Wm. T. M. Forbes.
Pectinate Antennae in the Geometridae.
Psyche 32(2):106-112, 1925.

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106 Psyche [April
PECTINATE ANTENNA IN THE GEOMETRIDA
(LEPIDOPTERA).
BY WM. T. M. FORBES,
Cornell University, Ithaca, N. Y.
The present note is the summary of an analysis of antenna1 characters in the Ennomid Geonietridse, for which there seems to be no present hope of publication.
The Geometridae and especially the Ennomid (Boarmine) series, have always been notorious for the plasticity of their characters and the difficulty of their classification. The present study of antennal characters has developed out of Bodine's work (Trans. Am. Ent. Soc. 23, 1, 1896) on the Lepidopterous antenna, and has, I believe brought some significant and useful characters to light.
This discussion includes only the genera with pectinate male antennae. In the Geometridse the simple antenna is not primitive, but has been several times produced by reduction. For the present it is not possible to place all these genera, but pupal and other characters will throw light on their relationships. Bodine recognizes five types of antennal organs : three lengths of setae, cones, and pit-organs. In the Ennomids the pit-organs seem of little use in classification, and the two types of generally distributed setae (his types 1 and 3) are not easily distinguished; the presence and arrangement of the long single setae which he calls type 2, and the thin-walled sensory cones- type 5,-give good characters.
The following grouping may be made:
There are three subdivisions of this type, corresponding to two natural groups, and a somewhat heterogeneous remainder. 1. Pectinations each with a single distinct seta of type 2, not obscured by long set% of type 3; pupa with two strong spines on the cremaster, with dorsal groove (between abdominal seg- ments 9 and 10) well developed; moth slender, wing normally with fovea. Antenna usually with a long simple apex.



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19251 Pectinate Antinnce in the Geometridce (~e~ido~tera) 107 Protoboarmia (indicataria), Parexcelsa (inconspicuaria), Eu- fidonia, Neoalcis (californiaria), Melanolophia, Vinemina (opa- caria), Paraphia, Epimecis, Elphos, Boarmia, (e. g. rhomboidaria), Amraica. In Amraica the antenna is unipectinate, with a super- ficial likeness to Arichanna. The others form a homogeneous group.
2.
Pectinations very short, gradually running out to base and apex, slender, central on segments, not clubbed, with long sparse bristles, the apical one not distinct; segments lightly chitinized, with fine striations, transverse on outer part of seg- ments, as in Melanolophia.
Bapta (?) virginalis.
The remaining Baptas have simple
antennae, and resemble Cabera (Deilinea). This species has no likeness to Cabera and has no visible connection with any other. 3.
Pectinations more or less clubbed, with a terminal tuft of long curved setae, burying the single short apical or sub-apical seta of type 2. Body stout, mouth parts normally reduced; sculpture normally not unlike group 1, which also has a similar Pupa.
Biston (ursaria), Amphidasys (cognataria, robusta), Erannis (segments sometimes 4-pectinate, sometimes merely serrate), A rtiora (Therapis), Coniodes, Cochisea. This is a homogeneous group. Artiora has usually been widely separated from Erannis but hardly differs save in wing-form.
4.
Pectinations with at least two strong setae of type 2, which are typically apical and strongly divergent; and are fre- quently supplemented by one or more such setae on the outer sides of the pectinations. A miscellaneous group, held together mainly by the survival of a primitive condition. Prosopolopha.
Euchlama (including irraria), Lytrosis, Stenotrachelys, An- gerona (prunaria, cexaria), Xanthotype, Metarrhanthis, Cepphis (Priocycla).




