Tijdschrift voor Entomologie 159 (2016) 9-75
Revision of the genus Ochterus (Hemiptera: Heteroptera: Ochteridae) in Mesoamerica and the United States Dan A. Polhemus & John T. Polhemus1
The genus Ochterus is revised for Mesoamerica and the continental United States. The fauna as currently understood contains ten previously described species, and the following ten new species described herein: O. chiapensis from southeastern Mexico, O. costaricensis from Costa Rica, Panama and El Salvador, O. explanatus from western Mexico, O. mexicanus from central Mexico, O. obscurus from Costa Rica, O. ovatus from southeastern Mexico, O. panamensis from Panama, O. pseudorotundus from northeastern Mexico, O. shepardi from Costa Rica, and O. smaragdinus from southeastern Mexico. Ochterus flaviclavus Barber is shown to represent an intraspecific color morph of O. banksi Barber and is synonymized under the latter species. Color photographs of the dorsal body and frons, as well as line drawings of male genitalic structures, are provided for all species treated, accompanied by distribution maps. Keywords: Ochteridae; Mesoamerica; new species; new records; distribution maps Dan A. Polhemus*, Department of Natural Sciences, Bishop Museum, 1525 Bernice Street, Honolulu, HI 96817, [email protected] John T. Polhemus1, Colorado Entomological Institute, 3115 S. York Street, Englewood, CO 80112, USA.
Introduction radiation; in addition, all species occurring in North As noted by D. Polhemus & J. Polhemus (2012) the America have also been included for the sake of com- family Ochteridae exhibits two major geographic pleteness, and because in many cases broader distri- foci of species radiation, one in the Neotropical butional data are now available for many of these region (Champion 1901; Barber 1913; Schell 1943; taxa than is reflected in the published record. To pro- Drake 1952; J. Polhemus & M. Polhemus 1976; vide adequate context, extralimital material from Menke 1979; Cordiero 2014; Herrera Madrigal northern South America or the Caribbean is also 2014) and another in Melanesia (Kormilev 1971, listed in those cases where such records help to clarify 1973, Baehr 1990a, b) respectively, with a smaller the overall ranges of certain species. Mesoamerica as secondary radiations in Southeast Asia (Gapud & treated herein is defined as the region extending San Valentin 1977, Nieser & Chen 1992, Polhemus from Mexico to Panama, and including the interven- & Polhemus 2012) and the islands of the Indian ing countries of Belize, Guatemala, El Salvador, Ocean (Polhemus 1992, Polhemus & Polhemus Honduras, Nicaragua, and Costa Rica. This region UNCORRECTED PROOF 2008). The core of the current work is a treatment of contains at least ten endemic species that are not the Mesoamerican component of the Neotropical shared with North America, South America, or the
Tijdschrift voor Entomologie 159: 9–75, Table 1, Figs 1–64. [ISSN 0040-7496]. brill.com/tve © Nederlandse Entomologische Vereniging. Published by Koninklijke Brill NV, Leiden. Published 10 May 2016. DOI 10.1163/22119434-15901001 *Corresponding author 1Deceased Downloaded from Brill.com10/08/2021 06:09:55PM via free access
Caribbean Islands, and as such stands as a discrete live to see the project completed. In recognition of area of endemism for Ochteridae within the Western his substantial contribution, the first author has Hemisphere. therefore taken it upon himself to finish this work, Previous synthetic work on the Ochteridae of while retaining Dr Polhemus as a posthumous sec- North America and Mesoamerica consists of the ond author. foundational work of Champion (1901) in the Biologia Centrali-Americana, followed by the monograph of Schell (1943). By 1913 the eastern Morphological Characters North American ochterid biota had been fully Figs 1–15. described, and was well covered by Schell (1943), with only supplemental locality records added Characters defining Ochteridae among the nepo- here. In Mesoamerica, two additional species, one morphan families are their large eyes; the presence of of which also has limited occurrence in the south- ocelli; slender four-segmented antennae; a long western deserts of the United States, were described labium with segment III much longer than the oth- subsequent to Schell’s monograph, by Drake ers; slender legs with fore femora enlarged; a tar- (1952) and J. Polhemus & M. Polhemus (1976). sal formula of 2-2-3; the forewing membrane with In the current work, ten more new species are multiple closed cells but no anastomosing veins; and described from Mesoamerica. It is worth noting uniquely formed genitalia, with the right paramere that among these latter taxa, a number have to our greatly prolonged and elaborated in relation to the knowledge not been recollected since their