Revision of the genus car valhomiris (hemiptera: miridae: orthotylinae).
Maldonado and Ferreira (1971) proposed Carvalhomiris to accommodate their new species C. brachypterus, described from specimens collected from areas nearby Bogotfi, Colombia.
During study of additional material collected by T.J. Henry in Ecuador and by R.T. Schuh in Colombia, two new species of Carvalhomiris were discovered. These additional species permit comparisons of the male and female genitalia, and support a revised diagnosis for Carvalhomiris that encompasses observed variation. Furthermore, these new species provide additional information that helps postulate putative synapomorphies for the species of the genus, and for understanding the relationships of Carvalhomiris with other Orthotylinae.
MATERIAL AND METHODS
Observations were made with a Nikon SMZ1500 stereoscope. Drawings of male and female genitalic structures were made using a Nikon Eclipse 80i compound microscope, with a camera lucida attachment. Scanning Electron Micrographs (SEM) were taken using a Hitachi S4700 Field Emission SEM. Digital dorsal color images were made using a Microptics-USA photomicrographic system, with Infinity K2 lens.
As part of the Plant-Bug Planetary Biodiversity Inventory project (PBI), all specimens studied are associated with a Unique Specimen Identifier (USI). These USIs are machine-readable matrix labels. Each USI has the prefix of the project (AMNH_PBI) followed by its unique number, for example AMNH-PBI 00123456. The USI allows for unambiguous identification of each specimen from the Plant-Bug PBI. Additional information can also be retrieved with each USI via the Discover Life website (http://www.discoverlife. org).
Measurements of specimens (Table 1) were made with an industrial micrometer mounted on a Wild M7 stereoscope with Mitutoyo displays (assembled by Stangert Corporation, Island Park, NY). USI numbers of those specimens measured are listed in "Appendix 1." Because species of Carvalhomiris are brachypterous and macropterous, an explanation is needed to clarify how the length measurements were taken. Total length was measured from the apex of the clypeus to the caudal margin of the membrane in macropterous specimens, and up to the caudal margin of the abdomen in the case of brachypterous individuals. The cuneo-clypeal length was measured from the apex of the clypeus to the caudal margin of the cuneus in the case of macropterous specimens, and from the apex of the clypeus to the caudal margin of the hemelytron in the case of brachypterous ones.
To study ovarian eggs, the abdomen of a mature female, for each one of the species, was removed and placed in warm distilled water. It was then dissected to expose the ovaries. Further dissection was done to expose the eggs, which were then placed in cold 10% KOH solution for a few minutes. After removing the surrounding tissue, the eggs were rinsed in distilled water, then in ethanol, and transferred to glycerin for further dissection and study. SEMs of eggs were made with ethanol rinsed, air-dried specimens.
General external morphological terms, in particular about the metathoracic scent gland efferent area, follow Schuh and Slater (1995). Terminology for male genitalia follows Kelton (1959), Konstantinov (2003), and Forero (2008). Terminology for female genitalia follows Davis (1955), Scudder (1959), and Forero (2008). For egg morphology I followed the terminology of Cobben (1968), in particular the use of the term "aft" which is the side of the egg mostly associated with the substrate (Cobben, 1968: 10).
All localities were georeferenced using local South American maps and gazetteers, and with the aid of GEOLocate (Rios and Bart, 2005). The distribution map for the species was generated with ArcMap 9.1 (ESRI, 2005), using a shaded relief base map derived from a digital elevation model of the Shuttle Radar Topography Mission (SRTM) from NASA.
In the following sections, species descriptions do not repeat invariable characters already mentioned in the generic description. I also describe in detail the brachypterous specimens first, and then the macropterous. Despite being an unusual practice in Miridae taxonomy, where most of the specimens are macropterous, in the case of Carvalhomiris I decided to describe the brachypterous specimens first because those are the most common specimens found, where macropterous forms are unlikely to be found (see the "material examined" section for numbers of macropterous versus brachypterous specimens).
In the "material examined" section, all interpolated data, in particular georeferences, are between square brackets. Unless otherwise stated, all measurements are in millimeters. Acronyms, names of institutions, and curators or individuals responsible of the collections mentioned in the text are as follow: AMNH, American Museum of Natural History, New York, Randall T. Schuh; CNC, Canadian National Collection of Insects, Agriculture Canada, Ottawa, Michael D. Schwartz; ICN, Instituto de Ciencias Naturales, Universidad Nacional de Colombia, Bogotfi, Carlos Sarmiento; JTP, John T. Polhemus Collection, Englewood, Colorado; MNRJ, Museu Nacional, Universidade Federal do Rio de Janeiro, Brazil, Luiz A. A. Costa; TAMU, Texas A&M University, College Station, Joseph C. Schaffner and Edward C. Riley; UNAB, Museo Entomol6gico, Facultad de Agronomia, Universidad Nacional de Colombia, Bogota, Francisco Serna and Erika V. Vergara; USNM, United States National Museum of Natural History, Washington D.C., Thomas J. Henry and Michele Touchet.
CARVALHOMIRIS MALDONADO AND FERREIRA Figures 1-11
Carvalhomiris Maldonado and Ferreira, 1971 [n. gen.]; Schuh, 1975: 17, figs. 52, 53, 61, 62 [femoral trichobothria]; Schuh, 1976: fig. 16 [pretarsus]; Schuh, 1995:89 [catalog].
TYPE SPECIES: Carvalhomiris brachypterus Maldonado and Ferreira, 1971 (by original designation).
DIAGNOSIS: Recognized by the pale green coloration; the usual strong brachyptery in both sexes (Fig. 1), with macroptery rare or uncommon; head with eyes removed from anterior margin of pronotum (Fig. 2); evaporatory area and peritreme on metathorax protruding (Figs. 3A-D); large, ovoid genital capsule (Figs. 3, 7); vesica with a single spiculum composed of a dorsal and a ventral portion (Fig. 5); right paramere clubbed in lateral view, ventral prolongation apically with two prongs (Fig. 6); lobe of interramal sclerite greatly elongated and paddle-shaped, located at the middle of sclerite (Fig. 8); and by the egg with a basal process (Fig. 10).
Carvalhomiris is easily recognized among other Neotropical Orthotylini genera by having the eyes removed from the anterior margin of the pronotum and by the usual strong brachypterism. Some other Neotropical genera also have species with the eyes removed from the anterior margin of the pronotum (e.g., Itacoris Carvalho, Parachius Distant, Paraproba Distant), but none of these genera present brachypterous species, nor are they as robust as Carvalhomiris. Furthermore, the particular configuration of the male genitalia, as described below, easily allows for the recognition of this genus.
