Revision of eminoculus schuh (heteroptera: miridae: phylinae) from South Africa, with the description of five new species.
Article Type: Report
Subject: Mirids (Research)
Authors: Schuh, Randall T.
Wu, Grace
Pub Date: 01/01/2009
Publication: Name: Entomologica Americana Publisher: New York Entomological Society Audience: Academic Format: Magazine/Journal Subject: Biological sciences; Science and technology Copyright: COPYRIGHT 2009 New York Entomological Society ISSN: 1947-5136
Issue: Date: Jan, 2009 Source Volume: 115 Source Issue: 1
Topic: Event Code: 310 Science & research
Geographic: Geographic Scope: South Africa Geographic Code: 6SOUT South Africa
Accession Number: 257814218
Full Text: Abstract--Eminoculus Schuh is revised and an updated diagnosis is presented for the genus. Seven species are recognized, five of which are described as new. Diagnoses, digital habitus photographs, illustrations of male and female genitalic structures, keys to both the males and females, and detailed locality are provided. Scanning micrographs of diagnostic morphological structures the head, pronotum, scent-gland evaporatory area, vestiture, and pretarsus--are included for four species. A phylogenetie analysis is presented, using the South African species Lasiolabopella capeneri Schuh and the Nearctic Chlamydatus keltoni Schuh and Schwartz as outgroups. Host data are presented for 6 species, and patterns of host association with the "mesem" clade of the Aizoaceae are discussed.

Key words: phylogenetic analysis, host plants, Aizoaceae.

INTRODUCTION

The present paper represents part of the most recent attempt to document the Miridae fauna of southern Africa. The first descriptive work on African Miridae was that of Poppius (1912, 1914). It was nearly 50 years before any additional remotely comprehensive work began to appear. Carvalho et al. (1960) contributed the South African Animal Life series chapter on the Miridae, a work that was based exclusively on the collections made by the Brinck and Rudebeck Lund University Expeditions of 1950--1951. Because those collections were not made by specialists in the group, and because they were not necessarily focused on areas of greatest diversity and endemism, the work of Carvalho et al. (1960) was limited in scope, dealing with only 41 species, many of which were described beginning with the work of Carl Stal (1855) and later in the works of Poppius cited above.

Schuh (1974) was the first to undertake specialized collecting of Miridae in southern Africa, in conjunction with field work by James A. Slater and Merrill H. Sweet in 19671968. That work concentrated primarily on the Blissidae and Rhyparochromidae, groups that had been shown to be extremely diverse through the South African Animal Life contribution of Slater on the Lygaeidae sensu lato (Slater, 1964); nonetheless, a substantial amount of new specimen materials for the Miridae was acquired. Schuh (1974) treated 103 species in 54 genera in the subfamilies Orthotylinae and Phylinae, 81 of the species and 20 of the genera having been described as new.

Randall Schuh and Gerasimos Cassis included the Cape Floristic Province of South Africa as part of their National Science Foundation (USA) funded Planetary Biodiversity Inventories (PBI) project in 2003, because of the tremendous diversity of phytophagous Heteroptera that had previously been revealed through the specialized collecting efforts of Slater, Sweet, and Schuh. They chose to focus their efforts on Namaqualand and the Fynbos, biotic elements that still remained very poorly sampled for Heteroptera. Forty person weeks of field effort have revealed substantially greater diversity in the Miridae than was previously known to exist and provides a much more extensive picture of host associations. In these pages we describe a limited part of that diversity.

Eminoculus, the subject of the present paper, was described by Schuh (1974) on the basis of very limited material and placed in the Phylini. Most of the known specimens were collected on ornamental plantings of Drosanthemum floribundum (Harv.) Schwart. in Pretoria. PBI-funded field work has made available extensive collections of five new species, all of them restricted to the "mesem" clade within the Aizoaceae (see further discussion below). Schuh's generic diagnosis is updated to reflect knowledge derived from the examination of this much more extensive geographic and taxic sample.

MATERIALS AND METHODS

During the course of this research project matrix code labels were affixed to the 1,800 specimens examined as a way to uniquely identify them; these codes are therefore referred to as "unique specimen identifiers" (USIs). The USI codes, e.g., AMNH_PBI 00094810, are composed of an institution and project code (AMNH_PBI) and a unique number (00094810). USI codes are included in the locality data and in the figures and captions.

All latitude-longitude data are presented in the "specimens examined" section of the present paper in degrees and decimal parts thereof. Altitude data are treated as metric. Please refer to the www.discoverlife.org website as a way of accessing additional information on specimens examined for the Planetary Biodiversity Inventories Project on Plant Bugs and the present paper.

Structural information is documented through the use of light and scanning electron microscopy. Color digital habitus images of the bugs were prepared using a Microptics-USA photomicrographic apparatus with Infinity optics. Habitus photos are proportional to the size of the actual specimens so that relative sizes can be deduced from comparison of the specimen images. Actual sizes of specimens can be determined by referring to table 1. Host field photos were made using a Nikon D1 SLR digital camera.

States of characters used in the cladistic analysis are indicated on the illustrations to facilitate appreciation of group-forming attributes.

All measurements are in millimeters and were made using a micrometer driven stage, micrometer output being written directly to a spreadsheet.

EMINOCULUS SCHUH

Eminoculus Schuh, 1974:160 (n. gen.).

TYPE SPECIES: Eminoculus drosanthemi Schuh, 1974

REVISED DIAGNOSIS: Recognized by the brownish black to black coloration of the body, the weakly to strongly stylate eyes, the strong alary sexual dimorphism, the ventrally directed hairs on the lateroventral surface of the claws, and the structure of the vesica in the male, particularly the field of spicules subtending the secondary gonopore.

Always breeds on species of Aizoaceae (Mesembryanthemoideae and Ruschioideae).

Form of the head and eyes of some species similar to that of Lasiolabopella Schuh and Lasiolabops Poppius, but easily distinguished from members of those groups by their lack of alary sexual dimorphism, by the presence of a pronotal collar in most species of Eminoculus, and by the structure of the male genitalia.

REDESCRIPTION: Male: Macropterous; range total length 2.52-4.15, range apex clypeus-cuneal fracture 1.72-2.81, range width pronotum 0.93-1.41. COLORATION (Fig. 1): Generally black or blackish-brown, very heavily sclerotized; femora and tibiae varying from entirely black to partially golden; coloration of tarsal segments variable. SURFACE AND VESTITURE: Body surface, most notably head, pronotum, and scutellum ranging from smooth and dull to highly polished (Fig. 1), shining, and transversely rugulose or rugose (Figs. 5A, B, 7A, 9A, 13A); coxae generally pruinose; vestiture variable, ranging from short and very sparse over much of dorsum in some species (E. drosanthemi; Fig. 9A, B) to very long and erect on head and pronotum in others (E. auratus Schuh and Wu, E. hirsutus Schuh; Figs. 7A, B); antennae with short semi-erect vestiture on all segments, segment 1 also with a few to many erect black spines; abdominal venter with long, reclining setae (Figs. 7E, 9E). STRUCTURE: Head: Vertical, very broad, eyes substylate to strongly stylate (Figs. 1, 5A, 7A); frons convex; head viewed anteriorly V-shaped below eyes; mandibular plates bulging; bucculae expanded; gula obsolete; antenna inserted below ventral margin of eye at about level of dorsal margin of clypeus; antennal segment 1 moderately enlarged, sometimes elongate, segment 2 usually stout and of uniform diameter, segments 3 and 4 of subequal diameter and about one-half diameter of segment 2. Thorax: Often with a flattened collar about as wide as diameter of antennal segment 1 (Figs. 7A, B, 9A), rarely absent (Fig. 5A, B); posterior margin of pronotum weakly to rather strongly concave (Figs. 7A, B, 9A); calli ranging from weakly elevated and separate (Fig. 5A) to more strongly inflated and confluent medially (Fig. 7A); mesoscutum broadly exposed and declining posteriorly; scutellum nearly fiat; supra-coxal lobe of metepisternum either lying fiat against coxa or more frequently projecting later ally as rounded or spinelike process: metathoracic scent-gland evaporatory area as in Figure SD. Hemelytra. Corial margin usually weakly convex, sometimes sinuous and concave on posterior halt': cells of membrane with smoothly, though sometimes sharply, rounded posterior margin (Fig. 1). Leg: Rather short, posterior femora noticeably bowed: tibiae with scattered slender spines on ventral surface, lacking rows of tiny closely-spaced spines: claws broad basally, nearly straight over most of length, sharply curving apically (Figs. SE, F, 7D, 9C): parempodia setiform, pulvilli relatively large, attached to most of ventral suface of claws: claws ventrolaterally with from one to six ventrally directed hairs (Figs. 5E, F, 7D, 9C, D. 13C, D). Abdomen: Elongate and slender in larger species, shorter and broader in smaller species (Fig. 7E). GENITALIA (e.g., Figs. 9E, F, 10, 14): Pygophore: Conical (Fig. 9E, F). Vesica: Formed of two sinuously curving straps, apex in form of a single, slender, apically-attenuated spine, dorsal surface of spine with at least weak subbasal indentation (notch), indentation frequently stronger: secondary gonopore well developed, ovoid, subtended by field of spicules pointing apicad. Phallotheca: L-shaped as in many Phylini, devoid of distinctive ornamentation. Left Paramere: Overall shape typical of Phylini: very heavily sclerotized over most of pararmere, opaque, except for elongate, vertically oriented, transparent area lying between base of posterior process and paramere shaft. Right Paramere: Elongate, with dorsal surface strongly transversely arched: apex usually with fingerlike process: very heavily selerotized, largely opaque in most species.

Female: Brachypterous, with abbreviated wings leaving 2 or 3 terminal abdominal terga exposed: of small to relatively large size: elongate to ovoid: range total length 2.1(>4.31, range length apex clypeusapex wing 1.72 2.81, range width pronotum 0.941.38. COLORATION: As in males, but antennal segments and legs often with extensive golden areas. SURFACE AND VESTITURE: As in males. STRUCTURE: Cuneus and costal fracture absent, membrane always present as narrow crescent. GENITALIA (Fig. 11): Posterior wall a simple sclerotized plate: sclerotized rings moderately infolded laterally with the lateral portion of the underlying dorsal labiate plate triangular plate: vestibulum relatively short and simple.

HOSTS: Host data are now available for six of the seven recognized species, showing clearly their strict association with members of the "mesem" clade, the subfamilies Mesembryanthemoideae and Ruschioideae, of the Aizoaceae. Detailed host records are given in each of the species treatments. Field photographs of representative hosts for each species are presented in Figures 2 and 3.

[FIGURE 1 OMITTED]

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[FIGURE 3 OMITTED]

Klak et al. (2003) discussed the history of classification of the Aizoaceae and the practice of some authors to break the group into a number of segregates, each elevated to the family level. The analysis of Klak et al. (2003) showed that the Mesembryanthemaceae of authors is a monophyletic group, formed of two monophyletic subgroups Mesembryanthemoideae and Ruschioideae. The remaining Aizoaceae do not form a monophyletic group, however, and are paraphyletic when the "mesems" are excluded. We therefore use the family name Aizoaceae throughout this paper, indicating subfamily placements under Mesembryanthemoideae and Ruschioideae. We note that the family name Ficoidaceae has also been used to refer to the "mesem" clade.

PHENOLOGY: Field experience in South Africa, particularly in Namaqualand, from Cistrusdal to Port Nolloth, suggests that species of Eminoculus appear early in the year. The majority of the specimens utilized in this revision were collected during the first two weeks of September, with the limited remaining material having been collected between mid-October and mid-November. Although the bugs are frequently collected on plants in flower, this does not seem to be a requirement, and at least one species is known only from sterile material.

DISTRIBUTION (Fig. 4): Known primarily from the Namaqualand and the Little Karoo areas of South Africa with some collections outside of this area apparently representing introductions with ornamental plantings, as in the case of specimens of E. drosanthemi from Pretoria.

DISCUSSION: Although Eminoculus was named for its stylate eyes, we have updated the original diagnosis to accommodate the less conspicuously stylate condition found in E. atrisetosus Schuh and Wu. We have also removed the pronotal collar from the generic diagnosis, as this feature is not found in E. atrisetosus. Nonetheless, E. atrisetosus possesses characteristics of the male genitalia, such as the indentation subbasally on the dorsal surface of the apical spine and the field of spicules subtending the secondary gonopore, that ally it with the remaining species here placed in Eminoculus. Furthermore, E. atrisetosus possesses ventrally directed hairs on the ventrolateral surface of the claw, a feature known to occur nowhere else in the Miridae except in the Eccritotarsini.