origi- left. In Old World taxa the apex of the right para nal discovery by the second author and his brother mere bears appendices (for examples see figures in during their early field expeditions to Mexico in D. Polhemus & J. Polhemus 2008), whereas in all the 1960s. Several more of the new species treated New World taxa the apex of the paramere is much herein are cryptic forms that appear to live in leaf litter, and are known from very few specimens. We consider it likely that additional undescribed spe- cies will be discovered in Mesoamerican leaf litter habitats, and pitfall trapping in the moist upland forests of the region would be a promising approach to testing this hypothesis. In the context of the present monograph, we con- sider previous species concepts applied to Mesoamerican Ochteridae, particularly by Schell (1943), to have been overly broad. Similarly, some series in the col- lections of the Smithsonian Institution determined by the late Dr Carl Drake also represent multiple species under a single taxon name. Overall, our species concepts for previously described taxa as employed in the present work are often more circumscribed than those used by the above authors, and this should be taken into account when comparing this work to previous publications dealing with Mesoamerican species. Finally, it should be noted that the current paper represents the culmination of a study begun decades ago by the second author, Dr John T. Polhemus, who passed away in 2013 before it could be completed. During a series of trips to Mexico and Central America in the 1960s and 1970s, he and his brother Martin Polhemus collected many Ochterus species new to science, which over the years were partially diagnosed, figured, and described. As with many projects of this type, other priorities and adventures intervened, but in his later years Dr J. T. Polhemus returned once again to this set of interesting species Fig. 1. Ochterus viridifrons (Champion), dorsal habitus. that he had discovered early in his distinguished career Female specimen from Mexico, Guerrero, Los Terreros, in heteropteran taxonomy. Unfortunately, he did not CL 1896. Downloaded from Brill.com10/08/2021 06:09:55PM via free access
in the Mesoamerican and North American species are the dorsal color pattern (Figs 1, 16, 23, 24, 31, 38, 41, 47, 54, 63, 64, 71, 74, 75, 83, 92, 98, 104, 111, 118, 119, 126, 134, 140, 147, 154); the color- ation and pattern of striation on the frons as seen in anterior view (Figs 17, 25, 32, 42, 48, 55, 65, 72, 76, 84, 93, 99, 105, 112, 120, 127, 135, 141, 148, 155); the shape of the male right paramere (Figs 18, 26, 33, 43, 49, 57, 67, 77, 86, 95, 100, 106, 113, 121, 129, 137, 142, 149, 156); and the shape of the medial process on the male pygophore in some instances (Figs 29, 36, 45, 52, 60, 61, 70, 81, 97, 103, 109, 123, 132, 146, 153). In addition, the New World taxa also have additional character states on the pronotum that are occasionally useful for species discrimination in certain taxa; these include small hooks at the anterolateral pronotal angles (Fig.116), pointed posterolateral angles (Figs 16, 140), and small lateral tubercles arising from the anterior pro- notal lobe adjacent to the lateral flanges (Figs 2, 3). The degree of dorsal punctation (or lack thereof) and the shape of the ventral metaxiphus may also be diagnostic for certain species. In the male genitalia of Ochteridae, the pygo- phore forms a ventrally keeled, cup-like capsule (Fig. 4), the posterior end of which has a projec- tion whose apex varies in shape, from acutely pointed to apically truncate (Figs 29, 36, 45, 52, 60, 61, 70, 81, 97, 103, 109, 124, 132, 146, 153). The proctiger forms a hinged lid over the top of this capsule, and is roughly triangular-shaped in many Mesoamerican species, although it may be elongated in some taxa (Figs 5–10). Both of these structures can aid in identification at the species level, although the differences are sometimes subtle since certain basic shapes are often shared among multiple taxa. The aedeagus has a cruciform phallobase, the lateral projections of which appear to be apodemes, possibly providing points of muscle attachment for the flanking parameres (Fig. 11). The distal portion of the aedeagus consists of an elongate flagellum with two distinct points of articulation. The first and most basal of these occurs at the point where the widened base of the flagellum attaches to the phallobase. The second of these arises distally on Figs 2, 3. Pronotal structures of Ochterus species, the flagellum, at the point where it transitions from showing shape and location of pronotal tubercle, and a gradually tapering, sclerotized cylinder to a more variable degree of expansion of the anterolateral mar- slender, sinuous sclerotized