[FIGURE 1 OMITTED]
REDESCRIPTION: Brachypterous male: Medium sized, oval shaped, robust, total length 2.48-3.53. COLORATION (Fig. 1): Overall coloration pale yellow with greenish areas. Head: Sometimes dark with postocular dark brown or reddish markings; first antennal segment usually dark brown, remaining segments with variable color pattern; apex of labial segment IV dark brown. Thorax: Pronotum pale green or with various degrees of dark brown markings; scutellum pale green, yellowish, or dark brown; sternum pale yellow. Hemelytron: Bright green. Legs: Third tarsomere dark brown. Abdomen: Pale yellow. SURFACE AND VESTITURE: Surface dull, covered with dense microtrichia. Head and pronotum covered with semi-decumbent or erect, simple setae (Fig. 2); antennal segment II on medial surface of basal third with a fringe of short, suberect, simple setae set in a dense patch, or with normal, decumbent, very short setae as on remaining surface of antenna; hemelytron with dark or yellowish, decumbent, simple setae (Figs. 3E, F), sometimes with a dense patch of these setae apically; femora with dense simple setae on dorsal surface, ventral surface with sparse, simple setae; fore femur on ventral surface with sparse, erect, long, simple setae; middle and hind femur on ventral surface with 8-9 trichobothria each, most of them with dense trichoma at their base (see also Schuh, 1975: Figs. 52, 53, 61, 62); tibiae with dense, decumbent, simple setae, and stout, sparse, semi-erect setae; abdominal sternites with sparse, semi-decumbent, simple setae, more densely set on ventral surface. STRUCTURE: Head (Fig. 2): Short, transverse. Clypeus protruded, basally strongly convex; frons strongly convex, slightly protruded over base of clypeus (e.g., Fig. 2B); vertex slightly convex; ocular carina slightly elevated, impressed on anterior margin (Figs. 2E, F); mandibular and maxillary plates taken together about half the height of the head; mandibular plate triangular, apically acute; maxillary plate elongate, subquadrangular, rounded at apex (Figs. 2A, B); buccula shorter than gula; gena elongate; gula about the same length as postocular region; eyes removed from anterior margin of pronotum, located approximately on longitudinal midpoint of head (Figs. 1, 2A, B), rounded in dorsal view, nearly reniform in lateral view, not surpassing dorsal margin of head in lateral view, and occupying upper half of head in lateral view, posterior area of eye glabrous (Figs. 2B, D), postocular region swollen ventrally with posterior margin continuing dorsally to meet ocular carina (Figs. 2A-D, 2C arrow); labrum triangular, as long as buccula (Figs. 2C, D); labium reaching metacoxa, all segments of approximately equal length; antennae long (Fig. 1), antennal segment I stout, longer than head length (table 1), II-IV of lesser diameter than I, II about 2.4 times as long as I (table 1), lengths of III and IV combined about the same length as II. Thorax (Figs. 2E-H; 3A-D): Collar fiat, narrow, anterior margin in dorsal view gently concave; pronotum trapezoidal, lateral margins nearly straight or sinuate, anterior lobe about as long as head length, surface flat or nearly convex, posterior lobe about half as long as anterior lobe, surface nearly fiat, posterior margin medially emarginate; calli on pronotum indistinguishable from surrounding pronotal surface, neither elevated nor rugose; mesoscutum greatly exposed, nearly fiat; scutellum triangular, anterior margin sinuate, not clearly separated from mesoscutum, apex acute, disc flat; proepisternum laterally protruding and visible in dorsal view; proepimeron slightly concave before ventral margin; propleural sulcus strongly arcuate anteriorly, with large medial depression (Figs. 2G, H); mesepisternum nearly fiat; mesepimeron concave on ventroposterior margin; mesopleural sulcus faintly impressed, dorsal portion ending in large pitlike concavity; opening of mesothoracic spiracle with wide mushroomlike cuticle, elevated with respect to surrounding surface (Figs. 3A, B); metepisternum with scent gland efferent system large, in contact with ventral area of mesothoracic spiracle, dorsal margin of mushroomlike cuticle of evaporatory area gently curved, not greatly inclined, evaporatory area on anterior half greatly protruded, curved upward, anterior margin of opening of channel strongly sinuate, peritreme relatively small, ovoid, slightly notched medially, situated on top of recurved portion of evaporatory area and not raised above it (Figs. 3A-D); prosternum triangular, lateral margins elevated. Hemelytron (Fig. 1): Oblong, short, apex not surpassing apex of genital capsule, the latter exposed dorsally; humeral plate mostly covered by posterolateral angles of pronotum, barely visible in dorsal view; costal margin gently curved; embolium and claval suture faintly impressed; membrane and cuneus absent; apex round or acute. Legs: Femora cylindrical, elongate; fore and middle femur about the same length, hind femur slightly longer and greater in diameter than fore and middle femur, and slightly curved posteriorly in dorsal view (Fig. 1); tibiae cylindrical, fore and middle tibia about the same length and diameter, hind tibia about 1.5 times as long as middle tibia, and wider in diameter; tarsi elongated, fore and middle tarsi about one third as long as tibial length, hind tarsi about 0.26 times as long as tibial length; pretarsus as in Figure 4, base of claw with 3 hairs directed ventrolaterad (Figs. 4A-C, 4A arrows), and numbered here I to III from apical to basal (Fig. 4D); hair I medium-sized, directed laterad, II the longest, directed ventrad, and partially sulcate (Figs. 4B, C), III the shortest, directed ventrad, bases of II and III located close together. Abdomen: Ovoid, large, about half the length of total body length. Genitalia: Genital capsule relatively large, ovoid, curved in lateral view (Figs. 3G, H); aperture semicircular, margin well-sclerotized, entire, without tergal or dorsal processes (Fig. 7); proctiger well-sclerotized on dorsal surface, reaching apex of genital capsule (Fig. 7); cuplike sclerite with medial area fiat, not raised from surface of genital capsule, its caudal margin not surpassing apex of genital capsule, raised left portion larger than right portion; insertion of parameres located on a horizontal plane; right paramere (Fig. 6) expanded on apical half and flattened laterally, body elongate, ventral portion of paramere elongate, its apex with two prongs, sometimes strongly bifurcate, apical margin finely serrate, either nearly straight or with a medial truncate process, dorsal portion of paramere acute; left paramere (Fig. 6) sickle-shaped in dorsal view, area caudal to dorsal sensory area either flat or with a serrate tubercle, apex of paramere with a large and acute or small and blunt ventral process, ventral preapical area either with a large protruding area covered with medium-sized spicules or with a nearly flat area covered with very small spicules; phallotheca (Fig. 5) nearly cylindrical, apex slightly curved to the right, opening dorsal, cephalic margin of opening slightly sclerotized, right margin of opening with a medial lobelike prolongation, preapical area of right margin elevated, with an acute process or rounded, near the base with lateral paired flaplike processes, heavily sclerotized vertical rodlike area (e.g., Fig. 5: C. truncatus lateral right view); vesica with a single C-shaped spicule (Fig. 5) composed of a dorsal and ventral portions, base of spiculum weakly sclerotized; dorsal portion of spicule shorter than ventral, from about one half to three-fourths the length of ventral portion, apex sclerotized, either straight or curved to the left in dorsal view, dorsal area weakly sclerotized; ventral portion of spicule with apex reaching or not apex of phallotheca, apex of ventral portion acute or expanded; sclerotized part of ductus seminis short, forming a single sclerotized unit, about half the length of dorsal portion of spicule and located close to its apex; when everted, spicule and sclerotized part of ductus seminis maintain relative position as in non-everted aedeagus (Fig. 5); opening of secondary gonopore directed ventrad.
[FIGURE 2 OMITTED]
Macropterous male: Similar to brachypterous male; total length 4.21-4.66. COLORATION: Thorax: Hemelytron fuscous. STRUCTURE: Thorax (Fig. 1): Posterior lobe of pronotum about as long as anterior lobe, inclined and elevated with respect to anterior lobe, transverse sulcus dividing the two lobes deeply impressed; mesoscutum greatly enlarged, disc convex, posterolateral sides flat and vertical; scutellum with disc slightly convex, apex rounded, anterior margin clearly delimited from mesoscutum; proepimeron strongly concave on ventral area. Hemelytron (Fig. 1): Elongate, completely covering the abdomen; lateral margin nearly straight, slightly wider at level of apex of embolium, clavus elevated with respect to corium surface; claval suture impressed; embolium well delimited, apex ending close to cuneal fracture, not greatly expanded inward; cuneus slightly longer than wide; membrane with two cells.
Brachypterous female: Similar in structure and coloration to brachypterous male but larger, more robust, and abdomen more ovoid; total length 3.11-4.10. VESTITURE: Head: Antennal segment II with normal, decumbent, very short setae as in remaining surface of antennae. STRUCTURE: Thorax: Hemelytron (Fig. 1): Oblong or ovoid elongate, apex reaching abdominal tergite VI or nearly reaching caudal margin of abdomen. Genitalia (Fig. 8): Subgenitai plate subtriangular, apex rounded; base of ovipositor located caudad to longitudinal midpoint of abdomen; posterior wall well sclerotized; interramal sclerite not differentiated from surrounding membranes, apparently occupying entire interramal space, produced anteriorly as a round, blunt lobe; medial margin of interramal sclerite on caudal half raised and corrugated forming a medial dorsal protuberance, dorsal margin smooth, without spines; lobe of interramal sclerite very long, about one half of its total length projected beyond anterior margin of posterior wall, paddle-shaped, apical half twisted about 45 degrees with respect to basal half, lobe located on vertical midpoint of interramal sclerite, covered with strong microtrichia; medial sclerite flat, well sclerotized, without ventral processes; dorsal labiate plate with a longitudinal median depression ventral to common oviduct, the latter weakly sclerotized on lateral margins, where lateral oviducts are inserted; accessory gland inserted in deep, pitlike depression, area of insertion not sclerotized, but surrounding area well sclerotized, produced ventrally as a conical protuberance; sclerotized ring oblong, wider on caudal portion than on cephalic one, cephalic portion and surrounding membranes with a field of microtrichia, medial margin curved inward, posteromedial angle with a small process directed caudad, lateral infolding of ring sclerotized on caudal half, with a lateral, long, sclerotized, and strongly acute process directed caudad on lateral margin; ventral labiate plate membranous; anterior medial margin of first gonapophysis enlarged and sclerotized (Fig. 9), either nearly symmetrical or highly asymmetrical, cephalic portion of margin strongly emarginate with respect to ventral labiate plate, medial wrinkled surface of gonapophysis with anterior portion lobelike, detached from main body of gonapophysis. Ovarian Egg: Elongate, ovoid (Fig. 10A); aft side (see Cobben, 1968: 10) basally with a prominent knoblike tubercle (Figs. 10A arrow, C-F), apex of tubercle with lateral prolongations (Figs. 10C, E arrow), posterior margin of tubercle rounded (Fig. 10F); chorion deeply punctate with elevated ridges forming nearly pentagonal shapes (Fig. 10B), surrounding area of insertion of knoblike process smooth, devoid of punctures (Figs. 10D, F); rim of corium with approximately 40-50 aeropyles, micropyles apically hook-shaped and not greatly projecting beyond margin of rim; operculum ovoid, with a large respiratory horn close to the fore side of egg (Figs. 10G, H arrows), expanded and tuberculate apically.
