E. L. Mockford.
Genus Eremopsocus McLachlan: Distinction from Cerastipsocus Kolbe and Review of Species (Psocoptera: Psocidae).
Psyche 82(2):244-258, 1975.

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GENUS EREMOPSOCUS McLACHLAN: DISTINCTION FROM CERASTIPSOCUS KOLBE AND REVIEW OF
SPECIES ( PSOCOPTERA : PSOCIDAE) *
BY EDWARD L. MOCKFORD
Department of Biological Sciences
Illinois State University
Normal, Illinois 61 761
During investigation on the systematics of Cerastipsocus and its close relatives, I have experienced difficulty in assigning species be- tween the two genera Erew~opsocus and Cerastipsocus. In this paper, I attempt an unequivocal definition of these genera. I also include diagnoses of the species placed by the new definition in Eremopsocus, and a reconsideration of geographic variation in the type species, E. infumatus McLachlan. The subgenera of Eremopsocus are also re- evaluated. One new species is described. Measurements (Table 11) were made on whole specimens either pinned or temporarily mounted in glycerine. They are in mm and have an error of å 1.04 microns. Abbreviations used with the mea- surements are explained as follows: fi, fa = lengths of first and second flagcllomeres; Fw =Z forewing length; F = length of pos- terior femur; T = length of posterior tibia; ti, ts length of first and second posterior tarsomeres; tict == number of ctenidia on posterior first tarsomeres; IO/D =: least distance between com- pound eyes in dorsal view divided by greatest antero-posterior di- ameter of eye in dorsal vkw; PO -= greatest transverse diameter of compound eye in dorsal view divided by greatest antero-posterior diameter of the eye in same view.
Characters of Eremopsocus and distinction from Cerastipsocus. - Eremopsocus McLachlan (1866) was erected for a single large Brazilian species, E. infumatus, with fuscous wing membranes. This species was reported to have incrassate flagellomeres in the male only, those of the female being slender. This antenna1 character, alone, was held to separate the genus from Psocus, which was then used in a very broad sense.
Pearman (1933) correctly aligned Eremopsocus with its close relatives by placing it in subfamily Cerastipsocinae of Family Psoddae. Pearman's material, from Venezuela, showed incrassate flagellomeres in both sexes, and on that account, he designated a distinct subspecies, E. infurnatus venezuelensis. *Manuscript received by the editor August 15, 1975. 244
Ps~rhe 82:244-238 t 1975). http:l/psycb cnlclub ore/SWI-W html



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19751 Mock ford - Erem<opsocus 245
I have examined 15 adult specimens of this species from two localities in central Brazil, ;and I find that all individuals of both sexes have antennae incrassate. I believe that McLachlan's observa- tion was incorrect and that the error arose from his having had more than one large, fuscous-winged Cerastipsocine species in his material. There are several such species in Brazil. Those with slender flagellomeres in both sexes are currently assigned to Cera- stipsocus. The argument for McLachIan's having had a mixture of species is strengthened by another of his observations: that mlales have wings shining, while females have them dull. Pinned specimens in the series of E. infunzatus which I studied have the wings shining in both sexes while pinned specimens of
Cerastipsocus fuscipennis
(Burmeister) received in the same shipment from a nearby locality in Brazil have the wings dull. I conclude that the flagellomeres of both sexes of E. infumatus are incrassate and that Pearman's Vene- zuelan subspecies has no validity.
Pearman (1933) described the genitalia of both sexes of E. in- fumatus. The hypandrium is symmetrical and bears an apically rounded lobe in the middle of the disc; distally, it belars a pair of blunt, posteriorly-directed prongs. This hypandrium differs from that of the type of Cerastipsocus (C. fuscipennis [Burmeister], designated by Roesler [1944], not C. venosus [Burmeister], erron- eously designated by Smithers [1967] ).
In C. fuscipennis, there is no trace of a rounded lobe in the middle of the disc and distal prongs are absent. This type of hypandrium has been illustrated for C. cubanus Enderlein (Mockford,, 1974: 164, Fig. I 18) and C. venoms Burmeister (Chapm'an, 1930, pi. XIII, Fig. 9). C. heaveri New (1972: 207, Fig. 22) is much' the same but its hypandrium has a pair of short, laterally directed distal prongs.
The distinctive hypandria of the types of Eremopsocus and Cera- stipsocus provide useful character complexes for distinguishing be- tween these two genera. Unfortun~ately, these complexes do not correlate with the distribution of incrassate and slender flagellomeres in the species as currently assigned in these two genera. Thus E. reductus (Banks) has the flagellomeres developed as in E. infumatus but the hypandrium as in C. venosus, completely lacking a lobe in the middle of the disc and without distal prongs (personal observa- tion). C. crassicornis (Kolbe) in which the male has only the first flagellomere incrassate and the flagellomeres of the female are slender has the hypandrium developed as in E. infumatus (personal obser- vation).




