Comparative Zoology there is a pair of specimens from Mexico (Apatzingan, Michoacan, 12,000 feet, Aug. 11. 1941, H. Hoogstral. and Jacda, Hidalgo, 4,500 feet, June 22, 1939, Ralph Haag) which are quite similar t,o macer in general characteristics, but which have more slender wings, with fewer presectoral cross veins in the fore wing (7 or 8, but about 10 in macer), and with more branches of CuP+Ai in the fore wing; the vertex is dark pruinose, with two well-defined rows of spots in the Jacala specimen. The genitalia are like those of macer. De- spite the differences from macer, it does not seqrn advis- able to consider them distinct without more material



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19561 Adams - Myrmeleontidue 87
from this area. Should it then appear that they are dis- tinct, the name mexicana Navas might be appropriate. The wing shape and markings are similar to those of mexi- cana; in the Apatzingan specimen the anal area in the fore wing is widened like Navas' figure (Figure Ib), but not to such a marked degree. The vertex and pronotal markings do not agree well with the figure.
Figure 1. Eremoleon mexicana
(Nav.) a. Head and prothorax, b. Axillary region of anterior wing, c. Axillary region of posterior wing (from Navas, 1925).
Segura vitreus has long been considered a synonym of macer; the description fits well except "Abdomen 8 cercis manifestis, cylindricis," and "Abdomen longum, in 8 alis longius . . .," characters which are not found in other species of Eremoleon or in related genera. Probably the type specimen, collected in 1871, has another abdomen affixed.
Eremoleon rnexicana (Navas)
Figure 1 a, b, c
Novuiqa mexicana Navas, 1925, Mem. R. Acad. Ci. Artes Barcelona 19 :189-190, fig. 17.
Eremoleon macer Banks, 1936, Carnegie Inst. Wash. Pub. 491 :235
In all probability, this is E. macer; however, as mention- ed above, the figure shows a differently shaped CuP+Ai in the fore wing. According to the description, the wings are acute, and less prominently marked than in macer, there being only a small spot at the connection of CuP+Ai and CuAs, at the rhegma, and at the hypostigmatic cell. The hind wing is immaculate.




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88 Psyche [September
Ekemoleon insipidus, new species
Figures 19, 20, 28
Head pale; a faint indication of an interantenna1 dark band; first row of vertex scars implemented - a dash on each side and double medial scar; behind is a pair of in- distinct brown submedian spots; last segment of labial palpi large, strongly swollen (Figure 28) ; antennae pale, club infuscate, flagellar segments all broader than long. Pronoturn gray-brown, a pair of small submedian spots be- fore furrow; behind furrow a pair of wide median brown bands, broadly separated by pale posteriorly ; a short lateral dark atripe each side, Nota mostly infuscate, a pair of broad brown stripes on mesoscutellum. Propleuron dark, mesane- pisternum with a brown stripe below, rest of pleura pale; mesepimeral wing process light ochraceous. Coxae pale;
a small basal brown spot; femora pale, very faintly spotted with brown ; large apical brown spot on anterior side ; tibiae lightly browndotted; tarsi pale. All setae, including long aeta on fore femur, pale, darkest ones a light yellow brown in color. Fore tibia1 spurs only as long as 2 tarsomeres, hind spurs, 1% tarsomeres.
First abdominal tergite pale, second dark with apical pale spot, third-sixth with small median pale spots at base and apex, and with large subbasal spot each side, broadly confluent medially and containing a dark spot each side. Apical tergites darker, with markings indistinct. Sternites pale to middle of 4th, beyond dark.
Venation largely pale; in fore wings most cross veins dark at ends; longitudinal veins dark where intersected by dark cross veins; each dark intersection surrounded by a small gray cloud in membrane; larger spots at apex of hypostigmatic cell, rhegma, and end of CuAa. In hind wing venation paler, membrane scarcely marked, dark spots at apex of hypostigmatic cell and rhegma.
EXPLANATION OF PLATE 3
Figures 3-7. Fig 3. Hesperoleon natnutus, Q , head and pronotum (Cathedral City, Calif.). Fig. 4. H. mfuscatits, g , head and pronotum (Antioch, Calif.). Fig. 5. H, @elitas, allotype, head and pronotum. Fig 5a. H, fidelihs, mesoscutellum. Fig. 6, E~emokow gradls, holotype, $ , head and thorax. Fig. 7. Tyttholeo'n puerilis, holotype, head and pronotum.