[FIGURE 3 OMITTED]
Macropterous female: Similar in structure and coloration to brachypterous female; total length 4.43-4.61. STRUCTURE: Thorax and hemelytron modified as that of macropterous male.
DISTRIBUTION: Carvalhomiris is known from middle to relatively high elevations in the Andes of Colombia and Ecuador (Fig. 11). The extreme gap between the localities of the known species in Ecuador and Colombia is probably a reflection of inadequate sampling of suitable habitats.
HOST ASSOCIATIONS: Two out of the three species of Carvalhomiris lack host plant data. The only recorded host plant is an unidentified species of Tropaeolaceae. Most of the species of Tropaeolaceae are found in the genus Tropaeolum, which has primarily an Andean distribution (Chavez, 2004). Future fieldwork in Andean montane forests should focus on accurate host-plant data for this genus.
DISCUSSION: MORPHOLOGY: The swollen postocular region as seen in Carvalhomiris has not been documented for any known Miridae taxa. Many Orthotylini genera worldwide have the eyes removed from the anterior margin of the pronotum to various degrees, but a conspicuous swelling has not been noted before. This swelling (Figs. 2A-D, 2C arrow) is apparently not part of the eye, but is located in the postocular region of the head. The most caudal part of the eye in Carvalhomiris is a smooth area (e.g., Fig. 2D) that occupies most of the concave portion of the eye as seen in lateral view (e.g., Fig. 2B). In Miridae, as in many other Heteroptera, the postocular region of the head does not have any sutures that allow delimitation of the head capsule sclerites (Spooner, 1938). Nonetheless, in other Orthotylini the caudal part of the eye is clearly delimited and differentiated from the postocular region, for instance in Orthotylus flavosparsus (Sahlberg) (Southwood, 1953: Figs. 1, 3, 7). In some orthotylines in which the eye is removed from the anterior margin of the pronotum, the postocular region is coplanar with the gular area, for instance in Hadronemidea echinata (Gruetzmacher and Schaffner) (Forero, 2008: Fig. 38A). In many other taxa, such as Itacorisi and Parachius, or some of the species of Zanchius (pers. obs.), a clearly delimited swollen postocular region is present. The presence of this structure needs to be surveyed in a broader sample of genera.
Schuh (1976) described and documented at length the pretarsal structure of Miridae. He presented several SEMs of various taxa, including some Phylinae and Orthotylinae (sensu Schuh, 1995). While discussing the base of the claw, he mentioned that most Phylinae, Orthotylinae, and Mirinae have a field of spicules that he called "claw spicules" (Schuh, 1976: 8, Figs. 15, 18, 21, 26, 42). Schuh also termed "claw hairs" those hairlike structures found on the outer surface of the claw (Schuh, 1976: 18; see also Wagner, 1952). He stated that no Orthotylini (as "Orthotylina") have these hairs, although he documented their presence in the Halticini and Phylinae. In Carvalhomiris, I found that the base of each claw does not have a field of small spicules, but instead there are three hairs of different length (Fig. 4A arrows). These hairs cannot be confused with "claw hairs," because they are located at the base of the claw and not on its more dorsolateral surface as documented by Schuh (1976). The only other Orthotylini, as far as I am aware, in which similar basal claw hairs have been documented are Zanchius breviceps (Wagner, 1951) (Doesburg, 1984) and Saileria bella (Van Duzee, 1916) (Forero, 2009). Doesburg (1984) presented some accurate illustrations of the external morphology of Z. breviceps (as Z. stami). He provided a lateral view of the pretarsus (Doesburg, 1984: Fig. 4), in which three hairs--and not a field of spicules--are clearly visible at the base of the claw. Doesburg, nonetheless, did not discuss the presence or describe the structure of these hairs. Forero (2009: Fig. 19E) presented a SEM of the pretarsus of S. bella in ventral view. Similarly to Doesburg, he did not recognize the novel structure of the pretarsus, and failed to further discuss it. Pretarsal mirid morphology has usually been documented using scanning electron micrographs in dorsal or anterior views (e.g., Schuh, 1976; Schwartz and Stonedahl, 1987; Schwartz, 2004; but see e.g., Stonedahl and Schwartz, 1986: Figs. 36-56; Weirauch, 2006: Fig. 8; Forero, 2008: Figs. 20E, 45C). In those views it is very difficult to examine the base of the claw (e.g., Figs. 4E, F). A thorough survey for this modification of the base of the claw in other Orthotylini is necessary, with the claw being viewed in the proper orientation.
[FIGURE 4 OMITTED]
The phallotheca of Carvalhomiris has a pair of lateral flaplike processes near its base and a vertical sclerotized rodlike area on the surface cephalad to them (e.g., Fig. 5, C. truncatus). The paired flaplike modification of the phallotheca is unique among Orthotylini. The phallotheca in Orthotylini is commonly nearly cylindrical, and usually its basal, lateral area is not modified. In some instances, a small protuberance may be found, for instance on the basal left side of Aoplonema Knight (Forero, 2008: Fig. 6). In this taxon the protuberance is neither flaplike nor paired. Because usually little attention has been paid to the detailed structure of the phallotheca in the description of Orthotylinae taxa, the occurrence of these flaplike structures in other Orthotylinae is not known. The same is true for the rodlike modification of the cuticle cephalad to the flaplike processes.
The female genitalia of Carvalhomiris have some novel morphological modifications (Fig. 8). The dorsal labiate plate is medially depressed to receive the common oviduct. Usually, the medial area of the dorsal labiate plate in Orthotylini is flat, and mostly membranous or with slightly sclerotized, paired areas (pers. obs.). The accessory gland in Carvalhomiris is inserted into a cuplike depression, in which the area of insertion is membranous and the surrounding area is well sclerotized. The depressed area forms a cuplike protuberance on the dorsal labiate plate that projects ventrad into the vestibulum. In other Orthotylini, the gland inserts into a flat, nondepressed area. The lateral infoldings over the sclerotized rings in Carvalhomiris have an acute caudad process. In Orthotylini, these lateral folds are strongly produced (Schuh, 1974: 267), and are variously sclerotized. These folds are connected laterally to the inner cuticle of the female abdomen by the muscle number 6 (i.e., M6), which originates in a tendon from the lateral infolding, and helps during oviposition (Davis, 1955: 148, Fig. 18). These M6 tendons are usually not strongly sclerotized (e.g., Slater, 1950; Davis, 1955), but in Carvalhomiris the lateral margin of the lateral infolding from which the tendon is produced, is greatly enlarged and sclerotized. The interramal lobe (="K" structure of Slater, 1950, and "processes" of Davis, 1955) in most Orthotylini is located on the dorsal-most area of the interramal sclerite (e.g., Schwartz, 2004; Schaffner and Schwartz, 2008; Forero, 2008). In Carvalhomiris, this lobe is located more medially than in other known species of Orthotylini. In addition, this lobe is elongate and strongly paddle-shaped, which is unique among the known species of Orthotylini (e.g., Slater, 1950; Davis, 1955; Schaffner and Ferreira, 1995; Schwartz, 2004; Schaffner and Schwartz, 2008; Forero, 2008).
The eggs of Miridae are characterized by having an elliptical operculum, lacking any other major modifications (Southwood, 1956). Species of Carvalhomiris present a relatively large knoblike process on the posterior pole at the aft side of the egg (Figs. 10A, C-F). This is a highly unusual character because mirid eggs are rounded on their posterior pole, without any protuberance or processes on their surface (Kullenberg, 1942, 1943; Southwood, 1956; Ren, 1992). Nonetheless, Cobben (1968) mentioned that Pseudoloxops coecineus (Meyer-Dur, 1843) and Malacoeoris chlorizans (Panzer, 1794) have an "indented" posterior pole (Cobben, 1968: Fig. 158A). The egg of M. chlorizans was first studied and illustrated by Kullenberg (1942: 10, plate 1, fig. 9), who observed the rugose chorion but not the basal process later pointed out by Cobben (1968).
[FIGURE 5 OMITTED]
In the egg of Carvalhomiris, the posterior pole is not indented, but has a distinctly protruded process, which may be homologous to the structure observed by Cobben (1968). Besides M. chlorizans and P. coccineus, a similar egg structure has been illustrated for Z. breviceps (Doesburg, 1984: Figs. 13a-c). Doesburg (1984) described the long, subcaudal, slender process of the egg, but failed to compare it to any other Orthotylinae or mirid species and argued that it was a unique structure for this species. He concluded that this character should be surveyed more broadly in similar species (Doesburg, 1984), a conclusion with which I also agree.