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19751 Mockford -- Erem~opsocus 247
It is my opinion that the hypandrium in these insects presents more information than the relative widths of flagellomeres and that the two genera can best be recognized on the basis of type of hypan- drium. This opinion is strengthened by the fact that no intermediate type of hypandrium has been found, whereas flagellomeres of different species show varying degrees of incrassation. In order to test this opinion further I have searched for additional characters showing essentially two states in the assemblage of species under consideration and noted how their states are distributed. The following characters were found :
(I) shape of posterior clunial margin of male in region of base of epiproct: slightly protruding and bilobed (Fig. 8) or decidedly protruding and not 'at all bilobed (Fig. 5) ; (2) presence (Fig. 5) or absence of a protuberance immediately distad of sense cushion on paraproct in male; (3) relative length of stem of dark T-shaped mark of female subgenital plate (stem of vase-shaped figure extending into distal process of plate) : either more than twice as long as broad or scarcely longer than broad to much shorter than broad. The following species were examined (assigned to genera accord- ing to current usage) : Cerastipsocus bogotanus (Kolbe), C. crassi- cornis (Kolbe) , C. cubanus Enderlein, C. fuscipennis (Burmeister) , C. ~ochraceocristatus Enderlein, C. siworii Ribaga, C. trifasciatus (Provancher), C. venosus (Burrneister), two undetermined species of Cerastipsocus, Ermopsocus infwnatus McLachlan, and E. re- ductus (Banks).
The data are summarized in Table I. From this table, the fol- lowing correlations are seen:
I. Incrassate flagellomeres correlate with hypandrial type of E. infumattis. The notable exception is E. reductus, while C. cubanus shows slight incrassation.
Fig. 1. Eremofsocz~s crassicornis (Kolbe) 8, forewing, scales in mm. Fig. 2. E. crassicornis (Kolbe) 8, hindwing. Fig. 3. E. crassicornis (Kolbe) 8, hypandrium. Fig. 4. E. cras~icornis (Kolbe) 8, phallosome. Fig. 5. E. crassicornis (Kolbe) 3, epiproct, left paraproct, and adjacent clunial margin, scale of Fig. 3. Fig. 6. E. crassicornis (Kolbe) 8, antenna (scape to base of fg), scale of Fig. 1. Fig. 7. E. crassicornis (Kolbe) $2, antenna (scape to base of fq), scale of Fig. 1. Fig. 8. Cerastipsocus venosus (Burmeister) 3, epiproct, base of right paraproct, and adjacent clunial margin.