filament enclosed by a gin – 2, O. explanatus n. sp., specimen from Mexico, transparent, membranous phallotheca. When at Durango, near Los Bancos; 3, O. banksi Barber, USA, rest, the basal portion of the aedeagal flagellum Texas, Columbus, CL 368. runs in the groove formed by the longitudinal ven- tral keel along the bottom of the pygophore cap- simpler in structure, with the cap and its associated sule, with the apical section, sometimes recurved appendices being absent. into a loop, often projecting through the proctiger As with Ochteridae from other parts of the world, via a basal sinus; as a result, when the proctiger is the primary characters used for species separation removed, the apical section of the flagellum may be Downloaded from Brill.com10/08/2021 06:09:55PM via free access
Proctiger Phallobase Distal Flagellum of Aedeagus Right Paramere Phallotheca
Pygophore Apex of Pygophore 4
Fig. 4. Ochterus manni Hungerford, male genitalia in lateral view, with major structures labelled. Specimen from Costa Rica, Guanacaste, 15 mi. N of Palmar Norte, CL 1302.
subject to damage. Interspecific differences in the on the structure of the right paramere, which is also form of the distal flagellum of the phallotheca exist, used as a key character in the present work. The distal particularly in regard to the length of the flagellum arm of the right paramere is frequently modified in relation to the length of the phallobase, the with lateral spurs, flanges or other types of projec- shape of the distal filament, and the degree of infla- tions, which may arise on either the inner or outer tion of the enclosing phallotheca (Figs 11–15), but margins. For the purposes of this work in regard to these have been difficult toevaluate on a compre- describing the locations of these processes, the term hensive basis in all species. Two distinct forms of ‘inner margin’ refers to the interior of the paramere’s phallotheca are present, the first type a closely U-shaped bend in lateral view, while the term ‘outer appressed envelope seen in species with a highly margin’ refers to the longer exterior side of the elongated flagellum, the second type more inflated paramere. and sack-like, generally seen in species with a shorter The ground color of most Ochterus species distal flagellum (Figs 11–15, 152). occurring in North and Central America varies The parameres flank the aedeagus to either side, from dark brown to black (Figs 1, 23, 24, 31, 38, and are oriented vertically when at rest, with the dis- 41, 47, 54, 63, 64, 71, 74, 75, 83, 92, 104, 111, tal section of the right paramere projecting slightly 118, 119, 126, 134, 140, 147, 154), although in beyond the dorsal margin of the genital capsule. some species, such as O. acutangulus and O. ova- The shapes of both parameres are interspecifically tus, the general coloration may trend toward a diagnostic, with the right paramere being large, more ruddy orange-brown (Figs 16, 98). The prominent, J-shaped, and easily examined (Figs 18, pale markings overlaying this ground color gener- 26, 33, 43, 49, 57, 67, 77, 86, 95, 100, 106, 113, ally consist of complex patches and flecks of pru- 121, 129, 137, 142, 149, 156), while the left para inose pale blue to lavender, usually symmetrically mere is small, finger-like, more translucent, and easily arranged (Figs 1, 24, 31, 38, 39, 54, 63, 64, 71, overlooked, or in some cases vestigial (Figs 21, 28, 74, 75, 83, 92, 104, 111, 118, 119, 126, 134, 35, 51, 57, 69, 79, 80, 88, 96, 102, 108, 115, 123, 140, 147, 154), and in some species may also 131, 139, 144, 151, 158). Consequently, taxonomy include dark orange ovate patches flanking the in the Neotropical Ochterus species has relied heavily claval apex, as seen in O. aeneifrons and O. barberi Downloaded from Brill.com10/08/2021 06:09:55PM via free access
Figs 5–10. Male proctiger of Ochterus species, illustrating variations in form among taxa – 5, O. schellae Drake, specimen from Mexico, Temescaltepec; 6, O. perbosci (Guérin-Menéville), specimen from Texas, Hidalgo Co., Bentson-Rio Grande State Park; 7, O. pseudorotundus n. sp., specimen from Mexico, Tamaulipas, Bocatoma; 8, O. banksi Barber, specimen from USA, Texas, Columbus, CL 368; 9, O. rotundus J. Polhemus & M. Polhemus, specimen from Mexico, Durango, near Los Bancos, CL 1017; 10, O. panamensis n. sp., specimen from Panama, Panama Prov., Serrania de Majé, CL 2779. Downloaded from Brill.com10/08/2021 06:09:55PM via free access
Basal Articulation Point Phallobase
Apex of Aedeagal Flagellum
Phallotheca
Apodemes
Distal Flagellum of Aedeagus
Distal Articulation Point Fig. 11. Ochterus chiapensis n. sp., male aedeagus, with major structures labelled. (Left) aedeagus; (right) detail of apex of distal flagellum. Specimen from Mexico, Chiapas, near Ixhuatan, CL 672.