The body and parameres in Eminoeulus are heavily sclerotized. The sclerotization is so intense in the left paramere that for the most part it is not possible to determine the location of setae on the body of the paramere. We have, however, been able to determine that a group of Eminoculus spp. has a distinctive grouping of setae at the apex of the anterior process of the left paramere.

Since males are macropterous, while females are brachypterous and flightless, this may have contributed to the fact that the latter were captured in greater abundance in almost all collecting events.

Eminoculus atrisetosus, new species Figures 1, 2, 4, 5, 6

DIAGNOSIS: Males and females recognized by the weakly stylate eyes, the absence of a flattened pronotal collar, the sericeous, golden, recumbent vestiture on the dorsum, the smooth, dull brownblack dorsum, and the many erect black spines on the medial and occasionally lateral surface of antennal segment 1 (Figs. 1, 5). All other Eminoeulus spp. with eyes much more strongly stylate and with a distinct flattened pronotal collar.

DESCRIPTION: Male: Total length 3.07-3.25; length apex clypeus-cuneal fracture 2.27-2.4; width pronotum 1.1-1.15. COLORATION (Fig. 1): Body and appendages brown black; margins of supracoxal lobes of pro-, meso-, and metathorax rather broadly yellow. SURFACE AND VESTITURE: Head, anterior lobe of pronotum, and hemelytra smooth and dull; posterior lobe of pronotum and scutellum rugulose and dull; calli smooth and dull; antennal segment 1 always with numerous, erect spines on mesial surface (<15) (Fig. 5A, B); genal region dorsad of bucculae with many long, erect, fine setae; head and pronotum with moderately long, sericeous, reclining, golden setae (Figs. 1, 5A, B); hemelytra with moderately long, appressed golden setae, primarily on anterior half (Fig. 5C), posterior half of corium with dark, simple, appressed setae; 3 hairs arising from lateroventral surface of claw (Fig. 5E, F). STRUCTURE: Head: Eyes sub-stylate; posterior margin of head and eyes curvilinear, contiguous with anterior pronotal margin (Fig. 5A, B); antennal segment 1 moderately swollen, convex on medial margin, short; labium reaching to foretrochanter. Thorax: Anterior margin of pronotum lacking a flattened collar (Fig. 5A, B); calli distinct, weakly elevated, not confluent; pronotum nearly flat longitudinally, very weakly convex transversely, lateral margins very slightly convex, posterior margin very shallowly concave; proepisternal supracoxal lying flat against coxa, not protruding; costal hemelytral margin weakly convex (Fig. 1). GENITALIA (Fig. 6): Vesica: Body relatively slender; apical spine with weak subbasal notch on dorsal surface, flange on dorsal surface of vesica truncate at level of base of secondary gonopore, angled toward gonopore. Phallotheca: As in generic description. Left Paramere: Anterior process modestly shorter than posterior, extending slightly beyond shaft of paramere, with a few scattered setae; posterior process parallel-sided, at more or less same level as dorsal margin of body of paramere. Right Paramere: Irregular elongate ovoid; apex with fingerlike projection located to left of center.

[FIGURE 4 OMITTED]

[FIGURE 5 OMITTED]

Female: Medium-sized, total length 3.12-3.54; length apex clypeus-apex wing 2.81 3.06; width pronotum 1.15-1.27. COLORATION (Fig. 1): General coloration of body brown-black to black. SURFACE AND VESTITURE: As in males. STRUCTURE: Abdomen ovoid and not distinctly acuminate apically, only slightly protruding beyond posterior margin of hemleytra. GENITALIA: See generic description.

[FIGURE 6 OMITTED]

ETYMOLOGY: From the Latin ater, black, and setosus, bristly, in reference to the numerous heavy black setae, especially on the mesial surface of antennal segment 1.

HOSTS: cf. Phyllobolus spp. (Fig. 2A, B) (Aizoaceae: Mesembryanthemoideae)

DISTRIBUTION (Fig. 4): Known only from northernmost Namaqualand in the area between Steinkopf and Port Nolloth.

HOLOTYPE: SOUTH AFRICA: Northern Cape: 11.7 km N of R382 toward Lekkersing, 29.22102[degrees]S 17.12044[degrees]E, 193 m, 07 Sep 2004, Schuh, Schwartz, Henry, Wyniger, Forero, cf. Phyllobolus sp. (Aizoaceae), det. Field ID, 1[male] (AMNH_PBI 00171859) (SANC).

PARATYPES: SOUTH AFRICA: Northern Cape:

11.7 km N of R382 toward Lekkersing, 29.22102[degrees]S 17.12044[degrees]E, 193 m, 07 Sep 2004, Schuh, Schwartz, Henry, Wyniger, Forero, cf. Phyllobolus sp. (Aizoaceae), det. Field ID, 4[male] (AMNH_PB1 00171909-AMNH_PBI 00171912), 4[male] (AMNH_PBI 00172042-AMNH_PBI 00172045) (AM). 101[male] (AMNH_PBI 00171854AMNH_PBI 00171858, AMNH_PBI 00171860AMNH_PB100171893, AMNH_PBI 00171897-AMNH_PBI 00171900, AMNH_PBI 00171917-AMNH_PBI 00171974), 147[male] (AMNH_PBI 00171974-AMNH_PBI 00172033, AMNH_PBI 00172046-AMNH_PBI 00172047, AMNH_PBI 00172054-AMNH_PB100172138) (AMNH). 4[male] (AMNH PBI 00171913-AMNH PBI 00171916), 3[male] (AMNH_PBI 00172051-AMNH_PBI 00172053) (CNC). 4[male] (AMNH_PBI 00171905-AMNH_PBI 00171908), 4[male] (AMNH_PBI 00172034-AMNH_PBI 00172037) (SAMC). 4[male] (AMNH_PBI 00171901-AMNH_PB1 00171904), 4[male] (AMNH_PBI 00172038-AMNH_PB1 00172041) (SANC). 29[male] (AMNH_PBI 00190279-AMNH_PBI 00190307), 49[female] (AMNH_PBI 00190308-AMNH PBI 00190356) (USNM). 3[male] (AMNH_PBI 00171894-AMNH_PBI 00171896), 3[female] (AMNH_PBI 00172048-AMNH_PBI 00172050) (ZISP). 70 km E of Port Nolloth, 29.248[degrees]S 17.59[degrees]E, 26 Nov 1974-26 Nov 1974, J. G. and B. L. Rozen, 2[male] (AMNH_PBI 00172857, AMNH_PBI 00172858), 2[female] (AMNH_PBI 00172859, AMNH_PBI 00172860) (AMNH). 9.4 km N of R382 toward Lekkersing, 29.23989[degrees]S 17.11204[degrees]E, 192 m, 07 Sep 2004, Schuh, Schwartz, Henry, Wyniger, Forero, cf. Phyllobolus sp. (Aizoaceae), det. P. Chesselet voucher-NYBG, 11[male] (AMNH_PBI 00172140-AMNH_PBI 00172150), 34[female] (AMNH_PBI 00172151-AMNH_PBI 00172184) (AMNH). 3[male] (AMNH_PBI 00190270-AMNH_PBI 00190272), 6[female] (AMNH_PBI 00190273-AMNH_PBI 00190278) (USNM).

OTHER SPECIMENS EXAMINED: SOUTH AFRICA: Northern Cape: 11.7 km N of R382 toward Lekkersing, 29.22102[degrees]S 17.12044[degrees]E, 193 m, 07 Sep 2004, Schuh, Schwartz, Henry, Wyniger, Forero, cf. Phyllobolus sp. (Aizoaceae), det. Field ID, 3 nymphs (AMNH_PBI 00190358-AMNH PBI 00190360) (USNM).

Eminoculus auratus, new species

Figures 1, 2, 4, 7, 8

DIAGNOSIS: Recognized by the large size, long erect setae on the head and pronotum, the strongly stylate eyes in both sexes, and weakly concave hemelytral margins in the male (Figs. 1, 7). Similar in size to E. rugosus and E. hirsutus. Distinguished from the former by the more exaggerated nature of the stylate eyes in E. auratus, by the long erect setae on the head and pronotum, and by the golden appendages in the female. Distinguished from E. hirsutus by the straight-sided to weakly convex lateral margin of the hemelytra in that species and the weakly convex condition found in E. auratus. Distinguished from E. laevigatus by the setae on the head and pronotum being sparse, only moderately long, and inconspicuous in that species. Distinguished from E. drosanthemi and E. lanuginosus by the significantly larger size of E. auratus.

DESCRIPTION: Male: Total length 3.8-4.15; length apex clypeus-cuneal fracture 2.55 2.81; width pronotum 1.22-1.41. COLORATION (Fig. 1): Body mostly black, posterior half of hemelytra somewhat brownish; femora black with pale apex; tibiae moderately infuscate; tarsal segments 1 and 3 black, 2 pale; margin of proepisternal supracoxal lobe yellow, margin of supracoxal lobes of meso-, and metathorax yellow. SURFACE AND VESTITURE: Head and pronotum highly polished, shining; posterior lobe of pronotum weakly rugose, polished, and shining (Fig. 7A); scutellum conspicuously transversely rugose, dull; hemelytra variably rugulose and dull; antennal segment 1 with few slender, erect spines on medial surface (<15); head and pronotum with very long, sericeous, erect, silvery setae (Fig. 7A); genal region dorsad of bucculae with long, erect, fine setae; hemelytra densely set with moderately long, sericeous, reclining golden setae (Fig. 7C); ~4-5 hairs arising from lateroventral surface of claw (Fig. 7D). STRUCTURE: Head: Eyes grossly stylate, almost spherical in outline when viewed from above; posteromedial angle of eye removed from anterolateral angle of pronotum by more than 2 times diameter of antennal segment 2 (Figs. 1, 7B); posterior margin of head and eyes curvilinear (Fig. 7B); antennal segment 1 parallel sided and relatively long; labium reaching to trochanteral joint of forecoxae. Thorax: Anterior margin of pronotum with a distinct flattened collar about as wide as diameter of antennal segment 1 (Fig. 7A, B); calli extremely pronounced, confluent anteriorly; pronotum steeply inclined posteriorly, with deep transverse impression dividing calli and posterior lobe; posterior margin weakly concave; costal margin of hemelytra weakly concave over most of length; proepisternal supracoxal lobe strongly projecting and spoonlike. Abdomen: Elongate, slender (Fig. 7E). GENITALIA (Figs. 7F, 8): Vesica: Body relatively broad; apical spine with very weak subbasal notch on dorsal surface; margin of flange on dorsal surface of vesica flowing smoothly to base of secondary gonopore. Phallotheca: As in generic description. Left Paramere: Anterior process modestly shorter than posterior, slightly extending beyond shaft of paramere, with diffuse clump of a few setae apically; posterior process not elevated above dorsal margin of body of paramere, moderately broad apically. Right Paramere: Irregular elongate ovoid, apex with centrally located fingerlike process.

Female: Large, elongate, total length 3.52-4.31; length apex clypeus-apex wing 2.80-3.18; width pronotum 1.17-1.25. COLORATION (Fig. 1): Body intensely black; appendages golden, except antennal segment 2 infuscate proximally and distally, antennal segments 3 and 4 infuscate, and tarsal segments 1 and 3 infuscate. SURFACE AND VESTITURE: As in males. STRUCTURE: Costal margin of hemelytra nearly straight; abdomen nearly parallel-sided over much of length, distinctly acuminate apically and projecting well beyond posterior margin of hemelytra. GENITALIA: See generic description.

[FIGURE 7 OMITTED]

ETYMOLOGY: From the Latin auratus, golden, for the color of the legs in the females.

HOSTS: Ruschia muelleri (L. Bolus) Schwantes and R. decurrens (L. Bolus) N.E. Br. (Fig. 2C, D) (Aizoaceae: Ruschioideae)

DISTRIBUTION (Fig. 4): Broadly distributed in Namaqualand from Clanwilliam in the south to near Port Nolloth in the north.

