N. Banks.
A Review of the Chrysopidae (Nothochrysidae) of Central America.
Psyche 52:139-174, 1945.

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PSYCHE
A REVIEW OF THE CHRYSOPIDE
(NOTHOCHRYSIDB) OF CENTRAL AMERICA
BY NATHAN BANKS
Museum of Comparative Zoology
Many years ago the writer described a few species from Central America, including Baja California. Since then Navas has published a large number of species. For some years the author has had a manuscript synopsis of the species in the Museum. In 1937 Professor Roger Smith visited the various European museums to study the types of Chrysopidae; he has given me a copy of these notes. With this most useful help I have felt emboldened to present tables to the species known to me, with descriptions of some new forms. Besides the specimens at the Museum of Comparative Zool- ogy I have seen those belonging to the American Museum of Natural History, the Academy of Natural Sciences of Phila- delphia, and the U. S. National Museum.
As to classification, I have made an attempt to get away from dependence upon the divisory veinlet.
In the Central American forms I seeloutside of the Apo- chrysinse, three groups, one those represented by Nadiva and allies, in which the joints of the antennae are very broad, the thorax broad, the venation more or less irregular, particularly in the discoidal cell; the anal area of the hind wings is large and the branches of anal veins sometimes forked. The second group is that based on Chrysopa and allies in which the stigma1 area is unmarked, and the medius of fore wing slopes down evenly to its marginal fork. This genus should be divided. The third group is that of Nodita and Leucochrysa, in which there is a dark mark at the base of stigma, and the medius curves to Published with the aid of a grant from the Museum of Comparative Zool- ogy at Harvard College.




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140 Psyche [Sept.-Dec.
join the outer gradates; if not curving it is at least bent some- what before forking. If one has assorted many specimens of Nodita and Leucochrysa he will find specimens that arouse doubt. Specimens put in Nodita sometimes have the divisory vein ending at the upper end of cell, and Navas has described one (notha) with the divisory vein ending as in Leucochrysa. A specimen that has the divisory cell as typical of Leucochrysa as in L. vayia was the last straw. For in other structures and in coloration it agrees closely with the large species of Nodita, azvedoi, maronica, eg~eg~ia, etc., having the radial sector much curved, partly black, the costal area rather narrow, the mar- ginal forks wholly brown, and in the hind wing the marginal vein on hind margin is brown.
I am convinced that the difference between Nodita and Leu- cochrysa does not reside in the divisory cell, but in the course of the radial sector.
In Nodita the costal area at broadest is rarely equal to the radial area (at broadest), and the radial sector at widest part of the radial area is at least as near to medius as to the radius. In Leucochrysa the costal area at broadest is about equal to or broader than the radial area (at broadest), and the radial sector at broadest part of radial area is plainly nearer to radius than to the medius.
Between Nodita and Chrysopa I have depended chiefly on the dark mark in stigma for Nodita. Many species of Nodita, on drying, tend to have the pronotum collapsed transversely, a deep groove along the middle; the transverse groove being close to the hind margin; some Nodita, however, show the transverse groove near middle of length.
Those species of Nodita and of Leucochrysa perhaps in which the medius does not so plainly curve to join the outer gradates might go into another genus; but I would prefer to find for it some other character. The width of the costal stigma1 area in the true Leucochrysas is usually more than those that would be split off by this division; but the width grades so much it is not dependable. It might even be suggested to unite the large and typical Leucochrysa with the large Nodita into one genus, there is much in common, but the comparative widths of the costal and radial areas appear to keep them apart. The genera are much the same as those of northern South America, and in several cases the species; in northern Mexico