(Figs 23, 47) or bright yellowish patches devel- Collection, the CL numbers following localities refer oped to varying degrees on the corium and clavus, to a collection locality-numbering scheme used by the as seen in certain morphs of O. banksi (Figs 40, authors to cross-reference photographs and other 41). In nearly all of the species treated there is metadata to specific collecting localities. some degree of intraspecific variation in regard to The majority of the specimens dealt with in this color pattern, such that not all specimens will paper were collected prior to the advent of modern exactly match the figures provided. global positioning systems. Many of these locali- ties, particularly for J. T. Polhemus Collection material where detailed field notes are available, Material and Methods have been retroactively georeferenced by using All measurements in the descriptions below are coordinates obtained from Web-based geospatial given in millimeters, and were made using a Wild browsers. These latitude and longitude data, as well M3Z dissecting microscope equipped with an ocular as other notations that add clarity to the locations micrometer. High resolution photographs of dorsal of particular localities in a modern context where habitus and head characters were taken using an the original label data are limited, have been added AutoMontage digital imaging system linked to a in brackets. Similarly, collection locality elevations Leica M165-C dissecting stereomicroscope, with originally taken in feet have also been converted to post-processing using Photoshop software. Line meteric values in brackets. Distances from major drawings of male genitalic structures were made towns or landmarks are given in miles because using a camera lucida attached to a Wild M3Z dis- these units are reflective of the vehicle odometer secting microscope. readings taken at the time of most of the original Synonymies provided under species are nomencla- collections. tural only, rather than comprehensive for all previous For localities where the retroactive georeference citations in the literature. For material in the Polhemus data could not be precisely constrained, the notation Downloaded from Brill.com10/08/2021 06:09:55PM via free access
Figs 12–15. Male aedeagus of Ochterus species, illustrating variations in form among taxa – 12, O. obscurus n. sp., specimen from Costa Rica, Cartago, Tapanti; 13, O. perbosci (Guérin-Menéville), specimen from Texas, Hidalgo Co., Bentson-Rio Grande State Park; 14, O. manni Hungerford, specimen from Costa Rica, Puntarenas, Boca de Barranca, CL 1306; 15, O. viridifrons (Champion), specimen from Mexico, Oaxaca, W of Tequisistlan, CL 1064.