[FIGURE 8 OMITTED]

Holotype: SOUTH AFRICA: Western Cape: 3 km S of Clanwilliam, E side of reservoir, 32.21115[degrees]S 18.90545[degrees]E, 185m, 03 Sep 2004, Schuh, Schwartz, Henry, Wyniger, Forero, Ruschia decurrens (L.Bolus) N.E.Br. (Aizoaceae), det. P. Chesselet VOUCHER-NYBG, 1[male] (AMNH_PBI 00170795) (SANC).

Paratypes: SOUTH AFRICA: Northern Cape: 19 km E of Port Nolloth on R382, 29.30934[degrees]S 17.06537[degrees]E, 168 m, 06 Sep 2004, Schuh, Schwartz, Henry, Wyniger, Forero, Ruschia stricta L.Bolus (Aizoaceae), det. P. Chesselet VOUCHER-NYBG, 1[male] (AMNH_PBI 00172863) (AMNH). 21 km W of Springbok on R355, 29.69466[degrees]S 17.71383[degrees]E, 516 m, 05 Sep 2004, Schuh, Schwartz, Henry, Wyniger, Forero, Ruschia muelleri (L.Bolus) Schwantes (Aizoaceae), det. P. Chesselet VOUCHER-NYBG, 10[male] (AMNH_PBI 00170810-AMNH_PBI 00170818, AMNH_PBI 00170821), 2[female] (AMNH_PBI 00170827, AMNH_PBI 00170828) (AMNH). 1[male] (AMNH_PBI 00170820) (SAMC). 1[male] (AMNH_PBI 00170819) (SANC). 1[male] (AMNH_PBI 00190435) (USNM). 5.2 km E of Kamieskroon on Kasmiesberg Rd, 30.1906[degrees]S 17.9753[degrees]E, 983 m, 27 Oct 2003, Schuh, Cassis, Weirauch, Ruschia sp. (Mesembryanthemaceae), det. P. Chesselet VOUCHER-NYBG, 2[male] (AMNH_PBI 00172871, AMNH_PBI 00172872), 1[female] (AMNH_PBI 00172873) (AM). 1[male] (AMNH_PBI 00171363), 2[female] (AMNH_PBI 00171366, AMNH_PBI 00171367), 1[male] (AMNH PBI 00172864) (AMNH). 1[male] (AMNH_PBI 00172865), 1[female] (AMNH_PBI 00172875) (CNC). 2[male] (AMNH_PBI 00172866, AMNH_PBI 00172867) (SAMC). 2[male] (AMNH_PBI 00172868, AMNH_PBI 00172869) (SANC). 1[male] (AMNH_PBI 00172870), 1[female] (AMNH_PBI 00172874) (Z1SP). ~3 km W of Wallekraal on Hondeklipbaai Rd, 30.36721[degrees]S 17.47401[degrees]E, 124 m, 26 Oct 2003, Schuh, Cassis, Weirauch, (Aizoaceae), 1[female] (AMNH_PBI 00170833) (AMNH). Western Cape: 3 km S of Clanwilliam, E side of reservoir, 32.21115[degrees]S 18.90545[degrees]E, 185 m, 03 Sep 2004, Schuh, Schwartz, Henry, Wyniger, Forero, Ruschia decurrens (L.Bolus) N.E.Br. (Aizoaceae), det. P. Chesselet VOUCHER-NYBG, 9[male] (AMNH_PBI 00170793-AMNH_PBI 00170794, AMNH_PB| 00170796-AMNH_PBI 00170802), 2[female] (AMNH_PBI 00170807, AMNH_PBI 00170808) (AMNH). 6[male] (AMNH_PBI 00190446-AMNH_PBI 00190451), 1[female] (AMNH_PBI 00190452) (USNM).

Other Specimens Examined: SOUTH AFRICA: Northern Cape: 21 km W of Springbok on R355, 29.69466[degrees]S 17.71383[degrees]E, 516 m, 05 Sep 2004, Schuh, Schwartz, Henry, Wyniger, Forero, Ruschia muelleri (L.Bolus) Schwantes (Aizoaceae), det. P. Chesselet VOUCHER-NYBG, 9 nymphs (AMNH_PBI 00170822-AMNH_PBI 00170826, AMNH_PBI 00170829-AMNH PBI 00170832) (AMNH). 5.2 km E of Kamieskroon on Kasmiesberg Rd, 30.1906[degrees]S 17.9753[degrees]E, 983 m, 27 Oct 2003, Schuh, Cassis, Weirauch, Ruschia sp. (Aizoaceae), det. P. Chesselet VOUCHER-NYBG, 2 nymphs (AMNHPBI 00171364, AMNH_PBI 00171365). 32 nymphs (AMNHPBI 00172876-AMNH_PBI 00172907) (AMNH). Western Cape: 3 km S of Clanwilliam, E side of reservoir, 32.21115[degrees]S 18.90545[degrees]E, 185 m, 03 Sep 2004, Schuh, Schwartz, Henry, Wyniger, Forero, Ruschia decurrens (L.Bolus) N.E.Br. (Aizoaceae), det. P. Chesselet VOUCHER-NYBG, 5 nymphs (AMNH_PBI 00170803-AMNH_PBI 00170806, AMNH_PBI 00170809) (AMNH).

Eminoculus drosanthemi Schuh

Figures 1, 2, 4, 9, 10

Eminoculus drosanthemi Schuh, 1974: 161, Figs. 65, 251-254 (n. sp.).

REVISED DIAGNOSIS: Males and females recognized by their small size (mean length male 2.63)

and highly polished, shining, black dorsum (Fig. 1). Abdomen in female more or less truncate posteriorly and not strongly projecting beyond posterior margin of hemelytra. Size in both sexes most similar to E. lanuginosus, but with other distinct differences between the two species. Posterior margin of head and eyes in E. drosanthemi more or less conforming to anterior margin of pronotum, whereas eyes distinctly protruberant in E. lanuginosus; dorsum of E. drosanthemi with evenly distributed, sparse, extremely short, black vestiture (Figs. 1, 7B), whereas dorsum in both sexes of E. lanuginosus with dense, long, silvery, woolly vestiture; femora in E. drosanthemi black, not pale or light brown with variable maculae, as in males and females of E. lanuginosus. Vesica in E. drosanthemi sharing with E. atrisetosus and E. lanuginosus the presence of a distinct subbasal notch on the dorsal surface of the apical spines and a truncate or toothed flange just proximal to secondary gonopore on the dorsal side of the vesica.

REDESCRIPTION: Male: Body relatively stout; total length 2.52-2.79; length apex clypeus-cuneal fracture 1.72-1.98; width pronotum 1.02-1.12. COLORATION (Fig. 1): Basic body coloration shining black; antennal segments 1 and 2 pale to light brown, sometimes distal quarter of antennal segment 2 and segments 3 and 4 entirely brown; femora black with pale apex; tibiae moderately infuscate; tarsal segments 1 and 3 dark, 2 pale; inner margin of proepisternal supracoxal lobe yellow. SURFACE AND VESTITURE: Head, pronotum, scutellum, and hemelytra smooth and highly polished, shining (Fig. 9A); posterior lobe of pronotum transversely rugose, contrasting with extremely smooth calli (Fig. 9A); antennal segment 1 with a few slender, erect spines on mesial surface (Fig. 9A); genal region dorsad of bucculae with long, erect, fine setae; head and pronotum with very short, sparse, simple, semierect, black setae (Figs. 1, 9A); hemelytra with evenly distributed, short, appressed, black setae (Fig. 9B); 1 hair arising from lateroventral surface of claw (Fig. 9C, D). STRUCTURE: Head: Eyes weakly stylate; posterior margin of eyes and vertex contiguous with anterior pronotal margin but projecting beyond its lateral margin (Fig. 9A); antennal segment 1 weakly swollen, parallel sided, relatively short; labium reaching to trochanteral joint of middle coxae. Thorax: Anterior margin of pronotum with distinct flattened collar about as wide as diameter of antennal segment 1 (Fig. 9A); calli weakly elevated and not confluent; pronotum nearly flat longitudinally, weakly convex transversely, posterior margin weakly concave (Fig. 9A); lateral corial margins weakly convex; hind tarsal segment 3 slightly longer than segment 2, segment 2 slightly longer than segment 1; supracoxal lobes of prothorax projecting. GENITALIA (Figs. 9E, F, 10): Vesica: Body relatively slender; apical spine with distinct subbasal notch on dorsal surface, flange on dorsal surface of vesica truncate at level of base of secondary gonopore. Phallotheca: As in generic description. Left Paramere: Anterior process very short, not extending beyond shaft of paramere, apparently devoid of setae; posterior process greatly elongate, elevated above dorsal margin of body of paramere. Right Paramere: Broadly ovoid, apex at most weakly acuminate and lacking distinct fingerlike process found in other Eminoculus spp.

Female: Small-sized, relatively stout bodied; total length 2.16-2.65; length apex clypeus-apex wing 1.86-2.24; width pronotum 0.94-1.06. COLORATION (Fig. 1): As in males. SURFACE AND VESTITURE: As in males. STRUCTURE: Abdomen rounded or truncate apically, not elongate and acuminate, only slightly projecting beyond apex of hemelytra. GENITALIA: See generic description.

[FIGURE 9 OMITTED]

HOSTS: Drosanthemum spp. (Fig. 2E) and Antimima cf. koekenaapensis (Aizoaceae: Ruschioideae). The single specimen recorded from Leipoldtia cf. schultzei (Aizoaceae: Ruschioideae) is almost certainly the result of misassociation with the host during the collecting process.

DISTRIBUTION (Fig. 4): The most widely distributed of the known Eminoculus spp., ranging broadly across Namaqualand and the Little Karoo. It is our judgement that the specimens

from Pretoria are all introduced on ornamental plantings and do not represent part of the natural range.

DISCUSSION: Schuh (1974) originally excluded several specimens from the paratype series on which the original description of E. drosanthemi was based because they differed in the structure of the head, surface texture of the pronotum, and type of pubescence. The availability of a much larger sample of material over a much broader natural range has caused us to conclude that all of the very small highly polished specimens of Eminoculus belong to a single species, E. drosanthemi.

[FIGURE 10 OMITTED]

PARATYPES: SOUTH AFRICA: Gauteng (formerly Transvaal): Meintjies Kop, Pretoria, 25.737[degrees]S 28.212[degrees]E, 1,419 m, 22 Oct 1967-22 Oct 1967, J. and S. Slater, Drosanthemum floribundum (Aizoaceae), 5[male] (AMNH_PBI 00172851-AMNH_PBI 00172855), 11[female] (AMNH_PBI 00172836-AMNH_PBI 00172846), 5[male] (AMNH_PBI 00172851-AMNH_PBI 00172855), 11[female] (AMNH_PBI 00172836-AMNH_PBI 00172846) (AMNH). National Botanical Garden, Pretoria, 25.737[degrees]S 28.212[degrees]E, 1,419 m, 28 Dec 1967, J.A. & S. Slater and R.T. Schuh, (Aizoaceae), 1[female] (AMNH_PBI 00172849) (AMNH). Pretoria, 25.7069[degrees]S 28.2294[degrees]E, 24 Oct 1967-24 Oct 1967, J.A. & S. Slater and R.T. Schuh, Drosanthemum floribundum (Harv.) Gilg (Aizoaceae), 1[male] (AMNH_PBI 00172856) (AMNH).