The chorion in Carvalhomiris is deeply punctured and has elevated ridges (Figs. 10A, B). In most Miridae the chorion is either smooth or with faint polygonal sculpturation (Kullenberg, 1942, 1943; Southwood, 1956; Cobben, 1968; Ren, 1992). Malacocoris chlorizans, P. coccineus (Orthotylini), and Strongylocoris leucocephalus (Linnaeus, 1758) (Halticini) are the only other Miridae known to have a rugose chorion (Kullenberg, 1942, 1943; Cobben, 1968), although in S. leucocephalus there are minute spines on the chorion (Kullenberg, 1943: pl. 8, figs. 8a, b), which are not present in M. chlorizans or P. coccineus (Kullenberg, 1942: pl. 1, fig. 9; Cobben, 1968: Fig. 158a). A rough chorion is not a consequence of having the egg removed from the oviduct because Cobben (1968) examined several other Miridae species using the same methodology without observing this condition. On the contrary, Cobben (1968: 160) suggested that this rough chorion is related to the semi-exposed condition of the egg in M. chlorizans and P. coccineus.
In Carvalhomiris the metathoracic scent gland efferent area is not coplanar with the remaining metepisternum, but is recurved dorsad at the level of the peritreme, the latter being flat relative to the surrounding mushroomlike cuticle (Figs. 3A-D). In most Orthotylini, the peritreme and surrounding evaporatory area are flat, with the peritreme usually elongate and located medially on the evaporatory area (e.g., Schwartz, 2004; Schaffner and Schwartz, 2008; Forero, 2008). In Ceratocapsini and Austromirini (Orthotylinae), the peritreme is elevated above the plane of the evaporatory area, and not elongate (e.g., Henry, 1994, 2006; Cassis and Moulds, 2002; Cassis, 2008). The notched anterior margin of the peritreme found in Carvalhomiris is rare among Orthotylini, but present in at least some Ceratocapsini (e.g., Schaffneria Knight, Henry, 1994), and common in Phylinae (e.g., Schuh, 2006: Fig. 6B; Weirauch, 2006: Fig. 7). Some Orthotylini, nonetheless, do not share a flat condition of the scent gland efferent system. In Osornocoris punctatus Carvalho, 1985, a species from Chile, the peritreme is tumescent, and the evaporatory area is nearly flat and not recurved dorsad (pers. obs.), a condition similar to what is found in Ceratocapsini and Austromirini. In some species of Itaeoris, the evaporatory area is nearly recurved dorsad and the anterior margin of the scent channel is sinuate, conditions that are very similar to those found in Carvalhomiris (pers. obs.). Species of Paraproba also have a slightly protruding evaporatory area with respect to the remaining metepisternal cuticle, but in the few species examined the mushroomlike cuticle is not dorsally recurved. Clearly, a more thorough documentation on the external morphology of the scent gland efferent system is needed for Orthotylinae.
PUTATIVE SYNAPOMORPHIES: Discussions on character variability are impossible for monotypic genera. The discovery of two new species of Carvalhomiris now makes it possible to assess and propose potential synapomorphies for the genus. The proposed putative synapomorphies should be tested in a phylogenetic analysis with broad taxon sampling. I propose the following putative synapomorphies for Carvalhomiris: large genital capsule with cephalic, lateral margin strongly emarginate; strong brachypterism in both sexes; fight paramere clubbed, with apical margin serrate; vesica with a single, large, C-shaped spicule, with its base nearly reaching the cephalic end of the aedeagus; lateral infoldings of the sclerotized tings on the dorsal labiate plate with a posteriad acute process; and dorsal labiate plate with a cuplike depression projecting into the vestibulum.
RELATIONSHIPS OF CARVALHOMIRIS: Because the Orthotylini is the largest tribe of the nominal subfamily, Schuh (1974) proposed four informal generic groupings: the Falconia, Orthotylus, Sericophanes, and Zanchius groups, thus providing the most current concept of suprageneric categories. Some of these generic groupings may deserve tribal ranking. For instance, the Ceratocapsini of Henry (1994) is the same as the Sericophanes group of Schuh exclusive of Sericophanes Reuter.
Schuh (1974) included in the Zanchius group those species having a delicate body, flattened appearance, usually hyaline hemelytra, and vesica without spicules. Henry (1995) modified the concept of the group, adding as diagnostic characters the exposed mesoscutum and blunt, ventral process on the aperture of the genital capsule, commenting also that some species may have at least one spicule. Henry (1995) also suggested that members of the Zanchius group might deserve to be placed in their own tribe.
[FIGURE 6 OMITTED]
Maldonado and Ferreira (1971) compared Carvalhomiris with Aetorhinella Noualhier, from the Canary Islands, and Parachius from the Neotropical region. They stated that Carvalhomiris shares with these two genera the same "shape and appearance," and that the brachypterism and long antennae distinguish Carvalhomiris from them. Nonetheless, the antenna in Aetorhinella is longer than in Carvalhomiris, being nearly as long as the total length of the specimen (pers. obs.). Brachyptery is present in various degrees in mirid species, where sexual dimorphism and polymorphic species are not uncommon (Wheeler, 2001). In Orthotylinae and Phylinae, and particularly in Neotropical Orthotylini, brachypterism is rarely found in both sexes, being males commonly macropterous and females brachypterous (Schuh, 1974: 257). Therefore, the brachypterous condition in both sexes is apparently unique for Carvalhomiris. Besides the brachypterous condition, without specific morphological details, the comparison made by Maldonado and Ferreira (1971) is not helpful.
Neotropical genera included in the Zanchius group are: Brasiliomiris Carvalho, Hyalochloria Reuter, Itacoris, Jobertus Distant, Parachius, Paraproba, Pliniella Bergroth, Proboscidotylus Henry, and Saileria Hsiao (Schuh, 1974; Henry, 1995). Of these, only Itacoris, some of the species of Parachius (i.e., P. luteolus Distant, 1884, and P. virescens Carvalho and Gomes, 1968), and Paraproba have the eyes far removed from the pronotum. In these species, the eyes present a postocular swelling similar to the structure found in Carvalhomiris. In addition to the Neotropical taxa, some Palearctic and Oriental taxa, for instance Aetorhinella and some of the species of Zanchius, also possess a distinctive swelling (pers. obs.). Despite not having been mentioned by Schuh (1974), Aetorhinella can be placed in the Zanchius group because of its delicate body and flattened appearance. The structure of the metathoracic efferent area in Carvalhomiris, with the evaporatory area raised and recurved dorsad, is similar to what is found in some of the species of Itacoris, and with a slightly modified condition in species of Paraproba (pers. obs.). The conspicuous paired lateral flaplike protuberances on the base of phallotheca are also present in the examined species of Paraproba and in some undescribed members of the Zanchius group from Australia (pers. obs.), but its presence in other taxa of the Zanchius group is not known. The structure of the egg, with a basal process on the aft edge, is present in M. chlorizans, P. coccineus (Cobben, 1968), and Z. breviceps (Doesburg, 1984). Of these, Malacocoris and Zanchius are included in the Zanchius group, and Pseudoloxops in the Orthotylus group (Schuh, 1974). The deeply punctate chorion is also present in Malacocoris and Pseudoloxops (Cobben, 1968), but apparently is not present in Z. breviceps (Doesburg, 1984). Furthermore, the structure of the base of the claw, with three hairs, is present as far as is known to me, only in Carvalhomiris and Z. breviceps.
All the characters mentioned above for Carvalhomiris, rarely occur in other Orthotylini outside the Zanchius group. Carvalhomiris occupies an apparently isolated position among the Neotropical genera currently assigned to the Zanchius group. The robust aspect of the body, which is usually delicate in the other genera, and some differences found in the male genitalia are particularly significant. For instance, the genital capsule of several of these Neotropical genera (e.g., Itacoris, Paraproba, Proboscidotylus, Saileria) has a blunt ventral prolongation on the aperture of the genital capsule (Henry, 1994; D. Forero, pers. obs.). This character is not shared by Carvalhomiris, in which the ventral margin of the genital capsule is entire and the whole genital capsule is shortened (Fig. 7). Similarly, many of these genera have reduced and not well-sclerotized spicules on the vesica. In Carvalhomiris, by contrast, the vesica has a large, well-sclerotized spicule (Fig. 5). Therefore, because Carvalhomiris shares at least some of the characters discussed above with other genera included in the Zanchius group, I am proposing to include Carvalhomiris in this generic group as currently delimited.
Because of the lack of comprehensive documentation (e.g., no illustrations of vesical structures, Schuh, 1974) of the characters used to define the Zanchius group, Carvalhomiris is best placed in this suprageneric grouping until a phylogenetic analysis is carried out for the entire group.