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‘vic.’ is provided preceding the coordinates, indicat- – Anterolateral margins of pronotum not so ing the locality lies in the vicinity of the site given, strongly expanded, not extending forward of but not necessarily at the exact spot indicated. In posterior margins of the eyes, instead angling particular, because distances from towns and other backward behind eyes (Figs 1, 3, 16, 23, 24, 38, features were determined using automobile odome- 39, 40, 41, 47, 54, 63, 64, 74, 75, 83, 98, 104, ters reading to the nearest mile, these ‘vic.’ localities 111, 118, 119, 126, 134, 140, 147, 154), should be plotted as center points surrounded by although sometimes with small transverse 1.0 mi. buffers if entered into a computerized geo- ‘shoulders’ immediately behind eyes (Figs 38, graphic information system. 39, 111, 134) ������������������������������������������������ 4 The following abbreviations are used for specimen 2. Pronontum very broad, width approximately depositories: 2× that of head (Fig. 71); posterior portion of pronotum bearing deep punctations ������������ BMNH The Natural History ������������������������������������������O. explanatus n. sp. Museum, London, United – Pronotum narrower, width less than 1.8× that of Kingdom. head (Figs 31, 92); posterior portion of prono- DAPC Dan A. Polhemus Collection, tum lacking obvious punctations �������������������� 3 Englewood, Colorado, USA. 3. Frons with lower half bearing a raised, roughly EMEC Essig Museum of Ento triangular tumescence with depressed center mology, University of (Fig. 93); hemelytra strongly raised along claval California, Berkeley, suture when viewed laterally; central section of California, USA. costal margin on hemelytra with conspicuous MSUC M ichigan State University, O-shaped spot (Fig. 92); male right paramere East Lansing, Michigan, USA. with sharp spur on innner margin of distal SEMK S now Entomological Museum, arm (Figs 94, 95); Costa Rica (Fig. 73) ������������ University of Kansas, ���������������������������������������������O. obscurus n. sp. Lawrence, Kansas, USA. – Frons with lower half not raised or modifed as TAMU Texas A & M University, above (Fig. 32); hemelytra not raised along cla- College Station, Texas, USA. val suture when viewed laterally; costal margin UMMZ University of Michigan, of hemelytra lacking a prominent O-shaped Museum of Zoology, Ann spot (Fig. 31); male right paramere with Arbor, Michigan, USA. rounded flange on outer margin of distal arm UNAM Universidad Nacional (Figs 33, 34); eastern and central United States Autónoma de México, (Fig. 37) ������������������������O. americanus (Uhler) Mexico City, Mexico. 4. Pronotum lacking lateral tubercles on anterior USNM ex JTPC J. T. Polhemus Collection, lobe (Figs 23, 24, 47, 74, 75, 83) ������������������ 5 now incorporated into the – Pronotum bearing lateral tubercles on anterior United States National lobe (Figs 1, 16, 38, 39, 40, 41, 54, 63, 64, 98, Museum of Natural History, 104, 118, 119, 134, 140, 147, 126) �������������� 9 Smithsonian Institution, 5. Anterolateral angle of pronotum produced into Washington, DC, USA. a small hooked point (Fig. 116); frons brilliant USNM U nited States National metallic green with reddish overtones (Fig. 112) Museum of Natural History, �������������������������O. perbosci (Guérin-Menéville) Line Collection and Drake – Anterolateral angle of pronotum sometimes Collection, Smithsonian sharply rounded, but never forming a small Institution, Washington, hooked point as above; frons shining black, DC, USA. sometimes with coppery or greenish overtones (Figs 25, 48, 76, 84) �������������������������������������� 6 6. Costal margin of hemelytra narrowly pale along Taxonomy its entire length, this pale coloration not inter- Family Ochteridae Kirkaldy 1906 rupted by dark markings (Figs 74, 75); frons and vertex sunken well below the level of the Genus Ochterus Latreille 1802 eyes when viewed anteriorly; male right para Key to the Mesoamerican and North American mere with apex forming a small bulb bearing tiny species of Ochterus Latreille papillae (Figs 77, 78) ������O. manni Hungerford 1. Anterolateral margins of pronotum broadly – Costal margin of hemelytra pale at extreme expanded, extending forward of posterior margins base, then with alternating dark and yellowish of the eyes, then recurved back behind posterior or pruinose markings for varying degrees along eye margins (Figs 2, 31, 71, 92) �������������������� 2 remainder of its Downloadedlength (Figs from Brill.com10/08/202123, 24, 47, 83); 06:09:55PM via free access