OTHER SPECIMENS EXAMINED: SOUTH AFRICA: Gauteng (formerly Transvaal): Pretoria, 25.7069[degrees]S 28.2294[degrees]E, 09 Jun 1966-09 Jun 1966, G. du Plessis, Drosanthemum floribundum (Aizoaceae), 23[male] (AMNH_PBI 00186408-AMNH_PBI 00186430), 5[male] (AMNH_PBI 00186431-AMNH_PBI 00186435) (PPRI_ENT). Northern Cape: 135 km S of Kamieskroon on N7, 31.19567[degrees]S 18.48209[degrees]E, 365 m, 28 Oct 2003, Schuh, Cassis, Weirauch, Drosanthemum sp. (Aizoaceae), det. P. Chesselet voucher-NYBG, 1[male] (AMNH_PBI 00170835), 1[female] (AMNH_PBI 00170836) Phyllobolus nitidus (Haw.) Gerbaulet (Aizoaceae), det. P. Chesselet voucher-NYBG, 5[male] (AMNH_PBI 00172917-AMNH_PBI 00172921), 9[female] (AMNH_PBI 00172922-AMNH_PBI 00172930) (AMNH). 8.1 km SW of N7 toward Messelpad, 29.80868[degrees]S 17.7802[degrees]E, 745 m, 08 Sep 2004, Schuh, Schwartz, Henry, Wyniger, Forero, Leipoldtia cf. schultzei (Schltr. & Diels) Friedrich (Aizoaceae), det. Field ID, 1[male] (AMNH_PBI 00172862) (AMNH). NW of Gaffes toward Hondeklip Bay, 30.55718[degrees]S 17.9257[degrees]E, 410 m, 09 Sep 2004, Schuh, Schwartz, Henry, Wyniger, Forero, Drosanthemum sp. (Aizoaceae), det. P. Chesselet VOUCHER-NYBG, 1[female] (AMNH_PBI 00171033) (AM), 1[male] (AMNH_PBI 00170981), 1[female] (AMNH_PBI 00171032) (AM). 36[male] (AMNH_PBI 00170957-AMNH_PBI 00170977, AMNH_PBI 00170982-AMNH_PBI 00170996), 53[female] (AMNH_PBI 00170997-AMNH_PBI 00171029, AMNH_PBI 00171035-AMNH_PBI 00171054) (AMNH). 1[male] (AMNH_PBI 00171031) (SAMC). 1[male] (AMNH_PBI 00170978), 1[female] (AMNH_PBI 00171030) (SANC). 1[male] (AMNH_PBI 00170980), 1[female] (AMNH_PBI 00171034) (ZISP). NW of Gaffes toward Hondeklip Bay, 30.39854[degrees]S 17.53129[degrees]E, 70 m, 09 Sep 2004, Schuh, Schwartz, Henry, Wyniger, Forero, Drosanthemum sp. (Aizoaceae), det. Field ID, 11[male] (AMNH_PBI 00171369-AMNH_PBI 00171379), 8[female] (AMNH_PBI 00171059-AMNH_PBI 00171066) (AMNH). 8[male] (AMNH_PBI 00190241-AMNH_PBI 00190248), 21[female] (AMNH_PBI 00190249-AMNH_PBI 00190269) (USNM). Western Cape: 16 km NE of Vanrhynsdorp on R27, 31.52003[degrees]S 18.87532[degrees]E, 190 m, 12 Sep 2004, Schuh, Schwartz, Henry, Wyniger, Forero, Drosanthemum sp. (Aizoaceae), det. P. Chesselet voucher-NYBG, 2[male] (AMNH_PBI 00171094, AMNH_PBI 00171095), 2[female] (AMNH_PBI 00171194, AMNH_PBI 00171195) (AM). 52[male] (AMNH_PBI 00171067-AMNH_PBI 00171081, AMNH_PBI 00171085, AMNH_PBI 00171087-AMNH_PBI 00171092, AMNH_PBI 00171100-AMNH_PBI 00171129), 91[female] (AMNH_PBI 00171130-AMNH_PBI 00171184, AMNH_PBI 00171189, AMNH_PBI 00171196-AMNH_PBI 00171230) (AMNH). 5[male] (AMNH_PBI 00171082-AMNH_PBI 00171084, AMNH_PBI 00171086, AMNH_PBI 00171093) (CNC). 2[male] (AMNH_PBI 00171098, AMNH_PBI 00171099), 2[female] (AMNH_PBI 00171192, AMNH_PBI 00171193) (SAMC). 2[male](AMNH_PBI 00171096, AMNH_PBI 00171097), 2[female] (AMNH_PBI 00171190, AMNH_PBI 00171191) (SANC). 30[male] (AMNHPBI 00190126-AMNH_PBI 00190155), 58[female] (AMNH_PBI 00190156-AMNH_PBI 00190213) (USNM). 4[female] (AMNH_PBI 00171185-AMNH PBI 00171188) (ZISP). 2 km SW of Lutzville on R362, 31.58223[degrees]S 18.30587[degrees]E, 66 m, 13 Sep 2004, Schuh, Schwartz, Henry, Wyniger, Forero, Drosanthemum sp. (Aizoaceae), det. Field ID, 6[male] (AMNH_PBI 00171231-AMNH_PBI 00171236), 23[female] (AMNH_PBI 00171237-AMNH_PBI 00171259) (AMNH). 34 km SW of Nuwerus on N7, 31.34771[degrees]S 18.60776[degrees]E, 222 m, 04 Sep 2004, Schuh, Schwartz, Henry, Wyniger, Forero, Antimima cf. koekenaapensis (L.Bolus) H.E.K. Hartmann (Aizoaceae), det. P. Chesselet VOUCHER-NYBG, 1[male] (AMNH_PBI 00170837), 7[female] (AMNH_PBI 00170838-AMNH_PBI 00170844) (AMNH). 4 km S of Nuwerus on N7, 31.16744[degrees]S 18.39917[degrees]E, 332 m, 04 Sep 2004, Schuh, Schwartz, Henry, Wyniger, Forero, Antimima cf. koekenaapensis (L.Bolus) H.E.K. Hartmann (Aizoaceae), det. Field ID, 18[male] (AMNH_PBI 00170845-AMNH_PBI 00170862), 94[female] (AMNH_PBI 00170863-AMNH_PBI 00170956) (AMNH). 7[male] (AMNH_PBI 00190214-AMNH_PBI 00190220), 20[female] (AMNH PBI 00190221-AMNH_PBI 00190240) (USNM). 44.6km NE of Barrydale on R62, 33.64883[degrees]S 21.0198[degrees]E, 324 m, 02 Nov 2003, Schuh, Cassis, Weirauch, Drosanthemum sp. (Mesembryanthemaceae), det. P. Chesselet voucher-NYBG, 15[male] (AMNH_PBI 00172908-AMNH_PBI 00172911, AMNH_PBI 00172931-AMNH_ PBI 00172941), 13[female] (AMNH_PBI 00172912-AMNH_PBI 00172915, AMNH_PBI 00172942-AMNH_PBI 00172950) (AMNH). 7.8 km W of Barrydale on R62, 33.86013[degrees]S 20.78365[degrees]E, 496 m, 02 Nov 2003, Schuh, Cassis, Weirauch, Drosanthemum sp. (Mesembryanthemaceae), det. P. Chesselet voucher-NYBG, 1[male] (AMNH_PBI 00172951), 1 nymph (AMNH_PBI 00172952) (AMNH). Knysna Head, 34.03[degrees]S 23.03[degrees]E, 09 Feb 1968-09 Feb 1968, J.A. & S. Slater, T. Schuh, M. Sweet, (Aizoaceae), 1[female] (AMNH_PBI 00172850) (AMNH).

Eminoculus hirsutus Schuh

Figures 1, 4

Eminoculus hirsutus Schuh, 1974: 162, Fig. 66 (n. sp.).

DIAGNOSIS: Males recognized by the large size, long erect setae on the head and pronotum, the strongly stylate eyes, and straight to weakly convex hemelytral margins (Fig. 1). Similar in size to E. rugosus and E. auratus. Distinguished from the former by the more exaggerated nature of the stylate eyes in E. hirsutus. Distinguished from the latter by the straight-sided to weakly convex lateral margin of the hemelytra in E. hirsutus and the weakly convex condition in E. auratus. Distinguished from E. laevigatus by the setae on the head and pronotum being sparse, only moderately long, and inconspicuous in that species.

REDESCRIPTION (adapted from Schuh, 1974): Male: Total length 3.72, width pronotum 1.20. COLORATION (Fig. 1): Body and appendages generally black; ventral margin of mesepimeron and metepimeron, all femora distally, all tibiae, and tarsal segments 1 and 2 tan or light brown; membrane dark smoky brown. SURFACE AND VESTITURE: All coxae mostly pruinose; head, pronotum, and scutellum rugulose, polished and shining; hemelytra faintly rugulose, dull; remainder of body, finely granulose, dull; frons, vertex, head ventrad of eyes and dorsad of bucculae, head, pronotum, scutellum, and basal region of hemelytra with long, sericeous, erect or semierect setae; hemelytra with numerous, venter of thorax and abdomen with less numerous, appressed, pale setae; antennae with dense decumbent vestiture, segment 1 with several fine black spines on medial surface; femora with few, long, erect setae, particularly on ventral surface; tibiae with semierect, black spines. STRUCTURE: Head: Short, transverse, eyes strongly stylate, projecting laterally beyond anterolateral margins of pronotum by distance on each side about equal to three-fourths width of anterior margins of pronotum; eyes nearly spherical; posterior margin of vertex forming fine, rounded carina medially, grading into cylindrical eye stalks laterally; vertex depressed on either side of midline anterior to posterior margin; antennal segment 1 parallel sided and relatively long; labium just surpassing forecoxae. Thorax: Pronotum with anterior margin nearly straight, lateral and posterior margins sinuate; anterior margin of pronotum with distinct flattened collar about as wide as diameter of antennal segment 1; pronotum with very deep, wide, transverse impression medially; calli elevated, pronounced, largely confluent; cuneal incisure shallow, fracture angled anteromedially; hind tarsal segment 1 about one-third length of segment 2, segment 3 about two-thirds length of segment 2; hind femora weakly bowed. GENITALIA: not examined

Female: unknown.

HOST: Unknown.

DISTRIBUTION (Fig. 4): Known only from the type locality near Oudtshoorn, in the Western Cape Province.

DISCUSSION: The single known specimen of this species has not been re-examined during the course of this study.

Holotype: SOUTH AFRICA: Western Cape: Oudtshoorn, Zebra, 33.5833[degrees]S 22.2[degrees]E, 01 Oct 1951, Museum Expedition, 1[male] (SAMC).

Eminoculus laevigatus, new species

Figures 1, 3, 4, 11

DIAGNOSIS: Recognized by the moderate to large size, the elongate body in the males, posteriorly protruding, acuminate abdomen in the female, the strongly stylate eyes with a weakly curving posterior margin in the male, and the highly polished dorsum with inconspicuous, scattered, erect or semi-erect setae on the head and pronotum in contract to the short, recumbent weakly golden setae on the scutellum, and hemelytra (Fig. 1). Both sexes readily separated from E. auratus and E. hirsutus by the sparse erect or semierect setae of moderate length on head and pronotum in laevigatus, whereas E. auratus and E. hirsutus with numerous long, silver, erect setae on head and pronotum. Males and females distinguished from E. rugosus by the distinctly golden, subappressed vestiture covering the entire dorsum in that species and the yellow proepisternal supracoxal lobe of E. laevigatus, in contrast to black or dark brown proepisternal supracoxal lobe in E. rugosus. Both sexes separated from E. lanuginosus by the dense woolly setae covering the entire dorsum in that species. Males and females distinguished from E. drosanthemi by their larger size and the nearly straight posterior margin of the head and eyes in E. laevigatus. Posteriorly acuminate abdomen in female similar to that in E. rugosus and E. auratus.

DESCRIPTION: Male: Total length 3.18-3.50; length apex clypeus-cuneal fracture 2.16-2.37; width pronotum 1.02-1.17. COLORATION (Fig. 1): Body black; femora black with pale apex; tibiae moderately infuscate to largely golden; tarsi black; margin of proepisternal supracoxal lobe yellow. SURFACE AND VESTITURE: Head, and pronotum highly polished and shining and very weakly rugose; hemelytra matte; scutellum transversely rugose, dull; antennal segment 1 with a few slender, erect spines on medial surface; head and pronotum with sparse, moderately long, simple, semierect, silvery setae; hemelytra with numerous, short, simple, appressed, weakly golden setae; genal region dorsad of bucculae with long, erect, fine setae; ~4 hairs arising from lateroventral surface of claw. STRUCTURE: Head: Eyes stylate, posteromedial angle of eye removed from anterolateral angle of pronotum by about diameter of antennal segment 1; posterior margin of vertex and eyes nearly linear; antennal segment 1 weakly swollen, parallel sided, relatively short; labium reaching to posterior margin of mesepisternum. Thorax: Anterior margin of pronotum with distinct flattened collar about as wide as diameter of antennal segment 1; calli distinctly elevated, not conspicuously confluent; pronotum convex transversely, inclined posteriorly to mesoscutum, lateral margins very slightly concave, posterior margin shallowly concave; supracoxal lobes of prothorax projecting; costal hemelytral margin nearly straight over most of length. GENITALIA (Fig. 11): Vesica: Body relatively slender; apical spine with weak subbasal notch on dorsal surface; margin of flange on dorsal surface of vesica flowing smoothly to base of secondary gonopore. Phallotheca: As in generic description. Left Paramere: Anterior process modestly shorter than posterior, slightly extending beyond shaft of paramere, with tight clump of a few setae apically; posterior process weakly elevated above dorsal margin of body of paramere, tapering apically, apex fingerlike. Right Paramere: Irregular elongate ovoid, apex with a distinct fingerlike process located to left of center.