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19451 Chrysopidae of Central America 141 there are several species which occur in the southern part of the United States.
I have listed as "species" all forms that I could distinguish fairly definitely by color as well as by structure. The markings of the head and thorax and sometimes of abdomen are generally constant. There is always some variation, but it is often in extent or distinctness of development, rather than presence or absence. Specimens sometimes become discolored, and then it is difficult to determine face-marks. The study of male genitalia in this group will serve better to differentiate the species, but tends to neglect other structures which may be of value in de- fining subspecies, varieties, and races, which will surely follow more extensive studies. Taxonomy is not simply to show how species can be separated, but just as truly to show how they can be allied, - classified. Although the genitalia in many groups are the most valuable in distinguishing forms, one should remember that in Nature no species is kept separate by the structure of its genitalia.
The name Chrysopa is a pure synonym of Hemerobius, and no scrapping of rules can change that fact. But until Chrysopa can be split into a number of genera (only one small one will have to take the name Hemerobius) I shall utilize the old name. Rambur a hundred years ago provided a name for the Heme- robius of McLachlan and Hagen.
Key to Genera
1. Third cubital cell not divided; costal area very broad; no regular series of gradates; five or six branches of radius beyond subcosta; radials often connected by cross-veins . Apochysince Third cubital cell divided in some way; costal area much narrowed at stigma1 area; rarely more than three branches to radius beyond subcosta; cross-veins in middle area mostly in two or three series . . . . . . . . . Chrysopince 2 2. Antennae on basal half, at least, with the joints plainly broader than long;
a - Thorax moderately slender; antennae with basal joints rather widely separated, and slender; in male with a process between antennae . . . . . . Meleoma b -Thorax broad, basal joints of antennae stout and little separated, no process between them; in hind wing branches of anal often forked . . . . . . Nadiva



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142 Psyche [Sept .-Dec.
Joints of the antennae, except a few toward base, are plainly longer than broad, pronotum often more slender, and in hind wings the anal branches not forked . . . . . . . . 3 3. Medius of fore wing slopes down to its marginal fork, scarcely, if at all, bent; no dark spot at base of the stigma; antennae but little longer than wings . . . . . . . . 4 Medius before marginal fork usually curves up a little to run into the outer gradate series; if it does not it is at least bent at that place; nearly always a dark spot at the base of the stigma; antennae often very long; divisory vein often oblique and . . . . . . . . . . . . . . . . .
straight 8
4. Divisory vein ends on the end of third cubital cell and about parallel to both medius and cubitus . . Chrysopodes Divisory vein ends on the medius, before end of the cell, and . . . . . . . . . . . . . . .
is often curved 5
5. Inner series of gradates absent in either hind or both pairs of wings; radius usually has three or more branches to the margin beyond end of subcosta; small species with few cells and the cubital area broad . . . . . . . . . . . 6 Inner series present with at least one gradate in both wings 7 6. Wing veins and much of body greenish; no inner gradates in either wing . . . . . . . . . . . Chrysopiella Wing veins and much of body brownish; inner gradates usually present in fore wing . . . . . . . Eremochrysa 7. In male a process between the antennae, in female the basal joints of antennae more widely separate than in Chrysopa . . . . . . . . . . . . . . . . . Meleoma No process between the antennae, which are rather close to each other at base . . . . . . . . . . . Chrysopa 8. Divisory vein parallel to each side of the third cubital cell; a series of cross-veins connecting some of the radials . . . . . . . . . . . . . . . . . . Carcarulla Divisory vein oblique . . . . . . . . . . . . 9 9. A cross-vein before radial sector, usually but five cubital cross-veins beyond the divisory cell . . . . Berkmansus No cross-vein before radial sector; usually six or more cu- bitals beyond divisory cell . . . . . . . . . . . 10 10. With a more or less complete third gradate series be- tween the other two . . . . . . . . . . . . Neula No intermediate gradate series . . . . . . . . . 11