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90 Psyche [Sept,ember
Fore wing: broad, 5-7 presectoral cross veins, the last usually connected to first cross vein from RS+MA; 9-11 branches of RS+MA; 9-10 cross veins between CuA and CuP+Ai; 8-10 cross veins between CuP+Ai and wing margin. Hind wing: cubital area broad, with four rows of cross veins.
Parameres (Figure 19) similar to those of macer, but the plates much farther apart, the inner margin strongly concave.
Body length, 24.-27.5 mm.; fore wing, length 30.5-35. mm., width 7.6-9.6 mm. ; hind wing, length, 28.5-32. mm. ; width 7.1-7.6 mm.; labial palpus, length 0.78 0.88 mm.; width 0.20-0.25 mm. ; antenna 5.7-6.7 mm. Holotype 8 : 5 mi. S. of San Miguel, L. Cal. VII-20-38, Michelbacher and Ross, in the Museum of Comparative Zoology. Paratypes : Riggs, Calif. 5-9-34 Q , Sperry (MCZ) ; Borego Valley, Calif., VI-6-40, C. D. Michener (CIS) ; El Mayor, L. Calif., IV-19-39, 1 8 1 9, E. S. Ross (GAS) ; Mejia Is., Angel de la Guardia I., Gulf of Calif.; VI-28-21, 8, E. P. Van Duzee (GAS).
These specimens are an insipid pale brown in color, with few really distinct markings on head or thorax ; nowhere is there any structure which is strongly pigmented - black is absent. The large, heavily swollen labial palpi (Figure
28) are characteristic. The holotype and the Mejia Isd. specimens are paratypes of affine Banks. Eremoleon gracile, new species
Figures 3, 17, 29
Face pale ; shiny dark interantennal band present, widely separated from first vertex row. First vertex row with a curved, dull black dash each side, and a pair of indistinct submedian brown dots; second row similar but fainter; be- hind a shiny black spot each side near eye, and an indis- tinct brown median mark. Labial palpi short, pale. Scape and pedicel shiny-black, flagellum slender, the segments slightly longer than wide, reddish brown, the tip strongly swollen, much darker.
Thorax mostly pale, notum marked as in Figure 3; meso- and metepisterna largely infuscate, epimera largely pale; expansion of mesepimeral wing process light yellow. Fore



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19561 Adams - Myrmeleontidae 91
coxa pale with small basal fuscous spot; mid and hind coxae infuscate laterally; femora pale, very faintly dotted at base of some setae, apical shiny black bands; tibiae dotted with dark, bearing postbasal and apical black bands. Tarsi pale, 5th tarsomere narrowly dark-tipped. Fore spurs equal three tarsomeres, hind spurs two ; setae mostly black, many white on femora.
First abdominal tergite pale with small dark preapical dot; second with pale basal band and apical dot; third-sev- enth dark-fuscous, a large pale spot each side, broadly con- nected medially, apical segments pale. First sternite with a black spot each side, second, third and base of fourth pale, apex of fourth, fifth, sixth and seventh fuscous. Wings : venation as in Figure 17. Base of wing pale, vena- tion largely pale, many cross veins dark wholly or at ends; longitudinal veins interrupted with dark at intersections of many cross veins; dark spots as indicated on figure. Measurements (mm.) : body length 21 ; abdomen 15 ; head and thorax 6.3 ; pterothorax 4.0 ; third abdominal tergite 3.4; antenna 6.8; fore wing 25.5 long, 6.6 wide; hind wing 24.0 long 5.5 wide; third segment, labial palpus 0.48 long, 0.12 wide.
H'olotype Q : Riverside, California, August 31, 1939, Paul De Bock (U'CR) : Deposited in the California Academy of Sciences.
Eremoleon nigribasis Banks
Figures 24, 27
Eremoleon nigribasis Banks, 1920, Bull. Mus. Comp. Zool. 64 :329 ; 1928, ibid. 68 :71; Proc. Calif. Acad. Sci., 4th Ser. 24 :I43
Eremoleon affine Banks, 1942, Proc. Calif. Acad. Sci., 4th Ser. 24:144, new synonymy
Utah, New Mexico, Arizona, Ba ja California. The type of E. affine Banks, as well as the paratypes from 5 mi. S. of San Miguel and Venancio, Baja Calif., are pale individuals of this species in which the black mark on the wing base is indistinct or absent.