[FIGURE 11 OMITTED]

Female: Medium to large-sized, elongate; total length 3.04-3.35; length apex clypeus-apex wing 2.32-2.64; width pronotum 1.00-1.09. COLORATION (Fig. 1): Femora and tarsal segment 3 black; tibiae and tarsal segments 1 and 2 pale. SURFACE AND VESTITURE: As in male. STRUCTURE: Hemelytra broadened posteriorly, abdomen broadened medially and distinctly acuminate apically, projecting well beyond posterior margin of hemelytra. GENITALIA (Fig. 11): As in generic description.

[FIGURE 12 OMITTED]

ETYMOLOGY: From the Latin laevigatus, smooth, for the condition of the dorsum, including the hemelytra.

HOSTS: Leipoldtia schultzei (Schltr. & Diels) Friedrich (Fig. 3A, B), Ruschia muelleri (L. Bolus) Schwantes, and R. stricta L. Bolus (Aizoaceae: Ruschioideae).

DISTRIBUTION (Fig. 4): Northern Namaqualand, ranging from the area around Kamieskroon north to the latitude of Port Nolloth.

HOLOTVPE: SOUTH AFRICA: Northern Cape: 8.1 km SW of N7 toward Messelpad, 29.80868[degrees]S 17.7802[degrees]E, 745 m, 08 Sep 2004, Schuh, Schwartz, Henry, Wyniger, Forero, Leipoldtia cf. schultzei (Schltr. & Diels) Friedrich (Aizoaceae), det. Field ID, 1[male] (AMNH_PBI 00171563) (SANC).

PARATYPES: SOUTH AFRICA: Northern Cape: 15.7 km SW of Soebatsfontein on Wallekraal Rd (SW of Kamieskroon), 30.24064[degrees]S 17.55599[degrees]E, 258 m, 26 Oct 2003, Schuh, Cassis, Weirauch, (Aizoaceae), 7[male] (AMNH_PBI 00171551-AMNH_PBI 00171557), 2[female] (AMNH_PBI 00171558, AMNH_PBI 00171559) (AMNH). 19km E of Port Nolloth on R382, 29.30934[degrees]S 17.06537[degrees]E, 168 m, 06 Sep 2004, Schuh, Schwartz, Henry, Wyniger, Forero, Ruschia stricta L.Bolus (Aizoaceae), det. P. Chesselet VOUCHERNYBG, 2[male] (AMNH_PBI 00171390, AMNH_PBI 00171391), 7[female] (AMNH_PBI 00171392-AMNH_PBI 00171398) 6[male] (AMNH_PBI 00190438-AMNH_PBI 00190443), 2[female] (AMNH_PBI 00190444, AMNH_PBI 00190445) (USNM). 21 km W of Springbok on R355, 29.69466[degrees]S 17.71383[degrees]E, 516 m, 05 Sep 2004, Schuh, Schwartz, Henry, Wyniger, Forero, Ruschia muelleri (L.Bolus) Schwantes (Aizoaceae), det. P. Chesselet voucher-NYBG, 1[male] (AMNH_PBI 00171594) (AMNH). 4.5 km SW of N7 toward Messelpad, 29.78873[degrees]S 17.80387[degrees]E, 733 m, 08 Sep 2004, Schuh, Schwartz, Henry, Wyniger, Forero, Leipoldtia cf. schultzei (Schltr. & Diels) Friedrich (Aizoaceae), det. Field ID, 1[male] (AMNH_PBI 00171560), 1[female] (AMNH_PBI 00171561) (AMNH). 4.7 km W of Springbok on R355, 29.68108[degrees]S 17.84683[degrees]E, 866m, 05 Sep 2004, Schuh, Schwartz, Henry, Wyniger, Forero, Leipoldtia schultzei (Schltr. & Diels) Friedrich (Aizoaceae), det. P. Chesselet voucher-NYBG, 2[female] (AMNH_PBI 00171549, AMNH_PBI 00171550) (AMNH). 6.5km S of Kamieskroon, 30.17987[degrees]S 17.86935[degrees]E, 836 m, 27 Oct 2003, Schuh, Cassis, Weirauch, Leipoldtia schultzei (Schltr. & Diels) Friedrich (Aizoaceae), det. Field ID, 2[male] (AMNH_PBI 00172953, AMNH_PBI 00172954) (AMNH). 7 km NW of Kamieskroon, Agtertuin Farm, 30.17725[degrees]S 17.90125[degrees]E, 725m, 10 Sep 2004, Schuh, Schwartz,

Henry, Wyniger, Forero, Leipoldtia schultzei (Schltr. & Diels) Friedrich (Aizoaceae), det. Field ID, 28[male] (AMNH_PBI 00171702-AMNH_PBI 00171729), 120[female] (AMNH_PBI 00171730-AMNH_PBI 00171786, AMNH_PBI 00171791-AMNH_PBI 00171853) (AMNH). 4[male] (AMNH_PBI 00190361-AMNH_PBI 00190364), 70[female] (AMNH_PBI 00190365-AMNH_PBI 00190434) (USNM). 4[female] (AMNH PBI 00171787-AMNH_PBI 00171790) (ZISP). 7 km NW of Kamieskroon, Agtertuin Farm, 30.16869[degrees]S 17.90032[degrees]E, 690 m, 25 Oct 2003, Schuh, Cassis, Weirauch, Leipoldtia schultzei (Schltr. & Diels) Friedrich (Aizoaceae), det. P. Chesselet voucher-NYBG, 3[male] (AMNH_PBI 00058634, AMNH_PBI 00171468-AMNH_PBI 00171469), 6[female] (AMNH_PBI 00171480-AMNH_PBI 00171485) (AM). (Aizoaceae), det. J Manning voucher-NYBG, 2[male] (AMNH_PBI 00171410, AMNH PBI 00171411), Leipoldtia schultzei (Schltr. & Diels) Friedrich (Aizoaceae), det. P. Chesselet VOUCHER-NYBG, 81[male] (AMNH_PBI 00101883-AMNH_PBI 00101906, AMNH_PBI 00171412-AMNH_PBI 00171454, AMNH_PBI 00171459, AMNH_PBI 00171470-AMNH_PBI 00171471, AMNH_PBI 00172955-AMNH_PBI 00172965), 144[female] (AMNH PBI 00101907-AMNH_PBI 00101991, AMNH_PBI 00171486-AMNH_PBI 00171487, AMNH PBI 00171492-AMNH_PBI 00171548) (AMNH). 3[female] (AMNH PBI 00171477-AMNH_PBI 00171479) (ANIC). 3[male] (AMNH_PBI 00171455, AMNH_PBI 00171466-AMNH_PBI 00171467), 4[female] (AMNH_PBI 00171488-AMNH_PBI 00171491) (CNC). 3[male] (AMNH_PBI 00171463-AMNH_PBI 00171465) (SAMC). 3[male] (AMNH_PBI 00171460-AMNH_PBI 00171462), 5[female] (AMNH_PBI 00171472-AMNH_PBI 00171476) (SANC). 3[male] (AMNH_PBI 00171456-AMNH_PBI 00171458) (ZISP). 7 km NW of Kamieskroon, Agtertuin Farm, 30.1682[degrees]S 17.90032[degrees]E, 668 m, 10 Sep 2004, Schuh, Schwartz, Henry, Wyniger, Forero, Leipoldtia schultzei (Schltr. & Diels) Friedrich (Aizoaceae), det. P. Chesselet VOUCHER-NYBG, 4[male] (AMNH_PBI 00171674-AMNH_PBI 00171677), 24[female] (AMNH_PBI 00171678-AMNH_PBI 00171701) (AMNH). 2[female] (AMNH_PBI 00190436, AMNH PBI 00190437) (USNM). 8.1 km SW of N7 toward Messelpad, 29.80868[degrees]S 17.7802[degrees]E, 745m, 08 Sep 2004, Schuh, Schwartz, Henry, Wyniger, Forero, Leipoldtia cf. schultzei (Schltr. & Diels) Friedrich (Aizoaceae), det. Field ID, 15[male] (AMNH_PBI 00171562, AMNH_PBI 00171564-AMNH_PBI 00171577), 96[female] (AMNH_PBI 00171578-AMNH_PBI 00171673) (AMNH). ~15 km S of Springbok off N7, 29.76933[degrees]S 17.8305[degrees]E, 840 m, 08 Sep 2004, Schuh, Schwartz, Henry, Wyniger, Forero, Leipoldtia of. schultzei (Schltr. & Diels) Friedrich (Aizoaceae), det. P. Chesselet VOUCHER-NYBG, 1[male] (AMNH_PBI 00171380), 9[female] (AMNH_PBI 00171381-AMNH_PBI 00171389) (AMNH).

[FIGURE 13 OMITTED]

[FIGURE 14 OMITTED]

Eminoculus lanuginosus, new species

Figures 1, 3, 4, 12

DIAGNOSIS: Both males and females recognized by their small size and dense covering of long, silvery, woolly setae on the head, pronotum, and hemelytra (Fig. 1). Abdomen in female more or less truncate apically and not strongly projecting beyond posterior margin of hemelytra (Fig. 1). Size and general structure most similar to E. drosanthemi. Dorsum of E. drosanthemi with evenly distributed, sparse, extremely short black vestiture, whereas both sexes of E. lanuginosus with dense, long, silvery, woolly vestiture on head, pronotum, and hemelytra; femora in E. drosanthemi black, not golden or light brown with variable maculae, as in males and females of E. lanuginosus. Vesica in E. lanuginosus sharing with E. atrisetosus and E. drosanthemi the presence of a distinct subbasal notch on the dorsal surface of the apical spines and a truncate or toothed flange just proximal to secondary gonopore on the dorsal side of the vesica. Distinguished from remaining species of Eminoculus other than E. drosanthemi by their much larger size, males of E. atrisetosus averaging 2.60 in length, remaining spp. averaging at least 3.19 in length.

DESCRIPTION: Male: Total length 2.58-2.63; length apex clypeus-cuneal fracture 1.79-1.84; width pronotum 0.97-1.02. COLORATION (Fig. 1): Dorsum black; antennae golden; femora golden or light brown with variable dark maculae; tibiae golden; tarsi with segments 1 and 2 pale, 3 dark; margins of supracoxal lobes yellow. SURFACE AND VESTITURE: Dorsum polished, shining black; hemelytra variably rugulose; posterior lobe of pronotum rugose, polished and shining; calli smooth and highly polished; antennal segment 1 with few slender, erect spines on medial surface (<15); genal region dorsad of bucculae with many long, erect, fine setae; head, pronotum, scutellum, and anterior half of corium and clavus with dense silvery woolly setae; posterior half of corium and clavus with setae more closely appressed; ~3 hairs arising from lateroventral surface of claw. STRUCTURE: Head: Eyes distinctly stylate; posterior margin of eyes not contiguous with anterior pronotal margin; posterior margin of head and eyes linear or weakly curvilinear; antennal segment 1 weakly swollen, parallel sided, relatively short; labium reaching to posterior margin of mesepisternum. Thorax: Anterior margin of pronotum with distinct flattened collar about as wide as diameter of antennal segment 1; calli weakly elevated and not confluent; pronotum nearly flat longitudinally, weakly convex transversely, posterior margin weakly concave; proepisternal supracoxal lobe weakly projecting; hemelytra with costal margin weakly convex. GENITALIA (Fig. 12): Vesica: Body relatively slender; apical spine with distinct subbasal notch on dorsal surface, flange on dorsal surface of vesica truncate at level of base of secondary gonopore. Phallotheca: As in generic description. Left Paramere: Anterior process relatively short, barely extending beyond shaft of paramere and apparently devoid of setae; posterior process elongate, very slightly elevated above dorsal margin of body of paramere. Right Paramere: Irregular elongate ovoid, apex with a small centrally-located fingerlike process.

