Jornandes Distant

Schaffner, JC & Schwartz, MD, 2008, Revision Of The Mexican Genera Ficinus Distant And Jornandes Distant With The Description Of 21 New Species (Heteroptera: Miridae: Orthotylinae: Orthotylini), Bulletin of the American Museum of Natural History 2008 (309), pp. 1-87 : 18-22

Jornandes Distant

Jornandes Distant, 1884: 301 (orig. desc.); Atkinson, 1890: 48 (catalog); Kirkaldy, 1906: 146 (catalog); Reuter, 1910: 163 (catalog); Carvalho, 1952: 77 (catalog); Carvalho, 1955: 76 (key); Carvalho, 1958: 80 (catalog); Carvalho and Schaffner, 1973: 69 (note); Schuh, 1995: 127 (catalog).

TYPE SPECIES: Jornandes championi Distant, 1884 (by monotypy).

Rhinocapsidea Reuter, 1908: 54 (orig. desc.); Carvalho, 1952: 79 (catalog); Carvalho, 1955: 78 (key); Carvalho, 1958: 130 (catalog); Schaffner,

1979: 74 (note); Schuh, 1995: 191 (catalog). NEW SYNONYM.

TYPE SPECIES: Eccritotarsus genetivus Distant, 1884 (by monotypy).

DIAGNOSIS: Characterized by the declivous head (figs. 17B, 20B, 28B) having the posterior margin carinate, the usually relatively short labium, the shiny appearance of the head and body, the impunctate pronotum and head, the distinctive minute sculpturing on corium and clavus (figs. 1F, 17A, 20A, 28A) rarely extending onto pronotum ( J. cruralis fig. 17B), and features of the male genitalia, especially the vesica, parameres, and tergal process. The relatively long anteocular portion of the head in comparison to the height of the eye in lateral view of Jornandes is superficially reminiscent of other North American Orthotylini (e.g., Ilnacora , Ilnacorella ). In Jornandes the eye height is always greater than the anteocular region and the frons is at most gently convex, with the vertex deeply transversely concave. In Ilnacora and Ilnacorella the anteocular region is always greater than the eye height, the frons is usually tumid, sometimes greatly so, and the vertex is slightly concave. The male genitalia in these two orthotyline genera are of a different form and the presence of scalelike setae is not known for the species of Jornandes .

REDESCRIPTION: STRUCTURE: Head: Strongly declivous, smooth and shining; vertex weakly transversely sulcate, posterior margin marginate or carinate; frons slightly to moderately rounded, usually shallowly separated from clypeus; anteocular distance variable, length ranging from 1/2 of eye height to subequal to eye height; maxillary plate prominent; buccula narrow, eye elongate as seen from side, located at posterior margin of head; antennal socket touching margin of eye; antennal segment I usually shorter than vertex width; antennal segment II ranging from slender and somewhat clavate to linear and more robust; diameters of segments III and IV variable, sometimes as great as segment II but usually less; relative lengths of segments from shortest to longest 1–4–3–2 or rarely 4–1–3–2; labium relatively short, usually not reaching posterior margin of mesosternum, occasionally extending onto metasternum. Pronotum: Shining, ranging from smooth to faintly rugulose; more or less triangular in shape; not strongly declivous; lateral margins rounded, calli smooth and weakly delimited; pleural and sternal regions except prosternum shining; small evaporative area ventral to mesothoracic spiracle (figs. 17C, 20C, 28C); metepisternum dorsal to evaporative region of scent gland with microtrichia continuing dorsally (figs. 17C, 20C) (except in J. burserae , J. rachelleae (fig. 28C), and J. viridulus ) on at least part or all of episternum; mesoscutum not exposed or only slightly so; scutellum slightly convex; often faintly transversely rugose. Hemelytron: Corium minutely and characteristically sculptured (figs. 1F, 17A, 20A, 28A), shining; usually curved downward laterally; embolium often not clearly delimited except at base; Legs: Coxa usually with pollinose areas (composed of microtrichia). Pretarsus: Claw strongly curved; pulvillus minute or small and not extending beyond medial curve of claw; parempodium apically convergent. COLOR- ATION: Variable, ranging from entirely pale, pale green, or with discrete brown marks to entirely black; sometimes variable portion of head, antenna, pronotum, and base of hemelytron orange red, posterior margin of pronotum rarely pale; legs variable; membrane generally fuscous but paler on species with paler body color. VESTITURE: Head consisting of a few usually short, scattered setae; antennal segment I with several long erect setae, one or two of which about as long as diameter of segment; segments II–IV with decumbent setae and some species also with erect setae; pronotum with single erect seta arising from each anterior corner posterior to eye; hemelytra ranging from almost glabrous to with long semierect setae; membrane frequently with a few semierect setae (not seen on some species); hind tibia with long erect setae; abdomen shining with elongate setae. GENITALIA: Genital segment with variable tergal process(es) projecting from right of, or at midline of, dorsal margin of aperture; ventroposterior margin of capsule with either notch or flange; subgenital plate usually situated asymmetrically near left paramere fossa; distal width of subgenital plate narrow to moderately wide, either projecting just beyond or dorsal to aperture of capsule on right side. Left paramere Cshaped in dorsal view; usually without produced sensory lobe; diameter of variable width, usually with subapical constriction; apex with terminal notch, mittenlike. Right paramere variable, elongate or C-shaped; sensory lobe variable, not developed, with broad process, or with long narrow spine; marginal surface and apex variable: smooth, serrate, or with obvious spines. Phallotheca length and shape variable, conical or rectangular; aperture sinuate, jagged, or with deep notch on right side, dorsally with narrow slot or open widely. Vesica with one prominent spiculum usually attached to right dorsal surface of proximal region of sclerotized part of ductus seminis; spiculum usually with sinuate ‘‘trunk’’, medial part often expanded; distal region of spiculum with variably shaped ‘‘branch(es)’’; usually branch(es) recurved or projecting to right side, partially serrate, narrow or needlelike diameter, apically point- ed; sometimes narrow, curved, pointed spine protruding from middle of spiculum trunk or posterior edge of spiculum base; sometimes spiculum with flattened, truncate process projecting from anterior portion of base.