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19451 Chrysopidae of Central America 143 11. Some costal cross-veins above origin of radial sector strongly sinuous; divisory cell more or less of a V; costal area wider than radial; costal stigmal area four times as broad as the subcostal stigma1 area . . . . . . . . . . Vieira No such sinuous costals, costal stigmal area usually less broad . . . . . . . . . . . . . . . . . .12 12. Radial sector more strongly curved, so that the widest part of the radial area is nearer to the medius than to the radius; costal area usually rather narrow and not as broad as radial area; divisory vein usually ends on the medius plainly before end of cell . . . . . . . . . . . . Nodita Radial sector less curved so that at the widest part of the radial area it is as near or nearer to radius than to medius; costal area usually broader than radial area; divisory vein usually ends on the end of the cell . . . . . . . . . 13 13. Stigma with a large black spot extending back to the radial sector; costal stigmal area usually twice as broad as the subcostal stigma1 area . . . . . . . . . . Gonzaga Stigma1 mark much smaller; costal stigmal area rarely twice as broad as the subcostal stigma1 area . . . . Leucochrysa Two genera recorded from this region by Navas I have not been able to recognize in the material studied. Ancylochrysa 1928 from Costa Rica from the odd divisory vein it might be near Goliva; but he says nothing about short antenna1 joints, the radial sector is little curved and the costal area broad as in Leucochrysa; the radius has several branches to margin beyond end of the subcosta; the medius slopes evenly to its marginal fork.
Orlandsia 19 l4a from Chiriqui has the divisory vein as in Nodita, otherwise it is (according to figure) much like Ancylo- chrysa, the costal area being broad, the radial sector little curved, the medius running to its marginal fork without a bend, no mark in stigma, and the costal part of stigma much broader than the subcostal; both have a very broad post cubital area. Four forms are known from Central America. Lainius constellatus Navas 19 13, from Guatemala. Domenechus sigillatus Navas 19 13, from Guatemala. Kimmins 1940 states this is the Apochrysa mirifica Gerst.



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144 Psyche
Loyola croesus Gerst. 1893, from Chiriqui. Loyola mirifica Gerst. 1888, from Chiriqui. Kimmins 1940 puts this in Domenechus.
Table to Species of Chrysopa
1. Face with dark marks as in Ch. oculata, second joint of antennae dark, beyond pale . . . . . . . . mexicanus . . . . . . . . . . . .
No such marks . 2
2. Antennae beyond second joint black, at least for a short distance . . . . . . . . . . . . . . . . . 2 5 . . . . .
Antennae beyond second joint wholly pale 3 3. Basal joint of antennae with dark spot or stripe . . 4 Basal joint unmarked . . . . . . . . . . . . 12 4. Basal joint with 'two dark stripes, gradates parallel and near together . . . . . . . . . . . . . . a~ioles Basal joint with but one dark stripe . . . . . . . 5 5. A good-sized black spot each side on pronotum . discolor No such spots . . . . . . . . . . . . . . 6 . . . . . . . . . . . . . . 6. Palpi dark 7 Palpi pale . . . . . . . . . . . . . . . . 10 7. Inner gradates bowed up toward radial sector; branches of radial sector much bent by inner gradates .... annotaria
Inner gradates parallel to outer row; branches of radial sector scarcely bent by inner gradates . . . . . . . 8 8. Head, face, vertex, and basal antenna1 joint suffused with reddish; practically all cross-veins dark; about five inner gradates . . . . . . . . . . . . . . . dampfina Head not suffused with reddish; inner gradates often but three; third cubital cell much narrowed at base, and often but one branch to hind margin, two from fourth cell . . . . 9
9. Abdomen pale as the thorax; pronotum narrowed from . . . . . . . . . . . . .
base to the front valida
Abdomen darker than thorax; pronotum not narrowed, except near front . . . . . . . . . . . . tolteca 10. Reddish mark on cheeks, face broadly reddish; vertex red each side; gradates not parallel; pronotum with two red spots on each side before margin . . . . . . . . bates; No reddish or other mark on cheeks . . . . . . 11