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19751 Mockford - Erenz~opsocus 249
2. 7
the E.
m'argin
'he protruding male clunial margin tends to correlate with infumatus hypandrial type, while the bilobed male clunial tends to correlate with the C. venosus hypandrial type. This correlation is better than indicated on the table. In E. infumatus, C. crassicomis,, and C. sp. no. I, the protrusion is a thickened lip, while in C. cubanus and E. reductus it is only a thin rim, as in those species with the bilobed margin.
3. The protuberance distad of the male paraproctal sense cushion tends to correlate with the E. infumatus hypandrial type and absence of the protuberance with rhe C. venosus hypandrial type. Again, the correlation seems better than the table suggests, as the protuber- ance in E. infumatus, C. crassicornis, and C. sp. no. I is small and decidedly pointed, while that in C. bogotanus is low, larger, and rounded.
4. The long stem of the T-shaped m'a,rk of the female subgenital plate shows con~plete correlation with the E. infumatus hypandrial type and the shorter stem with the C. venosus hypandrial type. From the above correlations, I conclude that the genera Eremop- socus and Cerastipsocus may be differentially defined as follows: Eremopsocus. - Antennae incrassate at least in male ; hypandrium with central lobe of disc; male clunial margin protruding above epiproctal base as a rounded, thickened lip; a pointed protuberance distad of male paraproctal sense cushion; stem of T-shaped mark of female subgenital plate more than twice as long as broad. Cerastipsocus. - Antennae incrassate or not ; hypandrium lacking central lobe of disc; male clunial margin bilobed before base of epiproct or slightly protruding above epiproctal base but never de- veloped as a decidedly protruding, thickened lip ; generally lacking a protuberance distad of male paraproctal sense cushion, or with a low, rounded protuberance; stem of T-shaped mark of female sub- genital plate little longer than broad to much shorter than broad. Assignment of species to Cerastipsocus and Eremopsocus. - I can affirm, either through examination of specimens or by existing pub- lished descriptions, the generic assignment of the following species according to the 'above definitions:
Cerastipsocus. - C. f uscipennis (Burmeister ) (type species) , C. beaveri New, C. bogotanus (Kolbe), C. cubanus Enderlein, C. ochra- ceocristatus (Enderlein) , C. reductus (Banks), new combination, C. si~~orii (Ribaga), new combination, C. trifasciatus (Provancher), C. venosus ( Burmeister ) .




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19751 Mockford - Eren~~opsocus 25 1
Eremopsocus. - E. infumatus McLachIan (type species), E. cras- sicornis (Kolbe) new combin'ation, E. nigripes n. sp. (heretofore referred to in this paper as Cerastipsocus sp. no. I). Several species traditionally assigned to Cerastipsocus are unassign- able by the above definition, and I hope to review that genus and assign these species in the near future. Subgenera of Eremopsocus. - Roesler ( 1944) assigned Syngono- soma Kolbe, Din~opsocus Banks, and Podopterocus Banks as sub- genera of Eremopsocus. These assignments were res sum ably made on the basis of incrassate flagellomeres alone. As I have shown that this character is not a reliable guide to relationships, it seems advisable to retain the old-world genera Dinopsocus land Podopterocus at least until their external genitalia are known. The type of Syngonosoma being South American, it seems likely that this genus may be a synonym (not a subgenus) of Eremopsocus as was suggested by Enderlein ( I g r I ) and Pearman ( I 933), but is will be necessary to examine material of the type species in order to confirm this SUE- gestion. Of the species listed in Eremopsocus by Smithers ( 1967), only the type species can be retained with certainty. Systematics of Eremopsocus species. - In this section the species now assigned to Eremopsocus are diagnosed, the description of E. infumatus is augmented, a detailed description of E. crassicornis is presented and E. nigripes n. sp. is described. A key to the species is included.
Eremopsocus infumatus McLachlan
E. infumatus McLachlan, I866 : 348.
E. infumatus venezuelensis Pearman 1933 : 159. Diagnosis. - First four flagellomeres incrmassate in both sexes, the basal two decidedly so, the next two somewhat less. Female sub- genital plate with a single row of long setae along cross-piece of T-shaped mark (the row double to the sides and single in middle on specimen examined).
Fig. 9. Eremopsocus crass'icornis (Kolbe) 9, epiproct. Fig. 10. E. nigripes n. sp. $2, epiproct. Fig. 11. E. crassicornis (Kolbe) 9, subgenital plate, scale of Fig. 12. Fig. 12. E. nigripes n. sp. ?, subgenital plate. Fig. 13. E. crassicorn'is (Kolbe) 9, gonapophyses. Fig. 14. E. nigripes n. sp. 9, gonapophyses, scale of Fig. 9. Fig. 15. E. infumatus McLachIan 9, epiproct, scale of Fig. 9. Fig. 16. E. crasslcornis (Kolbe) Q, sperma- pore sclerite. Fig. 17. E. nigripes n. sp. 9, spermapore sclerite, scale of Fig. 16.