Female: Relatively small-sized, globose in outline; total length 2.29-2.62; length apex clypeusapex wing 2.04-2.29; width pronotum 0.99-1.09. COLORATION (Fig. 1): Femora golden with variable dark maculae; tibiae and tarsal segments 1 and 2 golden, segment 3 dark. SURFACE AND VESTITURE: Dorsum, including exposed portion of abdomen with dense, long, silvery, woolly vestiture; areas proximal to claval commissure typically without setae. STRUCTURE: Abdomen ovoid and not distinctly acuminate apically. GENITALIA: See generic description.

ETYMOLOGY: From the Latin lanuginosus, woolly, for the condition of the vestiture of the dorsum.

HOSTS: Delosperma spp. (Fig. 3C, D) and Drosanthemum sp. (Aizoaceae: Mesembryanthemoideae)

DISTRIBUTION (Fig. 4): Apparently restricted to the Little Karoo. Known only from localities near Barrydale and Herbertsdale.

HOLOTYPE: SOUTH AFRICA: Western Cape: 28.1 km NW of N2 on R327 toward Herbertsdale, 34.08242[degrees]S 21.77657[degrees]E, 37 m, 06 Nov 2003, Schuh, Cassis, Weirauch, Delosperma sp. (Aizoaceae), det. P. Chesselet VOUCHER-NYBG, 1[male] (AMNH_PBI 00058640) (SANC).

PARATYPES: SOUTH AFRICA: Western Cape: 28.1 km NW of N2 on R327 toward Herbertsdale, 34.08242[degrees]S 21.77657[degrees]E, 37 m, 06 Nov 2003, Schuh, Cassis, Weirauch, Delosperma sp. (Aizoaceae), det. P. Chesselet VOUCHER-NYBG, 1[female] (AMNH_PBI 00171404) (AM). 2[male] (AMNH_PBI 00171399, AMNH_PBI 00171400), 6[female] (AMNH_PBI 00171401, AMNH_PBI 00171405-AMNH_PBI 00171409) (AMNH). 1[female] (AMNH_PBI 00171402) (SAMC). 2[female] (AMNH_PBI 00058641, AMNH_PBI 00171403) (SANC). 44.6 km NE of Barrydale on R62, 33.64883[degrees]S 21.0198[degrees]E, 324 m, 02 Nov 2003, Schuh, Cassis, Weirauch, Drosanthemum sp. (Mesembryanthemaceae), det. P. Chesselet voucher-NYBG, 1[male] (AMNH_PBI 00172916) (AMNH).

Eminoculus rugosus, new species

Figures 1, 3, 4, 12, 14

DIAGNOSIS: Recognized by the large size, the dull hemelytra, the recumbent golden setae on the hemelytra, the nearly straight posterior margin of the stylate eyes, and the apically pointed abdomen in the female extending well beyond the posterior margin of the hemelytra (Fig. 1). Structure of eyes similar to E. laevigatus, but that species somewhat smaller (mean length 3.34 as compared to 3.76) with the head and pronotum strongly polished and shining. Similar in size to E. auratus and E. hirsutus, but distinguished the more exaggerated nature of the stylate eyes and the highly polished head and pronotum in both of those species.

DESCRIPTION: Male: Total length 3.63-3.89; length apex clypeus-cuneal fracture 2.46-2.7; width pronotum 1.25-1.31. COLORATION (Fig. 1): Body and appendages generally black; femora with apex paler than remainder, tarsal segment 2 pale; proepisternal supracoxal lobe varying from yellow to dark brown or black. SURFACE AND VESTITURE: Head not polished; posterior lobe of pronotum rugose (Fig. 13A), polished and shining, contrasting with smooth and highly polished calli; hemelytra variably rugulose and dull; antennal segment 1 with a few(<15) slender, erect spines on medial surface; genal region dorsad of bucculae with many long, erect, very fine setae; head and pronotum with long, sericeous, semierect, golden setae; hemelytra with dense, moderately long, sericeous, recumbent, golden setae; ~4-5 hairs arising from lateroventral surface of claw (Fig. 13C, D). STRUCTURE: Head: Eyes stylate; posteromedial angle of eye removed from anterolateral angle of pronotum by distance about equal to diameter of antennal segment 1; posterior margin of eyes nearly straight, posterior margin of vertex concave across anterior margin of pronotum; antennal segment 1 weakly swollen, parallel sided, relatively short; labium reaching to trochanteral joint of forecoxae. Thorax: Anterior margin of pronotum with distinct flattened collar about as wide as diameter of antennal segment 1; calli pronounced and moderately elevated, confluent anteriorly; pronotum convex transversely, inclined posteriorly, lateral pronotal margins very slightly concave, posterior margin weakly concave; proepisternal supracoxal lobe projecting; costal hemelytral margin nearly straight over most of length. GENITALIA (Fig. 13B, 14): Vesica: Body relatively slender; apical spine greatly elongate and slender, with very weak subbasal notch on dorsal surface; margin of flange on dorsal surface of vesica flowing smoothly to base of secondary gonopore. Phallotheca: As in generic description. Left Paramere: Anterior process moderately elongate and relatively broad, with diffuse clump of setae apically; posterior process relatively short, only very slightly longer than anterior, not elevated above dorsal margin of body of paramere, tapered apically. Right Paramere: Elongate ovoid, apex with distinct fingerlike process located to left of center.

Female: Relatively large, broad bodied; total length 3.97-4.19; length apex clypeus-apex wing 2.81-3.17; width pronotum 1.27-1.38. COLORATION: As in male, except femora variably infuscate; tibiae and tarsal segment 2 golden to pale. SURFACE AND VESTITURE: As in males. STRUCTURE: Abdomen broadest just posterior to midpoint, acuminate apically, projecting well beyond posterior margin of hemelytra. GENITALIA: See generic description.

ETYMOLOGY: From the Latin rugosus, wrinkled, in reference to the texture of the posterior lobe of the pronotum.

HOSTS: Sterile sample (Aizoaceae/Mesembryanthemaceae) (Fig. 3E, F).

DISTRRIUTION (Fig. 4): Known from a single locality in southern Namaqualand.

HOLOTYPE: SOUTH AFRICA: Western Cape: Doring Bay to Heerenlogement, 31.98387[degrees]S 18.55379[degrees]E, 170 m, 13 Sep 2004, Schuh, Schwartz, Henry, Wyniger, Forero, (Aizoaceae), det. P. Chesselet VOUCHER-NYBG, 1[male] (AMNH_PBI 00171290) (SANC).

PARATYPES; SOUTH AFRICA: Western Cape: Doting Bay to Heerenlogement, 31.98387[degrees]S 18.55379[degrees]E, 170 m, 13 Sep 2004, Schuh, Schwartz, Henry, Wyniger, Forero, (Aizoaceae), det. P. Chesselet voucher-NYBG, 3[male] (AMNH_PBI 00171285-AMNH_PBI 00171287), 3[female] (AMNH_PBI 00171319-AMNH_PBI 00171321) (AM). 22[male] (AMNH_PBI 00171260-AMNH_PB1 00171273, AMNH_PB100171276-AMNH_PB1 00171281, AMNH_ PBI 00171294-AMNH_PBI 00171295), 54[female] (AMNH_ PBI 00171296-AMNH_PBI 00171315, AMNH_ PBI 00171322-AMNH PBI 00171323, AMNH_PBI 00171331-AMNH_PB1 00171362) (AMNH). 2[male] (AMNH_PBI 00171274, AMNH_PBI 00171275), 2[female] (AMNH_PBI

00171324, AMNH_PB1 00171325) (CNC). 3[male] (AMNH_PB1 00171282-AMNH_PBI 00171284), 3[female] (AMNH PBI 00171316-AMNH PBI 00171318) (SAMC). 3[male] (AMNH_PBI 00171291AMNH_PB1 00171293), 3[female] (AMNH_PBI 00171328AMNH_PBI 00171330) (SANC). Sterile material sp. (Aizoaceae), det. P. Chesselet voucher-NYBG, 22[male] (AMNH PBI 00190453-AMNH_PBI 00190474), 13[female] (AMNH_PBI 00190475-AMNH_PBI 00190487) (USNM). (Aizoaceae), det. P. Chesselet voucher-NYBG, 2[female] (AMNH_PBI 00171288, AMNH PB1 00171289), 2[female] (AMNH_PBI 00171326, AMNH_PBI 00171327) (ZISP).

PHYLOGENETIC ANALYSIS AND DISCUSSION

The data shown in tables 2 and 3 were analyzed with NONA (Goloboff, 1998) using the mult* (with 10 iterations) and max* branch swapping commands; the hold command was set at 10,000, allowing space for a maximum of 10,000 trees. Treating all characters as unordered, two trees were found with a length of 121 steps, consistency index of 71, and a retention index of 63. The strict consensus of those two trees collapses E. drosanthemi and E. lanuginosus to a single node. Subjecting those two trees to successive weighting produces a single tree. Analysis of the same data with PIWE (Goloboff, 1993, 1997), which produces best fit trees by maximizing the sum of the average unit consistency index--using the same set of commands as above--produced the same single tree with a fit statistic of 376.5. Thus, in figure 15 we have shown this single fully resolved tree with unambiguous optimizations of character data, as provided by the program WinClada (Nixon, 2000).

As can be seen from figure 15, there is extensive character support for recognition of Eminoculus with an updated diagnosis as well as for the subgroup fitting the diagnosis originally provided by Schuh (1974). Although most of the groupings within Eminoculus are relatively clearcut, the relationship between the phylogenetic result and the known distributions of the species is not so clear. For example, E. auratus and E. hirsutus, which are obviously closely related, have a combined distribution that extends to the virtual extremes of the distribution of the group. Eminoculus drosanthemi is widespread, its known distribution also encompassing virtually the total range of the genus.

Comparison of the distributions of Eminoculus spp. with the distributions of their known host groups shows a strong correspondence between the centers of diversity in the two taxa--Namaqualand and the Little Karoo. Nonetheless, many mesem genera have distributions that extend into the western margins of the Great Karoo, an area that remains virtually unsampled for the Miridae in general and Eminoculus in particular. Nonetheless, genera such and Ruschia have a wider distribution in South Africa and it is possible that Eminoculus may track that pattern.

Further collecting should also help to clarify details in patterns of host association. Current data strongly ally all Eminoculus spp. to the "mesem" clade within the Aizoaceae. The absence of precise host identifications for E. rugosus, and the limited data for most species, limit our ability to determine if there is a more detailed correlation between phylogenetic relationships and patterns of host association.

ACKNOWLEDGMENTS

Field work for this project was supported by a National Science Foundation (NSF) Planetary Biodiversity Inventories award (DEB-0316495) to Randall T. Schuh and Gerasimos Cassis for the study of the Miridae subfamilies Orthotylinae and Phylinae. Coauthor Grace Wu was supported as a summer intern during 2006 with funding from a NSF Research Experiences for Undergraduates award (0353817) to the American Museum of Natural History.

Host plants were identified by the staff at the Compton Herbarium, Kirstenbosch Gardens, Cape Town, South Africa. We especially thank Edwina Marinus for her assistance in coordinating the acquisition of these invaluable data and Pascale Chesselet for the actual identifications.

Steve Thurston inked the genitalic illustrations, prepared their final digital renderings, and assembled all plates in digital form for publication. Michael D. Schwartz prepared the maps. Comments by Christiane Weirauch improved an early draft of the manuscript; Michael Schwartz and Gerasimos Cassis reviewed the submitted versions. To all of these individuals we offer our thanks for their contributions to improving the manuscript.

We thank Timothy Crowe, Chris Tobler, and Mike Picker of the University of Cape Town, Gerhard Prinsloo, Plant Protection Research Institute, Pretoria, and Lorenzo Prendini, American Museum of Natural History for logistical assistance in the conduct of field work in South Africa. We also thank the relevant authorities in the Western Cape and Northern Cape Provinces for the issuance of collecting permits. Gerasimos Cassis, Dimitri Forero, Thomas J. Henry, Michael D. Schwartz, Christiane Weirauch, and Denise Wyniger provided assistance in the field during 2003 and 2004.