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Psyche [April
Gonodontis (bidentata, formosa, ocellaria), Abbotana, Himera, Campcea (Metrocam pa), Ellopia (not Therina) , Selenia, Hygro- chroa, Stenaspilates, Hypoplectis, Scodiona, Hulstina, Amblychia (angeronaria), Xandrames (dholiaria), Arichanna (transitional to Melanolophia group), Sarcinodes (unipectinate) , Achlora (4- pectinate).
This series like the first is composed of three main groups, of which two are more like each other than to the third, and has some anomalous genera of uncertain position. Nacophora makes a group by itself, much like the Biston group, but differ- ing in egg-type as well as ant'enna.
1. Basal segment of antenna1 shaft with well-marked pec- tinations; posterior series of longer pectinations than anterior, especially toward apex; cones very few, sometimes irregular in position and deformed. Apical setae of pectinations strong, strictly apical, and not obscured by set= of type 3; sub-apical seta strong, often arising well back from apex. Pectinations stout and long, tapering, with tips turned dist,ad; strongly chitinized, especially at base, and longitudinally striate; shaft smooth or slightly granular.
Nacophora (including ((Amphidaqs" arnobia) , Phceoura. 2. Cones numerous, normally on both series of pectinations; apical setse usually two, and well-marked. Tropical species fre- quently very heavily chitinized. American genera. Therina (including quercivoraria and lceta), Nepytia, Zerene, Sicya, Philtrcea, Philedia, Nipteria (in part), Lissochares (ni- grovenata), Deutophlebia (radiata), Emplocia (bupaloides), Leucula (cillenaria) , Carpella (districta) , Sanqalopsis (beata) ,-also "Diop- tis" hesperioides, for which I have not happened to see a valid generic name,-Metanema (inatomaria and determinata, transi- tional to group IIc).
3.
Cones numerous on one series of pectinations, normally the posterior, usually absent on the other series, but present



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19251 Pectinate Antennae in the Geometridce (Lepidoptera) 109 on both series on Bupalus, Crocallis and Euctenurapteryx. Old-world genera.
Chemerinu, Cleogene, Dusydia, Epione, Bupalus, Euctenura- pteryx (maculicaudaria), Crocallis. The last genus on most characters is an outlier of series I.
This is far the smallest, and in antennal structure the most homogeneous of the four series, but the genitalic structure as well as the appearance seems to indicate it is not homogeneous. Gnophos is particularly troublesome, as with a great variety of antennal type the genitalia a.re homogeneous, while Platsea, which has practically the antenna of G. dilucidaria, has wholly different genitalia.
Bases of pectinations conspicuously transversely rugose, and often swollen.
Pectinations basal on segments.
Over 40 segments. ............... Gnophos, Putceu, Pherne Under 40 segments. ................. Glaucinu, Coenocharis Pectinations subapical.
Segments under 40, pectinations short. Carphoides, Burnesia Segments over 40, pectinations long. ............ Pterotcea Bases of pectinations smooth or nearly so. Antenna pectinate to apex.
Lightly chitinized; segments stout. .............. Ennomos .......
Heavily chitinized; segments very slender. Aspilates Antenna wit'h a long simple apex. ... (a few aberrant Nipterias) SERIES IV: PECTINATIONS SCALED, CONES BASAL. There are two main types involved in this group, one rep- resented by Cabera and Apicia (Caberodes), in which the cre- master of the pupa has eight hooks, and the other by the Cleora group with a bifid cremaster. There is a corresponding difference in the appearance of the moth and of the antennae, and the groups are no doubt natural, although at present impossible to define. Ametris, commonly put with the CEnochrominse, seems to belong



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110 Psyche [April
here, while the true (Enochromids fall in series I; Cleora and Ripula have doubly pectinate antennae, the pectinations al- ternately scaled and naked. Cleora may be ancestral both to the Cleora group of this series and the Melanolophia group of the first, the inner pectinations having disappeared in one case and the out,er in the other. Ripula has no obvious affinities. The diurnal forms Epelis and Ematurga seem to be separately derived from somet,hing near Itan~e, and should not be combined in a single genus, as has been commonly done, following Hulst in Dyar's list. The following grouping is largely artificial and for the convenient tabulating of some characters only. Synopsis of groups.
Pectinations basal
Sculpture strong on pectinations, setae of type 2 apical, not strong. .................................. Group A Sculpture weak on pectinations; setae subapical and obsoles- cent, lost among the set= of type 3. .......... Group B Pectinations central or apical
Strongest seta of type 2 subapical, short and weak as a rule, when near the apex lost among a mass of longer set* of type 3.
Subapical seta strong, comparable with the apical. Group C Subapical seta less than half as long as the apical and weak, .............................
but distinct. Group D
Subapical seta not recognizable, lost in t7he setae of type 3 or (more probably) absent. ................ .Group E Pectinations ending in a strong and conspicuous apical seta No subapical or dorsal seta, only the apical type-2 seta present. .................................. Group F Subapical seta well-marked, no dorsal seta Pectinations apical. ......................... Group G Pectinations central. ........................ Group H Subapical seta strong; a few, at least, of anterior pectinat- ions with a third dorsal seta at about a third their lengt,h. ................................... Group I