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252 Psyche [June
Pearman's description ( I 933 ) is augmented as follows : Measure- ments (Tlable 11). Lateral prongs of hypandrium joined along their length to sides of hypandrium by semimembranous cuticle. Female epiproct (Fig. I 5 ) with postero-lateral sclerotized strips turning slightly outward at their distal ends. Material examined. - Brazil : Goias : Jatai, October, 1972 I Q F. M. Oliveira; 20 Km. north of Sao Joao da Alianca, April 14, 1956,
13 Q, F. S. Truxal.
Eremop~ocus crassicornis ( Kolbe) , new combination Cerastis crassicornis Kolbe, 1883 : 70.
Cerastipsocus crassicornis (Kolbe), Smithers, 1967 : 96. Diagnosis. - See diagnosis of E. nigripes n. sp. Male. - Measurements (Table 11).
Morphology. - Antenna with basal flagellomere (Fig. 6) decid-
edly incrassate, second less so, third slightly incrass'ate basally; all flagellomeres densely beset with upright hairs. Compound eyes small (see Table 11). Hypandrium (Fig. 3) with pair of short distal prongs; these and distal ends of lateral thickenings beset with minute denticles. Discal lobe and regions to its sides and base bearing setae. Phallosome (Fig;. 4) a closed ring with truncate distal process, the process and weakly-sclerotized region to its sides bearing minute denticles. Paraproct (Fig. 5) with a long distal spur and a short spur distad of sense cushion. Epiproct and posterior clunial margin as in Fig. 5.
Color (in alcohol). -Compound eyes and inner rims of ocelli black. Head deep orange-brown with faint clypeal striations. Maxil- lary palpi dark brown. Antenna1 scape and pedicel dark chestnut brown; flagellum black. Mesonotal lobes deep to medium chestnut brown,
their borders and thoracic pleura orange-brown. Forewing membrane (Fig. I ) uniformly fumose-brown except colorless in base of areola postica and nearly so in basal region of cell RI from behind stigmasac nearly to apex of pterostigma. Pterostigma brown Fig. 18. Eremopsocus nigripes n. sp. $, forewing. Fig. 19. E. nigripes n. sp. 3, hindwing. Fig. 20. E. nigripes n. sp. 8, hypandrium. Fig. 21. E. nigripes n. sp. 8, phallosome. Fig. 22. E. nigripes n. sp. 8, epiproct, left paraproct, and adjacent clunial margin, scale of Fig. 20. Fig. 23. E. nigripes n. sp. 8, antenna (scape through base of f̣) scale of Fig. 18. Fig. 24. E. nigripes n. sp. 9, antenna (scape through base of fn), scale of Fig. 18.




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254 Psyche [June
outlined in white. Veins of forewing white except following brown: short segment of Rs before its fork, short segments of M before M-Cuia junction, R2 + 3, except its extreme base, R4 + 5 in its distal two-thirds, MI, M2, M3, Cuia beyond junction with M. Hindwings (Fig. 2) uniformly pale fumose-brown. Coxae, tro- chanters and femora orange-brown ; tibiae and tarsi duskier. Mem- branous portions of abdomen ringed with slender white stripes on a purplish-brown background (subcuticular pigment). Terminal abdominal segments medium to dark brown. Female. - Measurements (Table 11).
Morphology. - Flagellomeres slender (Fig. 7). Subgenital plate (Fig. I I ) with pigmented area roughly T-shaped 'as in E. infumatus but with narrow region of stem of T shorter and setae distad of cross-piece shorter and not forming a distinct row, being interrupted in middle. Gonapophyses (Fig. 13) with first valvula acuminate distally, bearing a few minute spines on its edges; second valvula tapering distally and spinulose over distal third; third valvula with basal, more sclerotized region bearing setae (this region shown folded over rest in figure), remainder membranous, distal lobe straight. Ninth sternum and spermapore as in Fig. 16. Epiproct (Fig. 9) with clear region surrounded by approximately quadrate frame of heavier sclerotization, the frame and clear area bearing setae of various lengths.
Color. - Same as in male.
Miterial examined. - Brazil : Prov. Minas Gerais : Sao Joao del Rei, Coll. Sello, I
8 (holotype) ; Santa Catarina: Nova Teu- tonia, November, 1970, Coll. F. Plaumann, 14 8, 23 9. Uruguay: Depto. de Treinta y Tres: Rio Olimar Chico, 25 Km. WSW Treinta y Tres, I I April 1963, Coll. J. K. Bouseman, 8 8, 2 $. Eremopsocus nigripes, new species
Diagnosis. - Differing from E. infumatus in having only the first two flagellomeres incrassate and only in the male. Differing from E. crassicornis in having first two male flagellomeres decidedly more incrassate and having in both sexes posterior vertex of ptero- stigma angular, forming approximately a right angle, rather than rounded.
Differing from the described fuscous-winged neotropical Cerastipsocus species in which external genitalia not known: from C. ocularis (Kolbe) , C. moestus (Kolbe) , C. pa1lidinervi.s (Kolbe) , C. vetustus (Kolbe) in having pterostigma deep and with decidedly angular posterior vertex.
Male. - Measurements (Table 11) .