[FIGURE 7 OMITTED]
KEY TO THE SPECIES OF CARVALHOMIRIS
1. Right paramere with ventral portion greatly elongated and apically bifurcated (Fig. 6); brachypterous male and female with apex of hemelytron acute (Fig. 1); medial margin of first gonapophysis nearly symmetrical (Fig. 9); second antennal segment dark brown with a broad, pale median band, sometimes band darker or indistinct against dark brown ends; base of third antennal segment pale ... bifurcatus sp. nov.
Right paramere neither greatly elongated nor strongly bifurcated on ventral portion (Fig. 6); brachypterous male and female with apex of hemelytron rounded (Fig. 1); anterior medial margin of first gonapophysis strongly asymmetrical (Fig. 9); second antennal segment with basal half pale brown or pale yellow, basal half of third antennal segment pale yellow ... 2
2. Apical serrated margin of right paramere nearly straight (Fig. 6); dorsal portion of vesica with apex small and straight (Fig. 5); dorsal process of left paramere relatively small (Fig. 5).. brachypterus Apical serrated margin of right paramere with a large, nearly truncate, median process (Fig. 6); dorsal portion of vesica with apex strongly curved to the left (Fig. 5); dorsal process of left paramere relatively large ... truncatus sp. nov. Carvalhomiris bifurcatus, sp. nov.
Figures 1; 2A, C, E, G; 3A, E, G; 4C-F; 5-9; 10 C, E, G; 11
DIAGNOSIS: Recognized by the relatively small size (Table 1); anterior lobe of pronotum nearly convex (Figs. 1, 2E, G); acute apex of the hemelytron in brachypterous specimens (Fig. 1); right margin of phallotheca with preapical acute process (Fig. 5); vesica with dorsal portion with a wide base and apex curved dorsad, ventral portion apically acute and barely longer than dorsal portion (Fig. 5); right paramere with ventral prolongation long, curved dorsad, and bifurcate apically (Figs. 3G, 6); left paramere with flat surface caudad to sensory area, apex with ventral large and acute process, and ventral preapical area protruding and covered with medium-sized spicules (Fig. 6); and by the nearly symmetrical anterior medial margin of the first gonapophysis (Fig. 9).
Carvalhomiris bifurcatus is easily distinguished from C. brachypterus and C. truncatus by the smaller size and strongly bifurcate apex of the right paramere in the male. Brachypterous males and females can be distinguished from C. brachypterus and C. truncatus by the acute apex of the hemelytron. Both macropterous and brachypterous males can be distinguished from males of C. brachypterous and C. truncatus by the basal fringe of short erect setae on antennal segment II. Macropterous specimens are readily distinguished from C. brachypterus and C. truncatus by the slightly convex anterior lobe of the pronotum.
DESCRIPTION: Brachypterous male: Total length 2.48-288. COLORATION: Overall coloration pale yellow. Head: Either pale yellow, or clypeus, frons, anterior part of vertex, maxillary plate, and gular area dark brown; a few specimens with reddish postocular region, in which case reddish marking extending to propleura and present also in medial surface of antennal segment I; antennal segment I dark brown, medial surface brown, II dark brown with a broad pale median band, sometimes not noticeable, III and IV brown, base of III pale. Thorax: Pronotum on lateral areas of anterior lobe dark brown; proepisternum and proepimeron brown; dorsal half of mesepisternum, mesepimeron, and metepisternum either pale yellow or dark brown. Legs: Sometimes femur, partially or entirely, greenish. VESTITURE: Thorax: Hemelytron: Surface with yellowish simple setae; apex with a dense patch of simple setae. STRUCTURE: Head: Antennal segment II on medial surface of basal third with a fringe of short, suberect, simple setae set in a dense patch. Thorax: Hemelytron: Relatively long, apex acute (Fig. 1). Legs: Hind femur on ventral surface with small sets of brochosomes (see Rakitov, 2002), and a few brochosomes on the trichoma at the base of the trichobothria. Genitalia: Right paramere with ventral portion of paramere elongated, strongly curved dorsad, apically strongly bifurcated with cephalic prong sinuate and longer than caudal prong, apical margin of paramere gently serrate, straight and reclined (Fig. 6); left paramere with area caudad to dorsal sensory area fiat, apex of paramere on ventral surface with a large and acute process, ventral preapical area with protruding area covered with medium-sized spicules (Fig. 6); right margin of phallotheca with a preapical acute process (Fig. 5); vesica with dorsal portion about three-fourths the length of ventral spicule, base wide, apex strongly curved upward; ventral portion of vesica not reaching apex of phallotheca, acute apically (Fig. 5).
Macropterous male (Fig. 1): Similar to brachypterous male in coloration and structure, except for thorax structure as stated in the generic description, and by the pronotal color variation; total length 4.21-4.66. COLORATION: Head: Antennal segment I completely dark brown. Thorax: Pronotum completely dark brown, sometimes with a longitudinal area of variable size pale brown; mesoscutum sometimes with posterolateral sides dark brown; scutellum dark brown, sometimes with anterior longitudinal pale area of variable size.
Brachypterous female (Fig. 1): Similar to brachypterous male in coloration and structure, but larger; total length 3.11-3.36. VESTITURE: Head: Antennal segment II with normal, decumbent, very short setae as in remaining surface of antennae. STRUCTURE: Thorax: Hemelytron: Relatively long, nearly reaching caudal margin of abdomen. Genitalia (Fig. 9): Anterior medial margin of first gonapophyses nearly symmetrical.
Macropterous female (Fig. 1): Similar to macropterous male in coloration and structure, except for thorax structure as stated in the generic description; total length 4.43-4.68.
DISTRIBUTION: Known from a single locality in Napo Province, Ecuador (Fig. 11).
HOSTS: No host plants have been recorded for this species.
ETYMOLOGY: The name is taken from the Latin "furcatus," forked, and alludes to the strongly bifurcate apex of the ventral portion of the fight paramere, a character that is easily observed in nondissected specimens (e.g., Fig. 3G).
[FIGURE 8 OMITTED]
Discussion: The ventral portion of the spicule in at least one specimen was slightly bent instead of being straight (Fig. 5 arrow, AMNH_PBI 00185284). I considered this configuration normal spicule variation, as has been documented for other species of Orthotylini (e.g., Aoplonema Knight, Forero, 2008: Figs. 7, 10, 13).
Both macropterous and brachypterous specimens show variability with respect to the coloration of the head, pronotum, scutellum (Fig. 1), and pleural sclerites. The coloration may vary from completely pale yellow to dark brown in these structures. Usually the variability is correlated in all of the structures at the same time, i.e., when the head is dark brown anteriorly, the pronotum and scutellum are nearly completely dark brown. If a specimen has a less strongly pigmented head, then the pronotum is usually paler medially, and the scutellum is pale anteriorly. Dark forms are most common in macropterous specimens, although relatively dark specimens may occur in brachypterous forms (Fig. 1).
Brochosomes (Rakitov, 2002) were observed on the surface of the hind femur of C. bifurcatus. Brochosomes are well documented for Cicadellidae (Hemiptera: Auchenorrhycnha) (e.g., Rakitov, 1999, 2002, 2004). Recent studies document the presence of brochosomes within other Hemiptera groups as well, mainly in Miridae (Heteroptera) (Wyniger et al., 2008; D. Forero, pers. obs.).
HOLOTYPE MALE: ECUADOR: Napo: Sierrazul, SW of Baeza, 0.66666[degrees]S 77.91666[degrees]W, 2200 m, 22 Jan 1996-30 Jan 1996, T. J. Henry, Holotype Carvalhomiris bifurcatus sp. nov. Forero det., 1 [male] macropterous (AMNH_PBI 00185790) (USNM).
PARATYPES: ECUADOR: Napo: Sierrazul, SW of Baeza, 0.66666[degrees]S 77.91666[degrees]W, 2,200 m, 22 Jan 1996-30 Jan 1996, T. J. Henry, 1[male] macropterous (AMNH_PBI 00185826), 2[male] brachypterous (AMNH_PBI 00185836-AMNH_PBI 00185837), 1[female] macropterous (AMNH_PBI 00185848), 2[female] brachypterous (AMNH_PBI 00185861-AMNH_PBI 00185862) (AMNH); 5[male] macropterous (AMNH_PBI 00185788--AMNH_PBI 00185789, AMNH_PBI 00185823--AMNH_PBI 00185825), 35[female] brachypterous (AMNH_PBI 00185791-AMNH_PBI 00185811, AMNH_PBI 00185827-AMNH_PBI 00185835, AMNH_PBI 00185838-AMNH_PBI 00185842), 14[female] macropterous (AMNH_PBI 00185843-AMNH_PBI 00185847, AMNH_PBI 00185778-AMNH PB100185786), 249 brachypterous (AMNH_PBI 00185777, AMNH_PBI 00185812-AMNH_PBI 00185822, AMNH_PBI 00185849-AMNH_PBI 00185860) (USNM).