frons and vertex not distinctly sunken below 126); dorsum bearing numerous short, pale transverse eye level when viewed anteriorly; setae (Figs 1, 38, 39, 40, 41, 63, 64, 98, 104, apex of male right paramere not bulb-like, lack- 118, 119, 134, 147, 126) ���������������������������� 13 ing papillae (Figs 26, 27, 49, 50, 86, 87) �������� 7 12. Clavus and inner corium with broad, extensive 7. Hemelytra with 4–6 distinct, evenly spaced lavender pruinose patches (Fig. 16); distal head of orange-yellow patches along each hemelytral male right paramere broadly expanded, apex margin in addition to pruinose patches (Fig. 47); blunt (Figs 18–20); Mexico, Belize and Guatemala frons black with greenish overtones (Fig. 48); (Fig. 22) ���������������������� O. acutangulus Champion apex of male right paramere forming a small, – Clavus and inner corium with only scattered, very transverse cap, not hooked (Figs 49, 50) ���������� small lavender pruinose patches (Fig. 140); distal ���������������������������������������������� O. barberi Schell head of male paramere moderately expanded, – Hemelytra with at most three distinct orange- apex more narrowed and pointed (Figs 142, 143); yellow patches along hemelytral margins, or such Costa Rica (Fig. 125) ����������������O. shepardi n. sp. spots lacking entirely, with only pruinose patches 13. Frons shining black with greenish overtones present (Figs 23, 24, 83); frons uniformly black, centrally (Fig. 42); male right paramere with without greenish overtones (Figs 25, 84); apex prominent curving flange on outer margin of of paramere slightly hooked, not forming a distal arm (Figs 43, 44); both macropterous and transverse cap (Figs 26, 27, 85–87) ���������������� 8 brachypterous forms known, with coloration of 8. Larger species, body length 4.2–4.9 mm; pro- clavus highly variable among these forms notum and inner corium bearing evident punc- (Figs 38–41) �������������������������� O. banksi Barber tations; male right paramere with distal arm – Frons metallic green, aquamarine or purple extremely elongate and slender; apical section of (Figs 65, 99, 105, 120, 127, 135, 148, 155); distal arm not expanded, lacking a lateral spur male right paramere lacking a curving flange on on inner margin (Figs 26, 27) �������������������������� outer margin of distal arm, although other types �����������������������������������O. aeneifrons Champion of spurs or projections may be present – Smaller species, body length 3.50–3.95 mm; (Figs 66–68, 100, 101, 106, 107, 121, 122, pronotum and inner corium lacking evident 128–130, 136–138, 149, 150, 156, 157); only punctations; male right paramere not as elon- macropterous forms known, with coloration of gate, apical section of distal arm slightly clavus relatively stable (Figs 1, 63, 64, 98, 104, expanded, bearing a lateral spur on the inner 118, 119, 134, 147, 126) ���������������������������� 14 margin (Figs 85–87) ����������O. mexicanus n. sp. 14. Frons blackish-green with red overtones 9. Small species, body length less than 3.75 mm (Fig. 65); pruinose coloration on hemelytra �������������������������������������������������������������������� 10 often forming a crescent-shaped mark curving – Body length greater than 3.75 mm �������������� 11 inward from base of clavus and down onto basal 10. Frons bright green (Figs 55); frons and vertex corium (although this coloration absent in cer- sunken, lying distinctly below lateral eye level; tain male specimens) (Figs 63, 64); male right male right paramere with sharp, recurved spur paramere expanded distally to form a distinct on inner margin of distal arm, apex expanded head with lateral projections on both inner and rounded (Figs 56–58) �������������������������������� and outer margins (Figs 67, 68) ���������������������� �������������������������������������������O. chiapensis n. sp. ����������������������������������������O. costaricensis n. sp. – Frons blackish-brown (Figs 99); level of frons – Frons brilliant metallic green, aquamarine or about even with, and vertex set distinctly above, purple, without red overtones (Figs 99, 105, 120, lateral eye level; male right paramere bearing a 127, 135, 148, 155); pruinose markings on basal small lateral spur on outer margin of distal arm, hemelytra more diffusely distributed, not form- apex tapering (Figs 100, 101) �������������������������� ing a crescent-shaped mark (Figs 1, 98, 104, 118, ������������������������������������������������O. ovatus n. sp. 119, 134, 147, 126); male right paramere with 11. Width of