Margaret K. Thayer and Alfred F. Newton, Jr.
Revision of the South Temperate Genus Glypholoma Jeannel, with Four New Species (Coleoptera: Staphylinidae: Omaliinae).
Psyche 85:25-63, 1978.

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REVISION OF THE SOUTH TEMPERATE GENUS
GLYPHOLOMA JEANNEL, WITH FOUR NEW SPECIES (COLEOPTERA: STAPHYLINIDAE: OMALIINAE)*
BY MARGARET K. THAYER AND ALFRED F. NEWTON, JR. Museum of Comparative Zoology, Harvard University Cambridge, Massachusetts 02 138 U.S. A.
The southern temperate silphid genus Glypholoma Jeannel was transferred to the staphylinid subfamily Omaliinae by Newton (1975), and Lathrimaeodes Scheerpeltz (originally placed in the Omaliinae) was then synonymized with it. Newton (op. cit.) also presented additional descriptive information, a new locality record, and some discussion of the affinities of Glypholoma within the Omaliinae. Since that time, four new species of the genus have come to our attention, including one from Australia which greatly enlarges the known range of Glypholoma (previously only parts of Chile and Argentina). The discovery of these new species and the availability of a wealth of material of the type species, pustuliferum Jeannel, for detailed study led to our decision to revise the genus. Measurements, made with an ocular micrometer in a Leitz binocular dissecting microscope, are defined as follows: ~e&gth: measured in lateral view from front of (closed) mandibles to apex of abdomen (excluding genitalia if exserted, and attempting to estimate "normal" degree of contraction of abdomen). Width: maximum body width, across closed elytra at widest point (usually near middle).
Head width: in dorsal view, maximum width including eyes. Head length: measured along midline from anterior margin of labrum to level of centers of ocelli, viewed perpendicular to line of measurement.
*Published with the aid of a grant from the Museum of Comparative Zoology, Harvard University.
Manuscript received by the editor October 6, 1978.



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26 Psyche [March
Antenna1 length: from constriction between scape and its basal articulatory process to apex of last antenna1 segment. Ocellar diameter: measured antero-posteriorly in dorsal view. Pronotal width: maximum width.
Pronotal length: along midline from base to apex, viewed per- pendicular to line of measurement.
Elytral width: same as Width, above.
Elytral length: measured along suture from apex of scutellum to a line tangent to elytral apices.
Prosternal process length: measured along midline from a trans- verse tangent to the anterior margin of the procoxal cavities to apex of prosternal process.
Procoxal length: measured in ventral view between transverse tangents to anterior and posterior faces of procoxae. Mesosternal length: along midline from anterior margin to apex of mesosternal process.
Mesosternal process length: along midline from transverse tan- gent to anterior margin of mesocoxal cavities to apex of mesosternal process.
Mesocoxal length: measured in ventral view between transverse tangents to anterior and posterior margins of mesocoxal cavities. Mesosternal procoxal cavity length: along midline from anterior margin of mesosternum to a line connecting the posterior edges of the cavities.
Metasternal length: along midline between extremities of inter- coxal processes.
Metasternal antecoxal sutures: measured from lateral limit to lateral limit.
Metasternal width: between posterolateral corners of metaster- num.
Hind coxal length: lateral edge to posteromesal corner. Hind femoral length: including trochanter, from posteromesal corner of coxa to most distal point of femur. Hind tibia1 length: along mesa1 side (not including any spines or setae).
Hind tarsal length: from point of insertion of tarsus on tibia to tarsal apex, not including claws or empodial setae. The dividing line between the fourth and fifth hind tarsal segments was taken to be the point of insertion of the fifth segment on the fourth.