Carvalhomiris brachypterus, Maldonado and Ferreira
Figures 1, 4B, 5-7, 9, 10F, 11
Carvalhomiris brachypterus Maldonado and Ferreira, 1971 [n. sp.]; Schuh, 1995:89 [catalog].
DIAGNOSIS: Recognized by the relatively moderate size (table 1); anterior lobe of pronotum nearly flat; apex of hemelytron rounded on brachypterous specimens (Fig. 1); right margin of phallotheca with preapical margin rounded, smooth (Fig. 5); vesica with apex of dorsal portion small, with a small preapical denticle, base nearly constricted; ventral portion of vesica about two times as long as dorsal portion, reaching caudal margin of phallotheca, tapering and expanded towards the apex (Fig. 5); right paramere with ventral prolongation straight, directed caudad, apex not bifurcated but with a small preapical spine, apical margin straight, vertical, interrupted by a medium-size spine located near the dorsal portion (Fig. 6); left paramere with a small tubercle caudad to dorsal sensory area, apex with a small rounded ventral process, and preapical ventral area with a nearly flat area of very small spicules (Fig. 6); and by the highly asymmetrical anterior medial margin of the first gonapophysis.
Carvalhomiris brachypterus is easily distinguished from C. bifurcatus by the structure of the right paramere, with its ventral portion straight and directed caudad, and relatively larger size. Brachypterous male and female specimens can also be distinguished from C. bifurcatus by the more rounded apex of the hemelytron. Carvalhomiris brachypterus is most similar to C. truncatus because of the size, coloration, and structure of the right and left parameres, and the vesica. Carvalhomiris brachypterus is nonetheless easily distinguished from C. truncatus by the straight apical margin of the right paramere, which has a medial, nearly truncate process in C. truncatus; by the smaller dorsal, serrate process on the left paramere, which is larger in C. truncatus; and by the straight and relatively small apex of the dorsal portion of the spicule, which is strongly curved in C. truncatus.
REDESCRIPTION: Brachypterous male: Total length 2.94-3.32. COLORATION: Overall coloration pale yellow. Head: Swelling of postocular region brown or dark brown; antennal segment I brown, medial surface pale brown, II dark brown with basal half pale brown, sometimes not clearly distinguished, III on basal half pale yellow, IV missing. Thorax: Pronotum with posterior margin sometimes pale green; proepistemum sometimes brown or dark brown; proepimeron sometimes pale brown; mesepistemum, mesepimeron, and metepistemum sometimes barely pale brown. Legs: Femora, mostly on dorsal surface, greenish. Abdomen: Green or dark brown. VESTITURE: Thorax: Hemelytron: Surface with yellowish simple setae; apex with sparse simple setae. STRUCTURE: Head: Antennal segment II on medial surface of basal third with decumbent, simple setae. Thorax: Hemelytron: Relatively short, apex rounded. Genitalia: Right paramere with ventral portion straight, directed caudad, with a small preapical spine, apical margin of paramere straight, vertical, serrate, with a medium-sized spine close to dorsal apex of paramere (Fig. 6); left paramere with area caudad to dorsal sensory area with a small, serrate dorsal process (Fig. 6, large black arrow), apex of paramere on ventral surface with a small and rounded process, ventral preapical area with a nearly flat area covered with very small spicules (Fig. 6); right margin of phallotheca with preapical area rounded, smooth (Fig. 5); vesica with dorsal portion about one half the length of ventral portion, apex short and straight, directed caudad, preapically with a small acute process, narrowing towards its base; ventral portion of vesica reaching or surpassing apex of phallotheca, apex tapering and expanded apically (Fig. 5, gray arrow).
[FIGURE 9 OMITTED]
Macropterous male." Unknown.
Brachypterous female (Fig. 1): Similar to brachypterous male in coloration and structure, but apparently larger, total length 3.27. STRUCTURE: Thorax: Hemelytron: Relatively short compared with abdominal length, reaching abdominal segment V. Genitalia (Fig. 9): Anterior medial margin of first gonapophyses highly asymmetrical.
Macropterous female: Similar to brachypterous female in structure and coloration, except for the structure of the thorax as stated in the generic description; total length 4.32.
DISTRIBUTION: Carvalhomiris brachypterus is known from at least two localities at middle elevations in Cundinamarca (Colombia), both of which are very close to Bogota (Fig. 11).
HOSTS: No host plants have been documented for this species.
DISCUSSION: Maldonado and Ferreira (1971) mentioned a macropterous female. I have examined this specimen, which is topotypic with the holotype. Females of C. brachypterus, both macropterous and brachypterous, are very similar to females of C. truncatus. Association with males is necessary in order to correctly identify the specimens.
Maldonado and Ferreira (1971) misinterpreted the structure of the male genitalia. The drawing provided in their paper for the aedeagus (Maldonado and Ferreira, 1971: Fig. 3) illustrated the dorsal portion of the vesica as a separate sclerotized element from the ventral portion of the spicule. They also overlooked the dorsal serrate tubercle of the left paramere because of the ventral orientation of the structure (Maldonado and Ferreira, 1971: Fig. 4). Both characters on the vesica and left paramere are important for species delimitation, and thus are redrawn in this paper to facilitate recognition of C. brachypterus.
HOLOTYPE MALE; COLOMBIA: Cundinamarca: [in] Oak forest, Mosquera, [4.7106[degrees]N 74.23397[degrees]W], 14 Aug 1965, J. A. Ramos, Holotype "Carvalhomiris brachypterus M & F det J. Maldonado C. 1971, 1[male] brachypterous (AMNH_PBI 00175090) (MNRJ).
PARATYPES: COLOMBIA: Cundinamarea: Mosquera, [in] Oak forest, [4.7106[degrees]N 74.23397[degrees]W], 14 Aug 1965, J. A. Ramos, 2[male] brachypterous (AMNH_PBI 00186010, AMNH_PBI 00186229), 1[male] brachypterous (AMNH_PBI 00186249) (USNM).
OTHER SPECIMENS EXAMINED: COLOMBIA: Cundinamarea: 30 km W of Bogota, Mosquera, [4.7106[degrees]N 74.23397[degrees]W], 1,800 m, Mar 1965, J. A. Ramos, 3[male] brachypterous (AMNH_PBI 0018601l, AMNH_PBI 00186230, AMNH_PBI 00186231) (USNM), 1[female] brachypterous (AMNH_PBI 00186247) (USNM). Mosquera, [in] Oak forest, [4.7106[degrees]N 74.23397[degrees]W], 22 May 1965, J. A. Ramos, 1[male] brachypterous (AMNH_PBI 00196221) (AMNH), 2[female] brachypterous (AMNH_PBI 00302184, AMNH_PBI 00302185) (JTP), 1[male] brachypterous (AMNH_PBI 00186248) (USNM). San Antonio de Tena, [4.61533[degrees]N 74.35269[degrees]W, 1,443 m], Oct 1965, J. A. Ramos, 1[male] brachypterous (AMNH_PBI 00302182) (JTP), 4[male] brachypterous (AMNH_PBI 00186012, AMNH_PBI 00186235-AMNH_PI 100186237) (USNM), 2[female] (AMNH_PBI 00246764, AMNH_PBI 00246765) (TAMU), 7[female] brachypterous (AMNH_PBI 00186240-AMNH_PBI 00186246) (USNM); 09 Oct 1965, J. A. Ramos, 1[male] brachypterous (AMNH_PBI 00196222) (AMNH), 2[male] brachypterous (AMNH_PBI 00317041, AMNH_PBI 00317042) (CNC), 3[male] brachypterous (AMNH_PBI 00186232-AMNH_PBI 00186234) (USNM), 1[female] brachypterous (AMNH_PBI 00302183) (JTP), 2[female] brachypterous (AMNH_PBI 00186238 AMNH_PBI 00186239) (USNM).
Carvalhomiris truncatus, sp. nov.
Figures 1; 2B, D, F, H; 3B-D, F, H; 4A; 5-9; 10A, B, D, H; 11
Carvalhomiris brachypterus: Schuh, 1976: 6, 9, Fig. 16 [misidentification, pretarsus].
DIAGNOSIS: Recognized by the relatively moderate size (table 1); anterior lobe of pronotum nearly fiat; apex of hemelytron rounded on brachypterous specimens (Fig. 1); right margin of phallotheca with preapical margin rounded, smooth (Fig. 5); vesica with apex of dorsal portion hooked, strongly curved to the left, base relatively narrow; ventral portion of vesica about two times as long as dorsal portion, reaching caudal margin of phallotheca, tapering and expanded towards the apex (Fig. 5); right paramere with ventral prolongation straight, directed caudad, apex not bifurcated but with a small preapical spine, apical margin interrupted medially by a large, nearly truncate process (Fig. 6); left paramere with a large, serrate tubercle caudad to dorsal sensory area, apex with a small rounded ventral process, and preapical ventral area with a nearly flat area of very small spicules (Fig. 6); and by the highly asymmetrical anterior medial margin of the first gonapophysis.