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Mockford - Erem~opsocus 255
1 - 1 m C O 1 - 1 ' - i C O C O C O i n I D m 0 ( 0 1 0 t - ? . ? ? ? ( ^ c o t - ?
0 0 0 0 0 0 0 0 0 0 0
Q I D O @ ~ " - l @ @ I D " - l l n o '
\in 1-1 o o N N ? 0 1 ? N
L i ; ; ; ; ( 0 ( 0 K l K l m m




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Morphology. - Basal flagellomere ( Fig. 23 ) more incrassate than in E. crassicornis, second about the same, third to tip slender. All flagellomeres beset with upright hairs. Compound eyes small (see Table 11). Hypandrium (Fig. 20) with distal prongs somewhat longer than in E. crassicornis; these and sides of hypandrium, to extreme base
beset with minute denticles. Discal lobe and regions to its sides and base bearing setae. Phallosome (Fig. 21 ) symmetrical, with somewhat flattened base; the inset, spatulate distal process larger thlan in E. crassicornis and bearing minute denticles. Para-
proct (Fig. 22) as in E. crassicornis. Epiproct and posterior clunial margin as in Fig. 22. The latter differing from that of E. crassicornis in having sides slightly indented.
Color (in alcohol). - Compound eyes and inner rims of ocelli black. Head yellow deepening to burnt orange on vertex with a spot of medium orange-brown in front of ocellar interval; clypeal striations faintly indicated by orange-brown mottling. Maxillary palpi pale brown on basal two segments, becoming darker distally on second; third segment dark brown, distal segment black. Antenna1 scape pale brown, rest of antenna black. Mesonotal lobes orange peripherally, deep orange-brown in a broad central band running length of mesonotum. Thoracic pleura violet. Forewing membrane (Fig. 18) uniformly fumose-brown except small region in base of areola postica extending distally along vein Cuia; pterostigma and narrow stigmasaum white mottled in base with fumose brown. Veins of foi ewing furnose brown except following : R I colorless from posterior apex of pterostigma to wing margin; Rs pale brown at and immediately before and in its junction with Cuia, colorless im- mediately beyond junction ; Cu I a colorless before and immediately beyond junction with M; Cuib colorless. Hindwing (Fig. 19) uni- formly pale fumose brown. Coxae, trochanters, and femora yellow; tibiae and tarsi black. Membranous portions of abdomen (note variation below) longitudinally striped with purple bands: a broad band along each side including spiracles, a narrower band along dorsal mid-line; an incomplete band ventrally from hypandrium, widest in 7th segment, narrowing to its anterior end in third seg- ment; abdomen creamy white between purple bands. Terminal abdominal segments largely dark brown, paler on poorly sclerotized portion.
Female. - Measurements (Table 11).
Morphology. - Flagellomeres slender (Fig. 24). Subgenital plate (Fig. 12) with narrow revion of stem of T-shaped pigmented area shorter than in E. infumatus; setae distad of cross-piece of T