Female: Coloration and body form, except for the smaller eye and wider vertex, usually same as for male. GENITALIA: base of ovipositor situated anterior to middle of abdomen; subgenital plate broadly triangular or shield shaped. In ventral view (fig. 19C) base of interior valvulae (gonapophyses 8, GP8) and adjacent vestibulum strongly sclerotized and complexly convoluted on each side of vulva (VUL), either side in repose overlapped by other side and sometimes overlapped by adjacent convoluted ventral margin of ventral labiate plate (VLP); dorsal surface of VLP with microtrichia, ventral surface of VLP sometimes broadly or narrowly produced ventrally into vestibular opening; in dorsal view; sometimes right gonapophyses 8 with tubercle projected immediately ventral to medial surface of subgenital plate (fig. 19C). Sclerotized rings moderately large to large, ovate, forming medial surface of folded, strongly sclerotized, dorsal labiate plate (DLP), folded medial region of DLP variable in size, triangular, moderately or strongly projected medially; DLP ventral to common oviduct sometimes formed by bilateral pair of irregularly shaped sclerotized plates, otherwise area membranous; DLP elongate in lateral view. Posterior wall in posterior view (fig. 19A, B) composed of paired interramal sclerites, merging with broad, well-sclerotized medial section; medial section sometimes with convex mound or tubercle projecting posteriorly, and large, interramal lobes (IRL) with strongly spinose dorsal surfaces, shape variable, either entire or with incised ventral margins; if IRL incised, then dorsal portion of IRL shorter than ventral portion, ventral portion sometimes narrow, apex blunt or pointed.

DISCUSSION: The species of this genus are diverse in color, shape, and size. Some species are primarily dark fuscous to black whereas others are relatively brightly colored and occasionally with attractive patterns. The costal margins of the hemelytron may be parallel or curved and the lengths of the insects vary from 1.75 to 5.00 mm. The vestiture is also quite variable in length and density. All species of Jornandes have the distinctive minute sculpturing of the clavus and corium (figs. 1F, 17A, 20A, 28A), declivous head, and relatively short labium.

The structure of the male genitalia of the included spp. is diverse with the shape of the right paramere and vesical spiculum appearing especially species specific. However, within this array of genitalic form are several features that we consider diagnostic for the genus. The vesica has a solitary spiculum that in the majority of the included spp. is recurved apically, regardless of the number of accessory branches. The apical region of the left paramere is always notched, providing for medial and lateral lobes and a mitten shape. All but three species ( J. crotoni , J. mimosae , and J. tehuacanensis ) have tergal processes on the dorsal margin of the genital aperture originating to the right of, or at, the middle.