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19781 Thayer & Newton - Revision of Genus Glypholoma 27 Abdominal segments are numbered according to their morpho- logical origin. (The first segment visible ventrally is therefore the third segment.)
Mean length and width are given for each species, k one standard deviation.
Preparation of specimens for scanning electron microscope pic- tures consisted of clearing heads, mouthparts, and prothoraces in hot 1 N potassium hydroxide, critical-point drying all parts except elytra, and coating with gold-palladium mixture. Cleared and dissected specimens of pustuliferum and rotundulum and of the aedeagi of other species were examined under dissecting and compound microscopes.
Drawings were made with the aid of a camera lucida attachment on a Leitz binocular dissecting microscope. This study probably would not have come about were it not for S. B. Peck's extensive collecting in Chile and his kindly making this material available to us; he later generously provided us with a multitude of Australian specimens as well. Specimens were borrowed from the following institutions (ab- breviated in the text as indicated) and we extend our thanks to the curators involved for their cooperation in lending specimens. CAS
California Academy of Sciences, San Francisco, California, U.S.A. (D. H. Kavanaugh)
CNC Canadian National Collection, Ottawa, Ontario, Canada (J. M. Campbell)
MCZ Museum of Comparative Zoology, Harvard Uni- versity, Cambridge, Massachusetts, U.S. A. NMVM National Museum of Victoria, Melbourne, Victoria, Australia (A. Neboiss)
Specimens are also deposited in the following collections: ANIC Australian National Insect Collection, Canberra, A.C.T., Australia
ANMT A. F. Newton, Jr. and M. K. Thayer, Cambridge, Massachusetts, U.S. A.
FMNH Field Museum of Natural History, Chicago, Illinois, U.S.A.
SBP
S. B, Peck, Ottawa, Ontario, Canada




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28 Psyche [March
Figs. 1-4. Glyphloma spp: 1. G. pusiuliferum. 2. G. pecki, holotype. 3. G, temporals, holotype. 4. G, tenkicorne, holotype. Scale lines = 1.0 mm.



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19781 Thayer & Newton - Revision of Genus Glypholoma 29 The scanning electron microscope work done for this paper was made possible by National Science Foundation grants BMS-7502606 (J. F. Lawrence, principal investigator) and BMS-7412494 (SEM operating grant) with the superb technical assistance of E. Seling. We also thank L. H. Herman for calling our attention to the Klinger and Maschwitz paper, H. S. Dybas for reading and commenting upon the manuscript, and N. Hinnebusch for typing the manuscript. Glypholoma Jeannel
Glypholoma Jeannel, 1962: 482; Newton, 1975: 53. Type species: Glypholoma pustuliferum Jeannel, 1962: 483, by original designation and monotypy. Lathrimaeodes Scheerpeltz, 1972: 58; (placed in synonomy by Newton, 1975: 54). Type species: Lathrimaeodes pustulipenne Scheerpeltz, 1972: 59, by original designation and monotypy.
Diagnosis: Separable from other known Omaliinae by the exca- vate hind coxae, each elytron with eleven, more or less distinct striae, male genital segment with a small "button" internally at the anterior end of sternite 9, and visible dorsal pleural-coxal articulation in the Fig. 5.
Glypholoma rotundulum. Scale line = 1.0 mm.



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30 Psyche [March
prothorax. In addition, the combined presence of all of the follow- ing characters serves to distinguish Glypholoma from other Omali- inae. (Some other genera share one or a few of these characters with Glypholoma, but no other genus examined possesses all of these characters as Glypholoma does.)
1.
Gular sutures widely separated, minimum separation equal to 0.12-0.18 times the head width.
2. Prothoracic pleural-sternal articulation present. 3.
Procoxa with mesa1 articulating groove.
4.
Mesosternal procoxal cavities long, 0.40-0.75 times the total mesosternal length.
5.
Hind femur relatively short, ranging from 0.92-1.05 times hind coxal length.
6.
Deflexed lateral portion of elytron short, only 0.64-0.67 times as long as total length of elytron (measured in lateral view). 7.
Humeral margin of elytron serrulate.
8.
Median lobe of aedeagus with membranous part of basal bulb allowing dorsal-ventral instead of lateral-lateral contraction and expansion.
9.
Epistomal suture present and complete (although with or without median stem).
Description: Ovoid (narrower posteriorly) to more or less oblong in dorsal view, slightly to strongly convex dorsally in cross section. Sparsely pubescent to nearly glabrous on dorsal surface, widely spaced macrosetae on alternate intervals of elytra in a fairly characteristic pattern (pattern varies some among the species; see figs. 26, 29). Microsculpture lacking on dorsal surface of head, pronotum, and elytra. Length 2.1-3.5 mm, width 1.0-1.5 mm. Head capsule about as in fig. 58, lacking postocular ridge, temples, and nuchal constriction except in temporale, which has temples and a lateral nuchal constriction (fig. 57). Head about 1.8 times as wide as long, about 0.58 times as long as pronotal length, with pair of distinct ocelli on dorsal surface (see especially fig. 6); no anteocellar grooves or pits. Epistomal suture with internal rein- forcing ridge present, angulate or arcuate, with or without median stem. Antennae usually about 1 to 1.5 times as long as head width, varying from filiform (tenuicorne, about 2.2 times head width) to having a moderately developed club of about three to five segments; basal five to eight antenna1 segments glabrous except for a few