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19751 Mockf ord - Erem~opsocus 257
shorter and forming a continuous subquadrate region. Lamp-globe shaped pigmented region distad of narrow portion of T-stem not as broad as in E. crassicornis. Gonapophyses (Fig. 14) with second valvula not as long from bro4adest region to tip as in E. crassicornis; inner lobe of third valvula relatively longer. Epiproct (Fig. 10) with heavily sclerotized sides of distal clear area not decidedly point- ing rnedially at their distal ends. Spermapore plate (Fig. 17) much as in E. crassioornis.
Color (in alcohol). -As in male except no spot of medium orange-brown before ocelli and three faint purple lines radiating from each compound eye medially.
Variation. -- Some individuals lack the longitudinal purple bands of the abdomen but have, instead, transverse purple bands, one per segment, each band dividing into two at the spiracle on each side and continuing ventrally 'as two narrow bands which dissipate before reaching the ventral miff-line. These individuals have the dark pig- mentation of the mesonotum more diffuse and, in males, the dark pigmentation of the pterostigma and stigmasaum more dispersed. I can detect no mor~hological difference between these and the form described above. The two forms were 'apparently collected together. Holotvpe 8, allotype Q, .? 8 paratypes and 2 $ paratvpes, Brazil: Santa Catarina: Nova Teutonia, November, 1970, Coll. I?. Plau- maim. Types will be deposited in the United States National Mu- seuin, Washington, D. C.
Other material examined. - (all from Nova Teutonia, Santa Catarina, Brazil, Fritz Plaumann collector). Same data as holotype, o\ I 3 9, October, 197 1, 4 d" , 4 Q; December, 1971, 2 8, 1 9. Key to the Species of Eremopsocus
First four flagellomeres incr,assate in both sexes. Female sub- genital plate with a single row of long setae along cross-piece of T-shaped mark ............................ E. infumatus McLachlan At most first two flagellomeres incrassate, and these only i,: male. Female subgenital plate with sclattered setae bordering cross-piece of T-shaped mark, these not forming a single row ..................................................................................................... 2 Pterostigma with angular posterior apex. Veins in basal half of forewing, especially M + CUI brown, not contrasting sharply with wing membrane. First male flagellomere decidedly in- crassate (Fig. 4) ............................................ E. nigripes n. sp.



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258 Psyche [June
2b. Pterostigma with rounded posterior apex. Veins in basal half of forewing, especially M + Cur white, contrasting strongly in well colored individuals with fumose wing membrane. First male flagellomere only slightly incrassate .................................. ............................................................... E. crassicornis (Kolbe). Acknowledgments. - Material examined in this paper was in part borrowed from the following institutions: American Museum of Natural History, New York City; Los Angeles County Museum, Los Angeles, California; Museum fur Naturkunde, Humboldt Uni- versitat, Berlin, D. D. R.; Snow Entomological Museum, Lawrence, Kansas; United States Nationtal Museum, Washington, D. C. I wish to thank the officers of these institutions for arranging the loans. I wish to thank the following individuals for gifts of speci- mens: Mr. John K. Bouseman, (Illinois Natural History Survey, Urbana, Illinois), Dr. J. M. Campbell (Department of Agriculture of Canada, Ottawa, Ontario, Canada), Dr. Henry F. Howden (Carleton University, Ottawa, Ontario, Canada). CHAPMAN, P. J.
1930. Corrodentia of the Unitejd States of America: I. Suborder Iso- tecnomera. Journ. N. Y. Ent. SOC. 38: 219-290, 318-402. ENDERLEIN, G.
1911. Die fossilen Copeognathen und ihre Phylogenie. Palaeontogra- phica 5 8 : 279-360.
KOLBE, H. J.
1883. Neue Psociden des Konigl. Zoologischen Museums zu Berlin. Stett. Ent. Zeit. 44: 65-87.
MCLACHLAN, R.
1866. New genera and species of Psocidae. Trans. R. Ent. Soc. Lond. 5 : 345-352.
MOCKFORD, E. L.
1974. Records and descriptions of Cuban Psocoptera. Ent. Americana 48: 103-215.
NEW, T. R.
1972. A collection of Psocidae (Psocoptera) from central Brazil. Arq. 2001. 22: 193-237.
PEARMAN, J. V.
1933. A psocid allied to Eremopsocus infumatus McLachlan (Psocop- tera). Stylops 2: 159-161.
ROESLER, R.
1944. Die Gattungen der Copeognathen. Stett. Ent. Zeit. 105: 117-166. SMITHERS, C. N.
1967. A catalogue of the Psocoptera of the World. Austr. 2001. 16: 1-145.




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