The following institutions provided material for this study or material is deposited in them. We thank the persons indicated for their assistance. Institutional abbreviations used in the specimens examined sections are presented in the following list:

AM Australian Museum, Sydney, Australia; Gerasimos Cassis

AMNH American Museum of Natural History, New York

CNC Canadian National Insect Collection, Agriculture Canada, Ottawa; Robert G. Foottit

SANC Plant Protection Research Institute, Pretoria, South Africa; Ian Millar

SAMC South African Museum, Cape Town, South Africa; Simon van Nort

USNM United States National Museum of Natural History, Washington, D.C., Thomas J. Henry, Michele Touchet

ZISP Zoological Institute, Russian Academy of Sciences, St. Petersburg, I. M. Kerzhner

REFERENCES

Carvalho, J. C. M., J. A. P. Dutra and J. Becker. 1960. Hemiptera Heteroptera" Miridae. In: South African Animal Life. Stockholm: Swedish National Research Council, 7: 446-477.

Goloboff, P. A. 1993. Estimating character weights during tree search. Cladistics 9:83 91.

Goloboff, P. A. 1997. PeeWee, version 2.6. Parsimony and Implied Weights. Program and Documentation.

Goloboff, P. A. 1998. NONA, version 2.0. Program and Documentation.

Klak, C., A. Khunou, G. Reeves and T. Hedderson. 2003. A phylogenetic hypothesis for the Aizoaceae (Caryophyllales) based on four plastid DNA regions. American Journal of Botany 90: 1433-1445.

Nixon, K. C. 2000. WinClada. Version 0.9.

Poppius, B. 1912. Die Miriden der Athiopischen Region - Mirina, Cylapina, Bryocorina. Acta Societatis Scientiarum Fennicae 41(3): 203 pp., 1 pl.

Poppius, B. 1914. Die Miriden der Athiopischen Region II - Macrolophinae, Heterotominae, Phylinae. Acta Societatis Scientiarum Fennicae 44(3): 136 pp.

Schuh, R. T. 1974. The Orthotylinae and Phylinae (Hemiptera: Miridae) of South Africa with a phylogenetic analysis of the ant-mimetic tribes for the two subfamilies for the world. Entomologica Americana 47:332 pp.

Slater, J. A. 1964. Hemiptera (Heteroptera) Lygaeidae. In." South African Animal Life. Stockholm: Swedish National Research Council, 10:15-228.

Sta1, C. 1855. Hemiptera fran Kafferlandet. Ofversigt af Kongliga Vetenskaps3-akademiens forhandlingar, Stockhom 12: 27-46.

RANDALL T. SCHUH (1) AND GRACE Wu (2)

(1) Division of Invertebrate Zoology, American Museum of Natural History, New York, New York 10024, USA e-mail: schuh@amnh.org

(2) pomona College, Claremont, California, USA
KEY TO MALES OF EMINOCULUS

1. Eyes weakly stylate, posterior margin of eyes
   contiguous with anterior pronotal margin
   (Fig. 1); pronotal collar absent; antennal segment
   1 with numerous (> 15) erect black spines
   on medial surface (Fig. 5A, B); dorsum with
   long, decumbent, sericeous setae (Fig. 5A-C);
   breeds on Phyllobolus spp. (Mesembryanthemoideae)
                                             atrisetosus, new species

   -- Eyes stylate to grossly stylate, not contiguous
   with anterior pronotal margin; pronotal collar
   present; antennal segment 1 with one or a few
   erect black spines on medial surface; setae on
   dorsum variable length, coloration, and orientation              2

2. Smaller species (~2.5-2.7); apical vesical spine
   with a distinct subbasal indentation (notch) on
   dorsal surface (Fig. 6, character 43-2)                          3
   -- Larger species (>3.0); apical vesical spine with
   a very weakly developed subbasal indentation
   (notch) on dorsal surface (Fig. 8, character 43-1) 4
3. Dorsum appearing dull black; dorsum with
   dense silvery woolly vestiture; femora golden or
   light brown with variable dark maculae
   (Fig. 1); breeds on Delosperma spp. (Fig. 3C,
   D), Drosanthemum sp. (Ruschioideae)
                                             lanuginosus, new species
   --Dorsum polished, shining black; dorsum with
   evenly distributed short, black vestiture; femora
   black; breeds on Drosanthemum spp.
   (Fig. 2E) and Antimima cf. koekenaapensis
   (Ruschioideae) ... drosanthemi Schuh
4. Entire dorsum dull; posterior lobe of pronotum
   distinctly rugose; breeds on unidentified Mesembryanthemaceae
   (Fig. 3E, F)
                                                 rugosus, new species
   --Head and pronotum polished and shining,
   hemelytra dull; posterior lobe of pronotum at
   most weakly rugose                                               5
5. Posterior margin of vertex and eyes nearly
   straight; head and pronotum with inconspicuous,
   erect or semi-erect setae of moderate
   length; breeds on Ruschia muelleri and Leipoldtia
   cf. schultzei (Fig. 3A, B) (Ruschioideae)
                                              laevigatus, new species
   --Posterior margin of vertex and eyes curvilinear;
   head and pronotum with conspicuous, very
   long, erect, silvery setae                                       6
6. Lateral margin of hemelytra weakly concave;
   host unknown ... hirsutus Schuh
   --Lateral margin of pronotum weakly convex;
   breeds on Ruschia muelleri and R. decurrens
   (Fig. 2C, D) (Ruschioideae) . auratus, new species

KEY TO FEMALES OF EMINOCULUS

1. Eyes weakly stylate, posterior margin of eye
   partially contiguous with anterior pronotal
   margin; antennal segment 1 with numerous
   black spines on medial and lateral surfaces;
   width of head not exceeding width of pronoturn;
   dorsum with long, golden, decumbent,
   sericeous setae; breeds on el. Phyllobolus spp.
   (Fig. 2A, B) (Mesembryanthemoideae)
                                             atrisetosus, new species
   --Eyes stylate to grossly stylate, posterior margin of
   eye extending beyond anterior margin of pronorum;
   antennal segment 1 or a few black spines on
   medial surface; width of head exceeding width of
   pronotum; setae on dorsum variable in length,
   orientation, and coloration                                      2
2. Small, stout bodied, total length 2.1-2.7;
   abdomen ovoid and not distinctly acuminate
   apically (Fig. 1, drosanthemi, lanuginosus)                      3
   --Larger, body elongate, total length 3.0-4.5;
   abdomen distinctly acuminate apically; (Fig. 1,
   auratus, laevigatus, rugosus)                                    4
3. Dorsum with dense, long, silvery, woolly
   vestiture; femora pale or light brown with
   variable dark maculae; breeds on Delosperma
   spp. (Fig. 3C, D) and Drosanthemum sp.
   (Ruschioideae)                            lanuginosus, new species
   --Dorsum with evenly distributed, extremely
   short, black vestiture; femora black, except
   distally; breeds on Drosanthemum spp.
   (Fig. 2E) and Antimima of. koekenaapensis
   (Ruschioideae)                                   drosanthemi Schuh
4. Entire dorsum dull; posterior lobe of pronotum
   conspicuously rugose; breeds on unidentified
   Aizoaceae (Fig. 3E, F)                        rugosus, new species
   --Head and pronotum polished, shining black;
   scutellum and hemelytra moderately to highly
   polished; posterior lobe of pronotum at most
   weakly rugose                                                    5
5. Head and pronotum with inconspicuous,
   sparse, erect or semierect setae of moderate
   length; femora at least partially dark; hemelytra
   and exposed abdominal segments with
   inconspicuous, short, appressed, black or
   weakly golden setae; breeds on Ruschia muelleri
   and Leipoldtia cf. schultzei (Fig. 3A, B)
   (Ruschioideae)                             laevigatus, new species
   --Head and pronotum with numerous, conspicuous,
   long, erect, silvery setae; femora golden;
   hemelytra and exposed abdominal segments
   with numerous, conspicuous, decumbent, golden
   setae; breeds on Ruschia muelleri and R.
   decurrens (Fig. 3C, D) (Ruschioideae)
                                                 auratus, new species


Table 1.

                                                    Length

                      Total
Species               Leng    Cun-Clyp   LenHead   LenProno   LenScut

atrisetosus
M (N = 6)     Mean    3.19      2.33      0.26       0.52      0.31
              SD      0.06      0.05      0.05       0.03      0.02
              Range   0.18      0.13      0.12       0.08      0.06
              Min     3.07      2.27      0.22       0.49      0.29
              Max     3.25      2.40      0.34       0.57      0.35
F (N = 6)     Mean    3.32      2.96      0.33       0.57      0.33
              SD      0.15      0.10      0.05       0.04      0.04
              Range   0.42      0.25      0.12       0.10      0.10
              Min     3.12      2.81      0.27       0.51      0.30
              Max     3.54      3.06      0.39       0.61      0.39
auratus
M (N = 6)     Mean    3.98      2.72      0.27       0.80      0.32
              SD      0.13      0.10      0.06       0.03      0.06
              Range   0.35      0.27      0.16       0.08      0.14
              Min     3.80      2.55      0.23       0.76      0.25
              Max     4.15      2.81      0.39       0.84      0.39
F (N = 5)     Mean    3.96      3.01      0.27       0.85      0.30
              SD      0.33      0.15      0.03       0.06      0.04
              Range   0.79      0.38      0.08       0.14      0.11
              Min     3.52      2.80      0.21       0.79      0.23
              Max     4.31      3.18      0.29       0.93      0.35
drosanthemi
M (N = 7)     Mean    2.63      1.85      0.23       0.57      0.23
              SD      0.10      0.08      0.05       0.03      0.02
              Range   0.27      0.26      0.15       0.09      0.05
              Min     2.52      1.72      0.17       0.52      0.21
              Max     2.79      1.98      0.32       0.61      0.26
F (N = 7)     Mean    2.40      2.09      0.22       0.55      0.23
              SD      0.16      0.13      0.04       0.03      0.03
              Range   0.50      0.38      0.11       0.08      0.07
              Min     2.16      1.86      0.18       0.53      0.19
              Max     2.65      2.24      0.29       0.60      0.27
hirsutus
M (N = 5)     Mean    0.74      0.00      0.03       0.12      0.11
              SD      1.66      0.00      0.07       0.28      0.25
              Range   3.72      0.00      0.16       0.62      0.56
              Min     0.00      0.00      0.00       0.00      0.00
              Max     3.72      0.00      0.16       0.62      0.56
laevigatus
M (N = 5)     Mean    3.34      2.26      0.17       0.62      0.33
              SD      0.13      0.09      0.04       0.03      0.01
              Range   0.32      0.21      0.09       0.06      0.04
              Min     3.18      2.16      0.11       0.58      0.32
              Max     3.50      2.37      0.20       0.64      0.35
F (N = 6)     Mean    3.24      2.51      0.29       0.65      0.30
              SD      0.15      0.13      0.04       0.08      0.05
              Range   0.32      0.32      0.12       0.21      0.13
              Min     3.04      2.32      0.22       0.50      0.27
              Max     3.35      2.64      0.33       0.72      0.39
lanuginosus
M (N = 3)     Mean    2.60      1.82      0.14       0.52      0.29
              SD      0.02      0.03      0.03       0.02      0.02
              Range   0.05      0.05      0.05       0.04      0.05
              Min     2.58      1.79      0.12       0.50      0.27
              Max     2.63      1.84      0.17       0.54      0.32
F (N = 6)     Mean    2.48      2.19      0.19       0.57      0.22
              SD      0.12      0.10      0.02       0.06      0.02
              Range   0.33      0.25      0.06       0.14      0.07
              Min     2.29      2.04      0.17       0.49      0.17
              Max     2.62      2.29      0.23       0.64      0.25
rugosus
M (N = 5)     Mean    3.76      2.59      0.29       0.68      0.38
              SD      0.12      0.11      0.07       0.03      0.04
              Range   0.26      0.24      0.20       0.06      0.08
              Min     3.63      2.46      0.19       0.65      0.34
              Max     3.89      2.70      0.39       0.72      0.42
F (N = 6)     Mean    4.10      3.03      0.34       0.74      0.36
              SD      0.15      0.13      0.08       0.02      0.02
              Range   0.38      0.36      0.17       0.05      0.07
              Min     3.96      2.81      0.28       0.70      0.33
              Max     4.34      3.17      0.45       0.76      0.39