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Psyche
[September
Eremoleon pallens Banks
Figures 18, 26
Eremoleon pallens Banks, 1941, Psyche 48: 101-102 The type series was collected from a mine shaft on Pichaco Peak, Arizona.
Eremoleon longior Banks
Figures 22, 23, 25
Erenzokon longior Banks, 1938, Carnegie Inst. Wash. Pub. 491 : 235
A cave-dwelling species from Yucatan, Mexico. Its slen- derness is combined with great fragility. Hesperoleon texanus Banks
Figures 42, 43
Hesperoleon texanus Banks, 1903, Proc. Ent. Sot. Wash. 5 :I75
This species has not previously been recorded from Cali- fornia. 'California specimens are larger than more eastern specimens, with longer antennae. Oklahoma specimens have the third abdominal tergite 1.5 times the length of the pterothorax, while in the California males this ratio was 1.7 and 1.9. The difference in color is striking, the Southern California specimens being much darker. Material from Riverside shows a gradation in size down to that of typical texanus, and a 9 from Olanche, California (MCZ) , is inter- mediate in its coloration between the paler texanus and the darker southern specimens. Some means of measurements of the California specimens follow (in mm.) : EXPLANATION OF PLATE 4
Figures 8-17. Fig. 8. Tyttholeon puerilis, 8 , wings: b- basal piece of MA, ps- presectoral cross veins, he- hypostigmatic cell, rh- rhegma (San Felipe Can., Calif.). Fig. 9. Maracandula bellula, gonarcus and para- meres, posterior view (Skyforest, San Bemardin0 Co., Calif.). Fig. 10. Same, lateral view. Fig. 11. Tyttholeon puerilis, gonarcus and parameres, lateral view (San Felipe Canyon). Fig. 12. Same, posterior view. Fig. 13. Hesperoleon minutus, holotype, apex of abdomen, lateral view. Fig. 14. Same, ventral view. Fig. 15. H. minutus, gonarcus and parameres, posterior view. Fig. 16. Same, lateral view.
Fig. 17. Eremoleon gracile, holotype,
wings.




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94 Psyche
[September
Body length, 8 37, 24; abdomen 8 30.2, Q 17.2; pterothorax 8 3.9, Q 3.7; fore wing, length 8 25.3, Q 24.2. width 8 6.6, Q 6.1; labial palpi (terminal segment) 0.76 long, 0.20 wide; antennae 8 7.25, Q 4.8; third abdominal tergite s 7.2, s 5.1.
In California, texanus is most likely to be confused with H. niger Currie, from which it can be separated by the dots on the mesoscutellum, and by the markings of the coxae (banded with pale in texanus, mostly black in niger) . The gonarcus and parameres of these species are shown for comparison; niger (Figures 36, 37) has a conspicuously longer mediuncus, and lacks bristles on the lateral margin of the paramere between the hook and the hinge. Calif. specimens examined : Inyo Co. : Olanche, Q , F. A. Eddy (MCZ). San Bernardino Co.: Mill Creek, 6,000 ft., VII-23-29, 8 , P. H. Timberlake (UCR) . Riverside Co. : Riverside IX-27, X-7, 8, 1948, 3 Q Q , P. H. Timberlake (UCR) ; Keen Camp, San Jacinto Mts., IX-2-1950, s (UCLA). Los Angeles Co.: IX-20-54, 8, R. X. Schick (UCLA) .
Hesperoleon fidelitas, new species