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Thayer & Newton - Revision of Genus Glypholoma



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32 Psyche [March
Figs. 10-13. Glypholoma pwiuiiferum. 10. Apical antenna! segments. 1 1. Base of fourth maxillary paipal segment, external view. 12. Hypopharynx and labium, dorsal view, 13. Left maxilla, ventral view. Scale lines: fig. 1 1 = 0.01 mm; others = 0.1 mm,




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19781 Thayer & Newton - Revision of Genus Glypholoma 33 scattered long setae. Apical antenna1 segments simple, without apical gutters or invaginations (see figs. 9, lo), with fairly dense short setae in addition to scattered long setae (see fig. 10, compare basal segments, fig. 7).
Labrum transverse, narrowly rectangular to slightly bilobed anteriorly, ventrally as in fig. 8. Mandible (of pustuliferum and rotundulum, at least) without preapical teeth, with a medial setose area about midway from base to apex and a well-developed molar lobe (see figs. 8, 14-1 7). Molar lobe apparently articulating dorsally and ventrally with mediobasal area of mandible proper. Maxilla more or less as in fig. 13, the palp generally filiform with fourth (apical) segment 2.5to 5 times as long as third, a group of sensilla as in fig. 11 on its dorsolateral surface near the base. Hypopharynx (of pustuliferum and rotundulum, at least) about as in fig. 12. Labium apparently bilobed, with three-segmented palps arising from separ- ate palpigers; segments of palp subequal in length and width (see fig. 7, also Newton, 1975, fig. 3). Mentum large and trapezoidal; gular sutures separate, their minimum separation 0.12 to 0.18 times the head width (see fig. 7).
Pronotum 1.5 to 1.8 times as wide as long, about 0.35 times as long as elytra; widest point variable, from posterior corners to just behind middle; with complete sharp lateral margins, explanate in at least basal half; lacking lateral foveae with internal pillars. Post- coxal process of pronotum acutely triangular, apparently a bar to coxal flexation (see figs. 18, 20). Prosternum with or without median longitudinal keel; prosternal intercoxal process extending one-half to two-thirds of the (antero-posterior) length of the procoxae. Procoxa with external longitudinal keel and mesa1 transverse articulating groove (see figs. 18-2 1). Protrochantin ex- posed, shorter than postcoxal pronotal process, dorsal pleural-coxal (trochantinal-coxal) articulation visible, pleural-sternal articulation present (the last possibly absent in tenuicorne), as in fig. 20. Mesosternum 0.5 to 0.6 times as long as metasternum, with a nearly acute process (except in rotundulum, see figs. 35, 37) extending between the mesocoxae for four to seven tenths of their length (figs. 34,36). Mesosternal process not medially longitudinally carinate. Anterior part of mesosternum with cavities for reception of procoxae 0.4 to 0.75 times as long as whole mesosternum.