As mentioned above for C. brachypterus, C. truncatus is easily distinguished from C. bifurcatus by the straight and caudally directed ventral portion of the right paramere, and the relatively larger size. Brachypterous male and females specimens of C. truncatus can be distinguished from C. bifurcatus by the rounded apex of the hemelytron. Carvalhomiris truncatus is most similar to C. brachypterus in size, coloration, structure of parameres, and the vesica. Carvalhomiris truncatus is easily distinguished from C. brachypterus by the large, nearly truncate medial process on the apical margin of the right paramere; by the larger serrate process on the left paramere, which is smaller in C. brachypterus; and by the strongly left-hooked apex of the dorsal portion of the spicule, which is straight in C. brachypterus.
DESCRIPTION: Brachypterous male: Total length 3.10-3.53. COLORATION: Overall coloration pale yellow. Head: Clypeus and maxillary plate dark brown; postocular swelling reddish brown; antennal segment I dark brown, II and III dark brown with basal half pale yellow, II sometimes slightly darker at base, IV dark brown with base pale. Thorax: Proepisternum and proepimeron usually pale reddish brown. VESTITURE: Thorax: Hemelytron: Surface with dark brown simple setae; apex with sparse simple setae. STRUCTURE: Head: Antennal segment II on medial surface of basal third with decumbent, simple setae. Thorax: Hemelytron: Relatively short, apex rounded. Genitalia: Right paramere with ventral portion straight, directed caudad, with a small preapical spine, apical margin of paramere medially with a nearly truncate process (Fig. 6); left paramere with area caudad to dorsal sensory with a large, serrate dorsal process (Fig. 6, large black arrow), apex of paramere on ventral surface with a small and rounded process, ventral preapical area with a nearly flat area covered with very small spicules (Fig. 6); right margin of phallotheca with preapical area rounded, smooth (Fig. 5); vesica with dorsal portion about one half the length of ventral portion, apex strongly hooked, directed to the left, base relatively narrow; ventral portion of vesica reaching or surpassing apex of phallotheca, apex tapering and expanded apically (Fig. 5, gray arrow).
[FIGURE 10 OMITTED]
[FIGURE 11 OMITTED]
Macropterous male: Unknown.
Brachypterous female: Similar to brachypterous male in coloration and structure, but much larger, total length 3.82-4.10. STRUCTURE: Thorax: Hemelytron: Relatively short compared with abdominal length, reaching abdominal segment V. Genitalia (Fig. 9): Anterior medial margin of first gonapophyses highly asymmetrical.
Macropterous female: Similar to brachypterous female in structure and coloration, except for the structure of the thorax as stated in the generic description; total length 4.03-4.61.
DISTRIBUTION: Carvalhomiris truncatus is known from a single area with two adjacent localities in Cundinamarca (Colombia), both located at a relatively high elevation (Fig. 11).
HOSTS: Carvalhomiris truncatus has been collected on an unidentified species of Tropaeolum (Tropaeolaceae). Species of Tropaeolum have a mainly Andean distribution (Chavez, 2004), which fits the observed distribution of the species of Carvalhomiris.
ETYMOLOGY: The name of this species is taken from the Latin "truncus," cut off, because of its large, nearly truncate medial process of the right paramere.
DISCUSSION: Schuh (1976) discussed the pretarsal structure of Miridae, including various species of Orthotylinae. He showed a dorsal view of the pretarsus of a species of Carvalhomiris, which he assigned to C brachypterus (Schuh, 1976: 6, 9). I have examined the specimens that Schuh used in his paper, and they actually are C. truncatus.
Carvalhomiris truncatus shares several structural characters with C. brachypterus (see above), more than with C. bifurcatus. If this is related to some biogeographical pattern it is still unknown and additional undescribed species from unsampled areas may clarify this.
HOLOTYPE MALE: COLOMBIA: Cundinamarca: 39 km SW of Bogota, [4.43826[degrees]N 74.30070[degrees]W], 2,700 m, 15 Oct 1972, R.T. Schuh, Tropaeolum sp. (Tropaeolaceae), Holotype Carvalhomiris truncatus sp. nov. Eorero det., 1[male] brachypterous (AMNH_PBI 00196302) (AMNH).
PARATYPES: COLOMBIA: Cundinamarca: 39 km SW of Bogota, [4A3826[degrees]N 74.30070[degrees]W], 2,700 m, 15 Oct 1972, R.T. Schuh, Tropaeolum sp. (Tropaeolaceae), 21[male] brachypterous (AMNH_ PBI 00196223-AMNH_PBI 00196228, AMNH_PBI 00196293-AMNH_PBI 00196301, AMNH_PBI 00196303-AMNH_PBI 00196304, AMNH_PBI 00196307 AMNH_PBI 00196309, AMNH_PBI 00196313), 1[female] macropterous (AMNH_PBI 00196229), 69[female] brachypterous (AMNH_PBI 00196233, AMNH_PBI 00196315-AMNH_PBI 00196346, AMNH_PBI 00196354-AMNH_PBI 00196386) (AMNH); 1[male] brachypterous (AMNH_PBI 00196305), 1[female] brachypterous (AMNH_PBI 00196347) (ICN); 1[male] brachypterous (AMNH_PBI 00196306), 2[female] brachypterous (AMNH_PBI 00196348, AMNH_PBI 00196349) (UNAB); 3[male] brachypterous (AMNH_PBI 00196310-AMNH_ PBI 00196312), 4[female] brachypterous (AMNH_PBI 00196350-AMNH_PBI 00196353) (USNM).
OTHER SPECIMENS EXAMINED: COLOMBIA: Cundinamarca: 39 km SW of Bogota, [4.43826[degrees]N 74.30070[degrees]W], 2,700 m, 15 Oct 1972, R.T. Schuh, Tropaeolum sp. (Tropaeolaceae), 2 nymphs (AMNH_PBI 00196234, AMNH_PBI 00196314) (AMNH). Paramo de San Miguel, [4.45165[degrees]N 74.30009[degrees]W], 06 Feb 1965, J. A. Ramos, 1[female] macropterous (AMNH_PBI 00185787) (USNM).
APPENDIX 1. USI NUMBERS (WITH PREFIX AMNH_PBI) OF MEASURED SPECIMENS. Species Brachypterous Carvalhomiris [male]: 00185800, 00185801, 00185802, bifurcatus 00185803, 00185807 sp. nov. [female]: 00185813, 00185814, 00185815, 00185852, 00185851 Carvalhomiris [male]: 00186010, 00186012, 00186011, brachypterus 00302182 [female]: 00302185, 00186239, 00186240, 00186242, 00186243 Carvalhomiris [male]: 00196302, 00196294, 00196295, truncatus 00196296, 00196297 sp. nov. [female]: 00196347, 00196344, 00196341, 00196339, 00196337 Species Macropterous Carvalhomiris [male]: 00185823, 00185825, 00185826, bifurcatus 00185788, 0018579 sp. nov. [female]: 00185778, 00185779, 00185781, 00185783, 00185784 Carvalhomiris [female]: 00186249 brachypterus Carvalhomiris [female]: 00185787, 00196229 truncatus sp. nov.
This paper is prepared in partial fulfillment of the requirements of the Graduate School towards a PhD in Entomology at Cornell University. This project
was supported by, and represents a contribution to, the NSF Planetary Biodiversity Inventory grant DEB-0316495 to Randall T. Schuh (AMNH) and Gerasimos Cassis (University of New South Wales, Australia).
I want to particularly acknowledge, and thank, Thomas J. Henry (USNM) and Randall T. Schuh (AMNH) for improving the knowledge of Carvalhomiris by collecting large series of specimens in seldom collected areas of South America, which produced most of the specimens examined in this paper, and for making those specimens available to me. The following curators and individuals responsible of the collections kindly loan me additional material for study: Michael D. Schwartz (CNC), John T. Polhemus (JTP), Luiz A. A. Costa (MNRJ), and Joseph C. Schaffner and Edward C. Riley (TAMU).
Thomas J. Henry (USNM), James K. Liebherr (Cornell University, Ithaca), Randall T. Schuh (AMNH), and Michael D. Schwartz (CNC) kindly reviewed an early version of the manuscript and provided useful comments. Michael D. Schwartz further read the manuscript in its final version. Thomas J. Henry and Gerasimos Cassis provided useful discussions on the structure of the metathoracic efferent system of several Miridae taxa. Denise Wyniger (Natur-Museum Luzern, Switzerland) discussed with me the presence of brochosomes in C. bifurcatus.