We examined the female genitalia for all the included spp. except J. championi and J. variabilis , for which only one female is known for each taxon. The structure of the female genitalia is as diverse as that encountered in the male genitalia. We provide photographs of selected features of the female genitalia for three Jornandes spp. (fig. 19) to document a small portion of the variation. Any statements concerning diagnostic characters of the female genitalia would be premature.

Comparative information on the female genitalia is not available for all members of the group of Mexican genera we hypothesize to be related to Jornandes based on the roughly similar structure of the male genitalia (see ‘‘Delimitation of Taxon’’). Based on our still incomplete survey of North American Orthotylini genera, we can report here that attributes of the posterior wall (including a diversity of structure in the lateral and medial portions of the interramal sclerite and the interramal lobes) and dorsal labiate plate (including variation in the conformation of the sclerotized rings) observed in the type species of Ficinus , Fulgenticapsus , Jornandinus , Ilnacora , Lopidella , Lopidea , Oaxacaenus , Rolstonocoris , Scalponotatus , and Slaterocoris are not found in the included spp. of Jornandes . Drawing further inference regarding the relationships of these genera to Jornandes is beyond the scope of the current paper.

Whether the fused lateral and medial sections of the interramal sclerite and the usually modified ventral surface of the ventral labiate plate projecting ventrally into the vulvar region are diagnostic for spp. of Jornandes , and to a more limited extent in Ficinus (see generic descriptions above), will await a comprehensive study. All the related genera listed above and Jornandes have extensively sclerotized and convoluted gonapophyses 8 and ventral labiate plates. This feature is widely distributed in the Orthotylini ( Pluot-Sigwalt and Matocq, 2006) and as such cannot be deemed diagnostic for any of these genera. When females of more species are examined we suspect that the vulvar area and the shape of the interramal lobe will prove to be as diverse and species specific as the right paramere and the vesical spiculum are in the male genitalia (see illustrations of the female genitalia for Orthotylus spp. and Nesiomiris spp. in Southwood, 1953, and Gagne, 1997, respectively).

Lacking a phylogenetic analysis we can only propose several tentative species groups for the genus based on external appearance and male genitalia. Jornandes ater and J. xochipalensis : similar highly polished color pattern, antennal segment II with few, long setae, short labium, and the right paramere with a narrow, basal tubelike process. Jornandes burserae and J. viridulus : pale greenish yellow with little fuscous coloration, subequal vertex width and length of antennal segment I, semierect setae dorsal vestiture, male genitalia with elongate apex of the right paramere, C-shaped left paramere, and vesica with trifid apical region and relatively long basal branch or process. Jornandes ceibae and J. susanae : patterned dorsal and head coloration, antennal segment II dark and segment I pale, eye sexually dimorphic (larger in male), subequal vertex width and antennal segments I and II, and male genitalia with medially placed tergal process, C-shaped left paramere with a small apical notch, and vesica with similarly curved and laterally serrate distal region. Jornandes cruralis , J. genetivus , and J. sinaloa : large, parallel-sided costal margins, vesical spiculum with bifurcated terminal branches, long right paramere, and mostly spinose tergal processes (in J. cruralis and J. sinaloa ) Jornandes jaredi and J. zapotecas : parallel-sided costal margin, long, decumbent, evenly distributed dorsal setae, pale antenna and legs, eye sexually dimorphic (larger in male), male genitalia with small, apically serrate vesical spiculum, and large tergal processes on right side of genital aperture. Jornandes mimosae and J. tehuacanensis : although with dissimilar dorsal vestiture, the head in both sexes entirely orange brown, female antennal segment II with pale annulus medially, and male genitalia including a small right paramere with spinose dorsal lobe, moderately long Cshaped left paramere with wide apical notch, vesical spiculum bifid, with one long and one short serrate process, and genital aperture without tergal processes.