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19451 Chrysopidae of Central America 145 11. Inner gradates not reduced, both series more or less . . . . . .
plainly bordered, and not parallel
infausta
Inner gradates often reduced about half, close to and nearly parallel to outer row, not bordered; radials dark only in . . . . . . . . . . . . . . .
middle caligata
12. Venation almost wholly pale greenish, gradates not dark ; divisory cell usually ends before the cross-vein above .
. 13
Venation with some cross-veins at least partly dark and the . . . . . . . . . . . . . . .
gradates dark 14
13. Cubital area more than one half as wide as the post cubital; reddish on cheeks usually extends upward by side of . . . . . . . . . . .
eye, no black streak comanche
Cubital area not quite one half as wide as the post cubital area; a black streak by side of the red on cheek . californica
14. Seven cubital cross-veins beyond the divisory cell; a dark spot on each cheek and one each side on clypeus; venation . . . . . . . . . . . . . .
largely green f acialis
. . . . . . . .
Six cubitals beyond divisory cell 15
15. Some veins behind radial sector bordered with yellowish, the gradates bordered with brown; no mark on cheeks; palpi . . . . . . . . . . . . . . . . .
pale parish!
. . . . . . . .
No veins bordered with yellowish
16
16. Palpi dark or lined; inner gradates arise before or very . . . . . . .
near the penultimate cubital cross-vein 19 Palpi pale; inner gradates arise much beyond the penultimate . . . . . . . . . . . . . .
cubital cross-vein 17
17. A reddish or dark spot at each corner of the pronotum; . . . . . . . . . .
no marks under eyes tetrasticta
Not four spots on pronotum, a mark under each eye . . 18
18. Pronotum red on sides; cubital area as broad as post cubital area; usually twelve radials, and but three or four . . . . . . . . . . . .
inner gradates brevihirta
Pronotum not red on sides; cubital area not nearly as broad as post cubital area, all cubital cells longer than broad; usually . . . . about ten radials and five inner gradates fairchildi
19. Inner gradates strongly divergent from the outer ones 20 Gradates parallel or nearly so; often a red or dark mark . . . . . . . . . . . . . . . .
under eye 21




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146 Psyche [Sept.-Dec.
20. Gradates converge behind so their bases are near each . . . . . . . . . . . . . . . .
other bouvieri
. . . . . Gradates have bases very far apart divergens
21. Pronotum as long as broad, with an interrupted red . . . . . . . . . .
stripe each side, not on margin
22
. . . . . . . . . .
Pronoturn broader than long 23
22. Pronotum much longer than broad; inner gradates nearer to radial sector than to outer, arise at or before penul- . . . . . . . . . .
timate cubital cross vein angztsta
Pronotum only a little if any longer than broad; inner gra- dates arise beyond penultimate cubital cross-vein . yucatanensis 23. Divisory cell ends at or before the cross-vein above; . . . . . . . . . . . . .
cheeks reddish exotera
Divisory cell ends well beyond the cross-vein above . . 24
24. Cheeks dark; most cross-veins dark at ends; inner gradates parallel, each veinlet far from next . . . perfecta Cheeks pale; cross-veins not dark at ends; inner gradates slightly divergent and each veinlet is close to next . . forreri 2 5. A dark mark in a curve across upper edge of clypeus; palpi marked with dark; gradates parallel . . . aztecana . . . . . . . .
No dark across on clypeus or face 2 6
26. Two dark lines on each basal antennal joint . bilineata But one dark line on basal antenna1 joint . . . . . 27 . . . . . . .
No dark line on basal antenna1 joint 31
2 7. Black spot on cheeks; palpi pale . . . . . . 28 . . . .
No dark on cheeks; pronoturn reddish on sides 29 28. Radial area not as wide as postcubital area; many cross- . . . . . . . . . . . . . .
veins dark gradata
Radial area about as broad as the postcubital; posterior half of wing with mostly pale veins (except gradates) . indicata 29. Inner gradates near to radial sector, gradates not quite . . . . . . .
parallel; many costals wholly dark sarta Inner gradates close to outer row and parallel thereto . 30
30. Some gradates and some branches of radial sector bor- dered; pronotum about twice as broad as long . . berlandi No gradates nor other veins bordered; pronotum only a little broader behind than long, narrowed toward front . . adoina 31. Palpi partly dark; cubital area as broad as the costal