Received 17 December 2008; accepted 6 April 2009
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DIMITRI FORERO (1,2)
Division of Invertebrate Zoology (Entomology), American Museum of Natural History, New York, New York 10024-5192; and Department of Entomology, Comstock Hall, Cornell University, Ithaca, New York 14853-2601, USA
(1) Present address: Department of Entomology, University of California Riverside, CA 92521, USA.
(2) E-mail address for correspondence: firstname.lastname@example.org
Table 1. Measurements of species of Carvalhomiris. Cun = Cuneus; Clyp = Clypeus; Pron = Pronotum; Scut = Scutellum; IntOcDi = Interocular distance; Ant1 = Antennal segment 1; Ant2 = Antennal segment 2. See "Material and Methods" section for details. Length Species Total Cun-Clyp Head Pron Carvalhomiris bifurcatus sp. nov. Brachypterous Mean 2.74 2.59 0.50 0.52 males (N = 5) SD 0.16 0.10 0.01 0.02 Range 0.40 0.25 0.03 0.05 Min. 2.48 2.42 0.49 0.50 Max. 2.88 2.66 0.52 0.55 Brachypterous Mean 3.21 3.35 0.49 0.55 females (N = 5) SD 0.09 0.09 0.03 0.02 Range 0.25 0.23 0.07 0.05 Min. 3.11 3.21 0.44 0.53 Max. 3.36 3.43 0.51 0.58 Macropterous Mean 4.45 3.08 0.47 0.61 males (N = 5) SD 0.20 0.10 0.01 0.02 Range 0.45 0.24 0.04 0.06 Min. 4.21 2.99 0.45 0.59 Max. 4.66 3.23 0.49 0.65 Macropterous Mean 4.54 3.25 0.51 0.63 females (N = 5) SD 0.10 0.05 0.01 0.02 Range 0.25 0.11 0.03 0.06 Min. 4.43 3.20 0.49 0.59 Max. 4.68 3.31 0.53 0.65 Carvalhomiris brachypterus Brachypterous Mean 3.10 2.68 0.51 0.52 males (N = 4) SD 0.16 0.06 0.03 0.01 Range 0.38 0.15 0.06 0.03 Min. 2.94 2.61 0.47 0.51 Max. 3.32 2.76 0.53 0.54 Brachypterous Mean 3.34 2.77 0.49 0.52 females (N = 5) SD 0.10 0.11 0.02 0.03 Range 0.25 0.27 0.04 0.06 Min. 3.23 2.57 0.47 0.49 Max. 3.48 2.85 0.51 0.55 Macropterous Value 4.32 3.36 0.44 0.57 female (N = 1) Carvalhomiris truncatus sp. nov. Brachypterous Mean 3.23 2.81 0.50 0.54 males (N = 5) SD 0.17 0.11 0.02 0.01 Range 0.43 0.22 0.06 0.03 Min. 3.10 2.71 0.48 0.53 Max. 3.53 2.93 0.53 0.56 Brachypterous Mean 3.90 3.06 0.55 0.55 females (N = 5) SD 0.12 0.05 0.02 0.01 Range 0.28 0.12 0.04 0.03 Min. 3.82 3.00 0.54 0.54 Max. 4.10 3.12 0.58 0.57 Macropterous Mean 4.32 3.36 0.44 0.57 females (N = 2) SD 0.41 0.20 0.02 0.03 Range 0.59 0.28 0.02 0.04 Min. 4.03 3.21 0.43 0.55 Max. 4.61 3.50 0.46 0.59 Length Width Species Scut Cuneus Head Pron Scut Carvalhomiris bifurcatus sp. nov. Brachypterous Mean 0.28 -- 0.64 0.70 0.36 males (N = 5) SD 0.01 -- 0.01 0.02 0.02 Range 0.03 -- 0.01 0.05 0.06 Min. 0.26 -- 0.63 0.67 0.34 Max. 0.29 -- 0.64 0.72 0.40 Brachypterous Mean 0.27 -- 0.85 0.97 0.33 females (N = 5) SD 0.02 -- 0.44 0.48 0.09 Range 0.05 -- 0.99 1.09 0.21 Min. 0.24 -- 0.65 0.73 0.17 Max. 0.29 -- 1.64 1.82 0.38 Macropterous Mean 0.35 0.83 0.65 1.07 0.47 males (N = 5) SD 0.02 0.06 0.02 0.05 0.02 Range 0.04 0.16 0.06 0.11 0.05 Min. 0.33 0.75 0.63 1.01 0.45 Max. 0.37 0.90 0.69 1.12 0.50 Macropterous Mean 0.35 0.79 0.66 1.09 0.48 females (N = 5) SD 0.02 0.04 0.02 0.02 0.02 Range 0.07 0.09 0.04 0.05 0.05 Min. 0.31 0.74 0.64 1.06 0.44 Max. 0.38 0.83 0.68 1.12 0.50 Carvalhomiris brachypterus Brachypterous Mean 0.28 -- 0.94 1.13 0.31 males (N = 4) SD 0.01 -- 0.56 0.70 0.08 Range 0.01 -- 1.14 1.41 0.19 Min. 0.28 -- 0.64 0.76 0.20 Max. 0.29 -- 1.78 2.18 0.39 Brachypterous Mean 0.27 -- 0.71 0.82 0.46 females (N = 5) SD 0.03 -- 0.01 0.04 0.04 Range 0.06 -- 0.04 0.09 0.11 Min. 0.25 -- 0.69 0.78 0.39 Max. 0.31 -- 0.72 0.87 0.50 Macropterous Value 0.40 0.65 0.72 1.22 0.61 female (N = 1) Carvalhomiris truncatus sp. nov. Brachypterous Mean 0.33 -- 0.71 0.88 0.35 males (N = 5) SD 0.03 -- 0.01 0.03 0.01 Range 0.09 -- 0.02 0.06 0.03 Min. 0.30 -- 0.70 0.85 0.33 Max. 0.39 -- 0.72 0.91 0.37 Brachypterous Mean 0.32 -- 0.73 0.93 0.35 females (N = 5) SD 0.04 -- 0.01 0.02 0.02 Range 0.08 -- 0.03 0.05 0.05 Min. 0.28 -- 0.71 0.90 0.32 Max. 0.36 -- 0.75 0.96 0.37 Macropterous Mean 0.40 0.65 0.72 1.22 0.61 females (N = 2) SD 0.03 0.26 0.02 0.03 0.05 Range 0.05 0.37 0.03 0.05 0.07 Min. 0.38 0.46 0.70 1.20 0.57 Max. 0.42 0.84 0.73 1.24 0.64 Width Species IntOcDi Ant1 Ant2 Carvalhomiris bifurcatus sp. nov. Brachypterous Mean 0.31 0.64 1.53 males (N = 5) SD 0.01 0.01 0.07 Range 0.03 0.03 0.15 Min. 0.29 0.63 1.47 Max. 0.33 0.66 1.63 Brachypterous Mean 0.53 0.65 1.43 females (N = 5) SD 0.45 0.02 0.06 Range 1.00 0.05 0.15 Min. 0.32 0.63 1.33 Max. 1.33 0.68 1.48 Macropterous Mean 0.32 0.63 1.49 males (N = 5) SD 0.02 0.03 0.07 Range 0.05 0.06 0.16 Min. 0.30 0.60 1.41 Max. 0.36 0.66 1.58 Macropterous Mean 0.33 0.63 1.45 females (N = 5) SD 0.01 0.04 0.05 Range 0.01 0.10 0.12 Min. 0.32 0.57 1.39 Max. 0.33 0.67 1.52 Carvalhomiris brachypterus Brachypterous Mean 0.66 0.63 1.55 males (N = 4) SD 0.63 0.01 0.11 Range 1.26 0.02 0.15 Min. 0.34 0.62 1.48 Max. 1.60 0.64 1.63 Brachypterous Mean 0.36 0.61 1.45 females (N = 5) SD 0.01 0.06 0.09 Range 0.03 0.13 0.26 Min. 0.35 0.54 1.33 Max. 0.37 0.67 1.59 Macropterous Value 0.36 0.62 1.40 female (N = 1) Carvalhomiris truncatus sp. nov. Brachypterous Mean 0.36 0.71 1.75 males (N = 5) SD 0.00 0.03 0.08 Range 0.01 0.09 0.21 Min. 0.36 0.67 1.63 Max. 0.36 0.75 1.84 Brachypterous Mean 0.37 0.68 1.62 females (N = 5) SD 0.01 0.01 0.05 Range 0.02 0.03 0.13 Min. 0.36 0.67 1.55 Max. 0.38 0.70 1.67 Macropterous Mean 0.36 0.62 1.40 females (N = 2) SD 0.01 0.02 0.09 Range 0.01 0.03 0.12 Min. 0.36 0.60 1.33 Max. 0.37 0.63 1.46
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