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19251 Pectinate Antennae in the Geonzetridae (~e~idoptera) 11 1 Pectinations with apical seta more or less distinct, but not fully apical, at least on some segments; subapical strong; sculpture of shaft weaker than in group D Group J
A: Selidosema (ericetaria, ambustaria but not the American species sometimes included).
B : Lychnosea
(helviolaria only), L. (?"Egperitis") triang- uliferaria, Stegania (trimaculata), Ripula. C : Caripeta, ~emero~hila (abruptaria).
A wholly artificial
pairing, I suspect. Caripeta has a pupa similar to Cabera and Apicia, Hemerophila is much more suggestive in all ways of Lytrosis in series I.
D : Vitrinella (pampinaria) .
E: Itame (Diastichtis, Cymatophora), Physostegania, Elpiste, Macaria (species with more or less pectinate antennae), Mericisca, Buzura (suppressaria), Eubolia, Enconista, Epelis, Exelis, Eu- macaria, Parapheromia (lichenaria), Tracheops, Merisme (spodo- dea) , Fidonia (limbaria), Euaspilates, The South American 'Aids" salmonearia, and "Ectropis" anaisaria also belong to this group, but not the genera in which they now stand. F : Glena (insaria, quinquelinearia), Anicainpto'iss, Emitur- ga (faxonii, atomaria), Hyposidra (talara) , Gynopteryx (ssriaria) , Hymenomima (tharpa) , A metris.
G: Pseudoboarmia (umbrosaria, punctinalis), Stenoporpia (polygrammaria), Tornos, Hesperumia, Chloraspilates, Somitolo- phia, Halesa (cenitusalis), Erebomorpha, Aplogompha (rioj,G), Molybdogompha (biseriata,) " Lychnosea" in!ernzicata, Neo'erpss, Epiplatymetra.
H : Ixala, Pterospoda, Enemera.
I: Cabera, Drepanulatrix, Catopyrrha, Apicia (Caberoies, without C. irraria).
J: Plagodis, Anagoga, Hyperitis.
This is a homogeneous
group on venation as well as antenna.
In- the following genera I have been unable t'o find any trace of cones, or at most a few on the simple terminal segments, so that it is impossible to group them unanlbiguously. I divide them into artificial groups to call attention to some of their characters.




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112 Psyche [April
GROUP I: PECTINATIONS NAKED, BASAL ON SEGMENTS, APICAL SETAE NORMALLY DISTINCT
Heliothea, Egea (no terminal setae) , Nychiodes, Eurrhanthis, GROUP 11: PECTINATIONS NAKED, APICAL, NO SETAE OF TYPE 2. Acalia, Brephos
Nepterotcea (compare Ccenocharis and ~n-ophos), Eucaterua (compare IV B or IV G) Narraga (compare Epelis), Fernaldella (very near Narraga) , Melanchroia (Compare IV F) . STERRHIM and HYDRIOMENINM
Pectinations slender, basal, naked with stiff, sparse and rat,her evenly distributed setae, and a minute apical seta of type 2 or none. Cones rare on the pectinate segments; on the shaft in Xanthorhoe ferrugata,, rudimentary on apices of pectinations in the Sterrhince.
Pectinations naked, with cones usually on the simple apical segments only, but sometimes also on the apices of some pectin- ations.




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