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1945 1 Chrysopidae of Central America 147 area; costals and radials wholly dark; pronotum not margined . . . . . . . . . . . . . . . .
with reddish 32
Palpi not marked with dark, costals and radials less dark; . . . . . . . . . . . . . .
wings rather slender 33
32. Pronotum with a pair of submedian dark stripe . incerta . . . . . . .
Pronotum without any stripes leptana
. . . . . . . .
33. Margin of pronotum reddish 34
. . . . . . . .
Margin of pronotum not reddish
35
34. Cheeks with reddish mark; basal joint of antennas red- . . . . . . . . . . . . . . . .
dish lateralis
. . . . . . . . . . .
Cheeks not marked clave~i
35. Inner and outer gradates very close to each other, and . . . . . . . . . . . . . . . .
parallel everes
Gradates widely separated, inner sloping upward, and not . . . . . . . . . . . . . . .
parallel everinu
Species that I have not recognized in the material and could not place from the descriptions and notes, some of which are probably synonyms.
Chrysopa sanguine0 Navas 1927, near to caligata, but it does not agree in various points.
Chrysopa rubricosa Navas 1914, the figure shows marks on head that I have not seen in any species. Chrysopa effusa Navas 19 1 1, may possibly be Ch. gradata ; the latter has some of the branches of cubitus as Navas figures for effusa, but the divisory cell is not dark. Chrysopa guatemalteca Navas 1914 is very near Ch. sarta Bks. There are fewer gradates in sarta than stated for guate- malteca, and sarta has no dark in hind wing, and antenna1 stripe broad. If the same, guatemalteca has a month or more priority.
Chrysopa bouvieri Navas 192 3 and Ch. divergens Navas 193 1 according to description and Smith's notes must be very similar if not identical; those we have agree possibly a little better with divergens.
Chrysopa bulbosa Navas 1926 will run in the key to Ch. in- fausta; however, there is nothing unusual about the basal joint of antennae in infausta; and infausta has the stripe on basal joint lateral, not dorsal, the pronotum not margined, etc.



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148 Psyche Lbept .-uec.
Chrysopa cajensis Navas 1930 in table goes to yucatanensis and I consider it the same.
Chrysopa hieronyma Navas 191 7 seems to be tetrasticta. Chrysopa obesa Navas 1929. A broad bodied species with wholly green wings; may be a Nadiva but not IV. balboana. Chrysopa senior Navas 1927, wing 20 mm. long, is different from any I have.
Chrysopa morrisoni Navas 1914, veins all green, has dark dots or lines on vertex or pronotum, and thus readily separated from the all green veined species known to me. Chrysopa varicosa Navas 1914 is stated to have a dark lunule below each antenna, and peculiar modifications of certain vein- lets of the inner gradates; I have seen similar modifications in one species, but that species has no lunules on face, nor does it agree otherwise with the description of Navas, so it is probably also a distinct species.
Chrysopa lafoni Navas 1911, and Ch. nativa Navas 191 1, both from Costa Rica and fore wings 17 mm. long, do not fit anything I have seen.
Chrysopa sulcata, Ch. salleana, Ch. ceratica of Navas and Ch. explorata Hagen belong to Nodita; Ch. dolicharthra Navas is a Meleoma.
Chysopa externa var. marginata Navas 192 7, from Guate- mala, is described in one line and a half. The pronotum is marked on the anterior middle with dark red; I have not seen it. Chrysopa josephina Navas 1926 is said to have twelve joints of antennae black, and outer gradates bordered; it agrees fairly well with what I have as berlandi, but here the gradates are not bordered, but this is not always dependable. Chrysopa josephma Navas 1930, appears to be Ch. angusta Navas.
Chrysopa longicella Navas 1914; I identify this with Ch. bimaczdata McClendon, described from southern Texas; I have specimens from Guatemala, Nicaragua, and Canal Zone; except for minor details and size there is little to separate this from tolteca Bks. and so I have united them. Ch. valida from Baja