                      Length                         Width

Species               LenCun   WidHead   WidProno   WidScut

atrisetosus
M (N = 6)     Mean     0.51     1.17       1.12      0.44
              SD       0.04     0.02       0.02      0.02
              Range    0.12     0.05       0.05      0.05
              Min      0.46     1.13       1.10      0.41
              Max      0.58     1.19       1.15      0.46
F (N = 6)     Mean     0.00     1.33       1.21      0.46
              SD       0.00     0.02       0.05      0.07
              Range    0.00     0.05       0.12      0.17
              Min      0.00     1.31       1.15      0.37
              Max      0.00     1.36       1.27      0.54
auratus
M (N = 6)     Mean     0.65     1.42       1.30      0.38
              SD       0.06     0.04       0.07      0.08
              Range    0.17     0.12       0.19      0.19
              Min      0.54     1.34       1.22      0.27
              Max      0.72     1.46       1.41      0.47
F (N = 5)     Mean     0.00     1.59       1.22      0.35
              SD       0.00     0.02       0.03      0.05
              Range    0.00     0.04       0.08      0.12
              Min      0.00     1.57       1.17      0.28
              Max      0.00     1.61       1.25      0.40
drosanthemi
M (N = 7)     Mean     0.32     1.08       0.98      0.31
              SD       0.02     0.03       0.05      0.03
              Range    0.06     0.10       0.13      0.09
              Min      0.29     1.02       0.93      0.27
              Max      0.35     1.12       1.06      0.37
F (N = 7)     Mean     0.00     1.23       1.00      0.31
              SD       0.00     0.05       0.05      0.05
              Range    0.00     0.14       0.12      0.15
              Min      0.00     1.15       0.94      0.23
              Max      0.00     1.29       1.06      0.38
hirsutus
M (N = 5)     Mean     0.14     0.26       0.24      0.14
              SD       0.30     0.59       0.54      0.30
              Range    0.68     1.32       1.20      0.68
              Min      0.00     0.00       0.00      0.00
              Max      0.68     1.32       1.20      0.68
laevigatus
M (N = 5)     Mean     0.50     1.25       1.12      0.46
              SD       0.04     0.03       0.06      0.02
              Range    0.09     0.07       0.14      0.06
              Min      0.46     1.21       1.02      0.43
              Max      0.54     1.28       1.17      0.49
F (N = 6)     Mean     0.00     1.37       1.06      0.32
              SD       0.00     0.07       0.04      0.05
              Range    0.00     0.18       0.11      0.14
              Min      0.00     1.27       1.00      0.23
              Max      0.00     1.45       1.11      0.37
lanuginosus
M (N = 3)     Mean     0.33     1.11       0.99      0.39
              SD       0.06     0.03       0.02      0.04
              Range    0.10     0.05       0.05      0.09
              Min      0.26     1.07       0.97      0.34
              Max      0.36     1.13       1.02      0.43
F (N = 6)     Mean     0.00     1.28       1.05      0.39
              SD       0.00     0.05       0.04      0.05
              Range    0.00     0.11       0.10      0.13
              Min      0.00     1.23       0.99      0.34
              Max      0.00     1.34       1.09      0.47
rugosus
M (N = 5)     Mean     0.58     1.38       1.28      0.48
              SD       0.03     0.05       0.02      0.07
              Range    0.08     0.13       0.06      0.15
              Min      0.56     1.32       1.25      0.41
              Max      0.63     1.45       1.31      0.56
F (N = 6)     Mean     0.00     1.64       1.34      0.43
              SD       0.00     0.04       0.05      0.04
              Range    0.00     0.10       0.11      0.10
              Min      0.00     1.60       1.27      0.39
              Max      0.00     1.70       1.38      0.49

                            Width

Species               InterOcDi   AntSeg2

atrisetosus
M (N = 6)     Mean      0.61       1.24
              SD        0.05       0.06
              Range     0.14       0.14
              Min       0.55       1.15
              Max       0.69       1.30
F (N = 6)     Mean      0.75       1.25
              SD        0.05       0.05
              Range     0.11       0.13
              Min       0.68       1.18
              Max       0.80       1.31
auratus
M (N = 6)     Mean      0.80       1.36
              SD        0.05       0.05
              Range     0.13       0.13
              Min       0.71       1.29
              Max       0.84       1.43
F (N = 5)     Mean      0.99       1.08
              SD        0.04       0.10
              Range     0.10       0.26
              Min       0.95       0.92
              Max       1.05       1.19
drosanthemi
M (N = 7)     Mean      0.59       0.68
              SD        0.02       0.06
              Range     0.04       0.17
              Min       0.56       0.58
              Max       0.60       0.75
F (N = 7)     Mean      0.73       0.69
              SD        0.03       0.05
              Range     0.08       0.17
              Min       0.69       0.62
              Max       0.77       0.78
hirsutus
M (N = 5)     Mean      0.15       0.21
              SD        0.33       0.47
              Range     0.74       1.06
              Min       0.00       0.00
              Max       0.74       1.06
laevigatus
M (N = 5)     Mean      0.72       1.07
              SD        0.03       0.05
              Range     0.08       0.12
              Min       0.68       0.99
              Max       0.75       1.11
F (N = 6)     Mean      0.86       0.87
              SD        0.06       0.08
              Range     0.17       0.21
              Min       0.77       0.74
              Max       0.94       0.94
lanuginosus
M (N = 3)     Mean      0.71       0.67
              SD        0.02       0.02
              Range     0.04       0.04
              Min       0.69       0.66
              Max       0.73       0.70
F (N = 6)     Mean      0.87       0.65
              SD        0.05       0.04
              Range     0.13       0.10
              Min       0.79       0.61
              Max       0.93       0.71
rugosus
M (N = 5)     Mean      0.78       1.42
              SD        0.02       0.07
              Range     0.05       0.17
              Min       0.75       1.32
              Max       0.80       1.49
F (N = 6)     Mean      1.01       1.29
              SD        0.03       0.06
              Range     0.07       0.16
              Min       0.99       1.21
              Max       1.05       1.38

Table 2. Character descriptions for Eminoculus spp.

Antennal coloration

0 males color of antennal segment 1: 0) pale, 1) infuscate, 2)
black;

1 males color of antennal segment 2: 0) pale, 1) infuscate, 2)
black;

2 females color of antennal segment 1: 0) pale/golden (excepting
base), 1) infuscate black;

3 females color of antennal segment 2: 0) pale/golden, 1) banded,
pale medially, 2) black;

Leg coloration

4 males color proepisternal supracoxal lobe: 0) dark brown or
black, 1) yellow;

5 males color femora: 0) black, 1) black with pale apex, 2) golden;

6 males color femoral maculae: 0) absent, 1) present;

7 males color tibiae: 0) pale, 1) golden, 2) moderately infuscate,
3) black;

8 males color tarsi: 0) pale, 1) banded (1 and 3 dark, 2 pale), 2)
banded (1 and 2 pale, 3 dark), 3) black;

9 females color femora: 0) golden, 1) black, 2) infuscate;

10 females color femoral maculae: 0) absent, 1) present;

11 females color tibiae: 0) golden/pale, 1) golden with tibial
spines with black bases, 2) black, 3) weakly infuscate;

12 females color tarsi: 0) pale, 1) banded (1 and 3 dark, 2 pale),
2) banded (1 and 2 pale, 3 dark), 3) infuscate black;

Surface texture

13 males surface texture head and pronotum: 0) dull black, not
polished, 1) highly polished, shining black;

14 males surface and texture calli: 0) smooth and dull, 1) smooth
and highly polished;

15 males surface posterior lobe of pronotum: 0) smooth and dull, 1)
weakly rugulose and dull, 2) rugose and polished and shining;

16 males surface texture hemelytra: 0) dull black, not polished, 1)
polished, shining black; Vestiture in male

17 antennal segment 1 spines: 0) few slender erect setae, 1) many
erect spines;

18 vestiture density head and pronotum: 0) sparse, 1) dense;

19 surface and vestiture length head and pronotum: 0) short, 1)
long, 2) very long;

20 vestiture type head and pronotum: 0) simple, 1) sericeous, 2)
woolly, 3) scale-like;

21 vestiture orientation head and pronotum:, 1) erect, 2)
semierect, 3) decumbent;

22 vestiture color head and pronotum: 0) silver, 1) golden, 2)
dark;

23 vestiture density hemelytra: 0) sparse (few), 1) dense;

24 vestiture length hemelytra: 0) very short, 1) short, 2) long;

25 vestiture texture hemelytra: 0) sericeous, 1) woolly, 2) simple,
3) flattened, scalelike;

26 vestiture orientation hemelytra: 0) decumbent (appressed), 1)
reclining;

27 vestiture color hemelytra: 0) silver, 1) golden, 2) black;

Structure

28 structure of antennal segment 1: O) moderately swollen (convex
on medial margin) and short, 1) weakly swollen and parallel-sided
and short, 2) parallel-sided and elongate;

29 males structure eyes: O) sub stylate, 1) weakly stylate, 2)
stylate, 3) grossly stylate;

30 males structure posterior margin of eyes: O) contiguous with
anterior pronotal margin, 1) not contiguous with anterior pronotal
margin;

31 males orientation of posterior margin of eyes: O) curvilinear
posteriorly, 1) nearly straight posteriorly;

32 males structure labium: O) extending well beyond trochanteral
joint of procoxae, 1) not extending beyond trochanteral joint of
procoxae;

33 males structure pronotum shape: O) nearly fiat, 1) swollen and
inclined longitudinally and transversely;

34 pronotal collar: O) absent, anterior margin of pronotum narrowly
reflexed, 1) present, broad and fiat;

35 males structure thorax calli: O) weakly elevated, 1) slightly
elevated, 2) distinctly elevated;

36 males calli contiguous across anterior margin: O) no, 1) yes;

37 females structure posterior pronotal margin: O) straight across
scutellum, 1) concave across scutellum;

38 males structure costal hemelytra margin: O) convex, 1) linear
over most of length, :2) concave;

39 females hemelytral condition: O) macropterous, 1) brachypterous,
membrane rudimentary;

40 supracoxal lobes of prothorax: O) lying flat against coxa O)
projecting laterally;

41 claw hairs/pulvillar combs: O) absent, 1) few (1--3), 2) many
(4-6);

42 females structure abdomen: O) ovoid and not distinctly acuminate
apically, 1) nearly parallel-sided and distinctly acuminate
apically;

Male genitalia

43 subbasal notch on dorsal surface of apical spine of vesica: 0)
absent, 1) weak, 2) strong;

44 flange on dorsal surface of vesica: 0) flowing smoothly to base
of secondary gonopore, 1) truncate at level of base of secondary
gonopore;

45 width of vesica body: 0) relatively slender, 1) broad;

46 field of spicules subtending secondary gonopore: 0) absent, 1)
present;

47 right paramere apical extension: 0) distinctly pointed
fingerlike and central, 1) fingerlike, distinctly left of center,
2) blunt;

48 left paramere development of anterior process: 0) abbreviated
(barely extending beyond shaft of left paramere), 1) moderately
long (extending beyond shaft of left paramere), 2) elongate
(extending well beyond shaft of left paramere);

49 left paramere posterior process: 0) elongate and decurved
elongate and straight, 1) broad and moderately elongate and
straight, 2) greatly elongate, elevated, and lanceolate;

50 left paramere setae on anterior process: 0) absent or scattered,
1) long, heavy setae clumped at apex of process.

Table 3. Character distributions in Eminoculus spp.

                                        Character number

                 000000000011111111112222222222333333333344444444445
    Taxon        012345678901234567890123456789012345678901234567890

Chlamydatus
  keltoni        2-2-01002100200000000220220200000000000000000000110
Lasiolabopella
  capeneri       00000000010000000000322123-00000000000000000000-000
atrisetosus      222210033102400101011111201100001000000101021012110
auratus          220111021000111200121001201123111112112112110111111
drosanthemi      000111021103111210000110000211000010010111021013030
hirsutus         22---0012---- 112001210011010231111121-1-1---------
laevigatus       220111023103311210010100100212111112011112110012121
lanuginosus      110012112011211200112101210011111010010111021011010
rugosus          221210031203101200111111200112111111111112110012111
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