We cannot be certain of the generic placement of two species currently assigned to Jornandes . Distant (1893) described J. dissimulans from Guerrero, Mexico based on a single female. Carvalho (1958) originally considered it to be the junior synonym of subalbicans , swayed by the similar facies and identical type locality of the two nominal taxa. This synonymy was maintained when Carvalho and China (1959) erected the new genus Amulacoris to accommodate subalbicans . Subsequently Carvalho (1981) restored the species status of dissimulans within Amulacoris , and later concluded ( Carvalho, 1988) that it was not congeneric with A. subalbicans . We cannot describe the male genitalia, having failed to associate the holotype of dissimulans with any known species of Jornandes , Scalponotatus , and Slaterocoris . Because its cuticular sculpturation (fig. 1A) is not formed by minute, densely set punctures we can only support Carvalho’s conclusion that J. dissimulans be considered incertae sedis.

The assignment of nordestina to Jornandes is suspect based on its Brazilian distribution, cuticular sculpturation, and male genitalia. None of the specimens examined for this study are from further south than northern Guatemala. Even though a photo of the holotype seems to indicate that the cuticular texture of the clavus and corium might be composed of densely set punctures, it is not conclusive. The color pattern of the dorsum with the scutellum concolorous with the head, pronotum, and base of the hemelytron is not found in species of Jornandes . If the scutellum is pale in any of the included spp., then the base color of the entire body is pale. The strongest evidence against continued placement of nordestina in Jornandes is the unique form of the male genitalia. Whereas most species of Jornandes have a single recurved vesical spiculum, with minor basal or medial branches, the spiculum of nordestina is broad basally with two large multidissected, recurved branches. Additionally, the parameres are unlike any Mexican taxon, especially the left paramere, which does not have a mittenlike apex, but conspicuously has a large basal process longer than the length of the entire paramere (see Carvalho and Wallerstein, 1978: figs. 8–10). The correct placement of nordestina may necessitate erecting its own generic-level taxon. Such action is beyond the scope of the present study, so we propose incertae sedis status for J. nordestina .

The distribution of the Jornandes is primarily Mexico with a single species ( Jornandes championi Distant ) occurring in Guatemala (fig. 10).

Host plant associations are known for 13 of the 27 species considered in this paper. A summary of the host genera and associated families are as follows: Montana and Parthenium (Asteraceae) , Bursera (Burseraceae) , Ipomoea (Convolvulaceae) , Croton (Euphorbiaceae) , Desmanthus , Mimosa ( Fabaceae : Mimosoideae ), Salvia (Lamiaceae) , Ceiba (Malvaceae) , Recchia (Simaroubaceae) , and Heliocarpus (Tiliaceae) . This diverse group ranges from roadside plants ( Croton ), to bushes ( Mimosa ), and small ( Ipomoea ) or large trees ( Ceiba ). Only Jornandes genetivus is found on more than one plant family, with hosts in the Asteraceae , Convolvulaceae , and Euphorbiaceae . The other species of Ficinus and Jornandes are known only from a single species of plant. Discerning a pattern of host utilization in the species of these two genera obviously awaits further knowledge of their plants hosts.

Of the 27 recognized species of Ficinus and Jornandes , 19 are distributed across an arid band extending from Tehuacan and Coxcatlan, Puebla, on the east to Iguala and Chilpancingo, Guerrero, on the west (figs. 9– 10). This is a distance of only approximately 240 km (150 miles) that ranges from roughly 1,200 to 1,850 meters (4,000 –6,000 feet), but spreads over portions of three biogeographic provinces as listed by Morrone (2006). The area is located at the southern edge of the Transmexican Volcanic Belt Province at its confluence with the Balsas Basin and Sierra Madre del Sur provinces.

Five species outside this area occur at localities in western Mexico extending northward to the states of Sinaloa, Sonora, and Baja California Sur. Two species occur only in southern Oaxaca, with Jornandes championi known only from the Guatemalan type specimen. Besides three specimens of J. genetivus , no other species were found from Chiapas, however a few unusable specimens on hand suggest that two undescribed species occur there. No members of the Ficinus and Jornandes are known to occur on the Mexican Plateau or in the eastern coastal region.

With three exceptions, specimens seen in this study were collected in the months of July and/or August. This is consistent with the rain patterns found commonly in most arid areas of Mexico. The rainy season, which initiates vegetation growth, usually starts in June and lasts into September. It would appear that only a single generation of these insects occurs per year. A limited amount of rain sometimes occurs in March and could conceivably produce a generation; however, no specimens were collected during the subsequent months.

CHECKLIST OF SPECIES- GROUP NAMES PROPOSED IN OR CURRENTLY USED IN JORNANDES (valid names are printed in bold)