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34 Psyche [March
Figs. 14- 17. Gly~holoma pusiul~erum, right mandible. 14-15. Mesal view, whole mandible and molar surface, respectively. 16-1 7. Dorsal view, whole mandible, and part of moia, respectively. Scale Sines: figs. 14,16=0.1 mm; figs. 15.17=0.01 rnm.



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19781 Thayer & Newton - Revision of Genus Glypholoma 35 Figs. 18-2f. Gfypholuma pus~uliferum. 18-19. Prothorax, ventrolateral view, with and without procoxa and trochantin, respectively. 20. Right procoxa and surrounding area, ventrolateral view, with cox8 rotated anteriorly. 21. Prothorax and head, antcroventral view. dpc = dorsal pleural-coxai articulation; g = mesa! articulating groove of coxa; k = external coxal keel; np = postcoxal process of pronoturn; ps = pleural-sternal articulation; sp = prosternal imercoxal process. Scale lines = 0.1 mm.




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19781 Thayer & Newton - Revision of Genus Glypholoma 37 Figs. 22-25,
Glvphotorna spp,, prolegs. 22-24, G. pu~tu/iferurn; 22, male tarsus, oblique ventral view; 23, male left tibia, posterior view; 24, female tarsus, oblique ventral view. 25. G. roiundulum, male tarsus, oblique ventral view. Scale iines = 0.1 rnm.




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38 Psyche [March
Figs. 26-29.
Glyphohmu spp. 26-28. G. pusiul~erum, right elytron; 26, dorso- lateral view 27, detail of fig. 26, including two pustules; 28, strial puncture (detaii of fig. 26). 29. G. roiwsdutum, right elytron, dorsolateral view. Scale Sines: figs. 26,29 = 1.0 mm; fig. 27 = 0.1 mm; fig. 28 = 0.01 mm.



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19781 Thayer & Newton - Revision of Genus Glypholoma 39 Figs. 30-33. G~photoma spp., internal elytral surface. 30-32. 17. pus~uliferum; 30, overall view: 3 I, detail of lateral area (" t" in fig. 30); 32, detail of apex ("a" in fig. 30). 33. G. rotundutum, same view as fig. 32. Scale lines: figs. 30, 3 1 = 0.1 mm; figs. 32, 33 = 0.01 mm.




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40 Psyche [March
of rotundulum, in which the wings only reach the apex of tergite 2 and are not folded. Distinct anal flap present on wings of pustuli- ferum and rotundulum, probably in other species also. Folding pattern of fully-developed hind wing similar to that illustrated for Anthobium (=Eusphalerum) sorbi by Forbes (1926: fig. 33), the first transverse fold being a hinge by which the costal margin is turned about 90櫻 (In an individual, the hinges of the two wings usually form slightly different angles.)
Abdomen with most of segment 8 and part of genital segment exposed; first visible sternite is sternite 3. Tergites 4 or 5 to 8 fairly well to well-sclerotized, spiracles located in tergites 4 or 5 to 8, in membrane beside tergites 1 to 3 or 4; one pair of paratergites on each of segments 3 to 6 or 7, may be partly or entirely fused with sternites laterally (as almost complete fusion in fig. 47); tergite(s) 4 or 4 and 5 with paired patches of medially-directed microtrichia which cover about half to nearly all of the surface of the tergites (figs. 40-43); tergite 7 with an apical fringe ("palisade fringe" of some authors) as in fig. 45. Intersegmental membranes with a brick wall pattern of irregular plates as in fig. 44, some dorsal plates with posterior teeth as shown. Sternite 2 extending up around sides of abdomen slightly, but in tenuicorne appearing to be membranous, its limits therefore not determinable. Sternites 2 and 3 with small intimately associated intercoxal processes, more or less as in fig. 48, sternite 3 (at least in pustuliferum and rotundulum) with a trans- verse fold across its middle one-fourth to one-half about at posterior margin of intercoxal process (this area not visible in other species because of telescoping of abdomens). Sternite 3 without distinct coxal cavities, metacoxae simply protruding parallel to sternite's surface; at least in some species with a ridge near basal margin and, like sternites 4 to 5 or 6, a curved ridge just inside each lateral margin. Sternite 8 with anterior median projection (see figs. 49, 61-65) associated with a gland system, similar to that described by Klinger and Maschwitz (1977). In at least three species (rotundulum, pecki, pustuliferum) gland reservoir extending anteriorly as far as anterior margin of segment 5.
Male: Peg setae apparently absent from trochanters, femora, and tibiae; first four segments of protarsus slightly broadened, spatulate setae in pairs on segments 1 to 3 of protarsus (figs. 22, 25), and singly on first two (possibly three inpecki) segments of mesotarsus



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Figs. 34-37. G1,yphoIoma spp., pterothorax. 34,36. G. pustu-, ventral and oblique lateral views, respectively. 35, 37. G. rotundulum, ventral and oblique lateral views, respectively. Scale lines = 0.1 ram.



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Figs. 38-41.
G1,vpholoma spp. 38-40. G. pustuliferum; 38, metastemum and right metacoxa, lateral view; 39, male left mesotarstis, ventral view; 40, abdomen, dorsal view. 41. G. mndiilum, pterothorax and abdomcn, dorsal view. t7 = tergite 7. Scale lines: figs. 38, 39 = 0.1 mm; figs. 40, 41 = 1.0 mm.



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19781 Thayer & Newton - Revision of Genus Glyphdoma 43



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44 Psyche [March




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19781 Thayer & Newton - Revision of Genus Glypholoma 45 (see fig. 39). Genital segment with tergite 9 narrowly continuous across dorsal midline, tergite 10 short, sternite 9 attenuate, fairly well sclerotized in posterior one-third or so, only lightly sclerotized anteriorly, with a small cylindrical protuberance on internal surface near anterior end (see figs. 66-75). Aedeagus of general staphylinid type (median lobe with basal bulb, small dorsal median foramen, ventro-apical median orifice); parameral side of aedeagus facing dorsally within abdomen; small basal piece present as a more or less C- or U-shaped strap on dorsal side of basal bulb (often very lightly sclerotized and difficult to see); basal bulb of median lobe lightly sclerotized, with a membranous band around the abparameral side, presumably allowing dorsal-ventral bellows-like contractions; in- ternal sac with dense armature of short, fine spines; apex of median lobe a finger-like projection, even with or slightly shorter than apices of parameres; parameres with or without subapical setae (see figs. 76-85).
Female: Pro- and mesotarsus without spatulate setae, as in fig. 24. Genitalia (based on pustuliferum and rotundulum, except as noted) about as in Newton, 1975, figs. 8-10: sclerotized spermatheca apparently absent, sternite 9 present, tergite 9 divided into two lateral sclerites, tergite 10 U-shaped; proximal gonocoxites separate, about half as long as distal gonocoxites; in all species in which females are known, distal gonocoxites separate, each bearing at its apex a short stylus with 1 (or 2?) long apical seta(e) and one or more subapical setae.
Immature stages unknown.
Range: Southern Chile and Argentina south to Tierra del Fuego; Victoria, Australia; see maps, figs. 50, 51. KEY TO THE SPECIES OF GLYPHOLOMA JEANNEL Antenna filiform (fig. 56); elytra without raised pustules; dorsal surface uniform in color; body form as in fig. 4 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. tenuicorne n.sp.
Antenna with gradual apical club (figs. 52-55); elytra with or without raised pustules; dorsal surface with various color markings; body form otherwise (figs. 1-3, 5) . . . . . . . . . .2 Elytra with raised pustules on alternate intervals (fig. 26) .3 Elytra without raised pustules (fig. 29) . . . . . . . . . . . . . . . . . .4



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46 Psyche [March
3.
Postocular lobe present (fig. 57); metasternal antecoxal sutures long, about three-fourths the width of the metasternum (fig. 59); elytral pustules concolorous with surrounding area . . ................................. G. temporale nsp. Postocular lobe absent (fig. 58); metasternal sutures short, no more than about one-third the width of the metasternum (fig.