head distinctly less than half the maxi- distal portion not forming a distinct head, either mum width of the pronotum (Figs 16, 140); pos- broad and paddle-like (Fig. 121, 122, 128–130, terolateral pronotal angles sharply acute, pointed 136, 137), or broadly expanded distally with an (Figs 16, 140); dorsum lacking short, pale setae angular projection on the inner margin only (Figs 16, 140); frons shining black on upper half, (Figs 106, 107, 149, 150, 156, 157) ������������� 15 shining metallic green on lower half ���������������12 15. Pronotum and hemelytra with greenish over- – Width of head more than half the maximum tones on dark ground color (Figs 104, 147); width of the pronotum (Figs 1, 38, 39, 40, 41, male right paramere with sharp, acute spur on 63, 64, 98, 104, 118, 119, 134, 147, 126); inner margin of distal arm (Figs 106, 107, 149, posterolateral pronotal angles either acute or 150) ������������������������������������������������������������ 16 rounded, but never sharply pointed (Figs 1, 38, – Pronotum and hemelytra without greenish over- 39, 40, 41, 63, 64, 98, 104, 118, 119, 134, 147, tones (Figs 1,Downloaded 118, 119, from 126, Brill.com10/08/2021 134, 156); 06:09:55PMmale via free access
right paramere lacking a sharp, acute spur on Material examined. Belize, Cayo: 1 ♂, 11 ♀, Mountain inner margin of distal arm, if projection present Pine Ridge, Rio Maya, 5 mi. S of San Luis, water temp. 65° F., on inner margin then this projection obuse and 30 December 1973, CL 643, J. T. Polhemus (USNM ex broadly angled, not sharp and acute (Figs 121, JTPC). Guatemala, San Marcos: 1 ♀, Rio Naranjo, 450 ft, 122, 128–130, 136, 137, 156, 157) ��������������17 G. C. Champion (holotype, BMNH). Mexico, Chiapas: 16. Frons metallic aquamarine (Fig. 148); apex of 36 ♂, 21 ♀, 5 immatures, rushing mountain stream with male right paramere tapering (Figs 149, 150); pools and waterfalls, 7 mi. N of Arriaga [vic. 16°19'10"N, apex of male pygophore blunt (Fig. 153); 93°52'18", elev. ~370 m], 1 May 1964, CL 1072, J. T. & far southern Mexico (Fig. 133) ������������������������ M. S. Polhemus (USNM ex JTPC, DAPC). ���������������������������������������O. smaragdinus n. sp. – Frons bright metallic green (Fig. 105); apex Diagnosis. Body form elongate ovate; male body of male right paramere blunt and thumb-like length 4.50–5.30, maximum width (across hemely- (Figs 106, 107); apex of male pygophore pointed tra) 2.50–2.70; female body length 5.10–5.40, max- (Fig. 109); Panama (Fig. 110) ���������������������������� imum width (across hemelytra) 2.80–3.00. �������������������������������������������O. panamensis n. sp. Description. General coloration dark brown to 17. Frons metallic purple with bluish overtones orange-brown with scattered yellowish markings, (Fig. 127); body form rotund, with costal mar- hemelytra flecked with greyish to lavender pruinose gins of hemelytra strongly curved outward patches; frons black with strong metallic reddish (Fig. 126); male right paramere with obtuse, overtones centrally, metallic green laterally (Fig. 16), broadly angled projection on inner margin of eyes silvery, areas basal to eyes brown, remainder of distal arm (Figs 128–130) �������������������������������� head black except for small bluish pruinose patches ���������O. rotundus J. Polhemus & M. Polhemus at posterolateral angles; pronotum black anteriorly, – Frons metallic green (Figs 120, 135, 155); body lateral flanges dark yellow margined with dark brown form not as rotund as above, with costal mar- along their outer edges, posterior half of pronotum gins of hemelytra only weakly to moderately orange-brown, central section with 4 small transverse curving (Figs 1, 118, 119, 134, 154) male right paramere lacking a projection on inner margin of distal arm (Figs 121, 122, 136, 137), or if such a projection present then paramere thick with distal section massively expanded (Figs 155, 156) �������������������������������������������� 18 18. Metaxiphus narrowly triangular with a pro- duced posterior apex; male right paramere thick, with distal section massively expanded, apex tapering to a point (Figs 155, 156) ���������� ���������������������������������� O. viridifrons Champion – Metaxiphus broadly triangular with posterior apex not produced; male right paramere slender, with distal section elongate, slightly expanded and paddle-like, apex blunt and broadly rounded (Figs 121, 122, 136–138) �������������� 19 19. Large species, body length 5.20–6.30 mm; male right paramere with low, elongate flange along outer margin of distal arm (Figs 137, 138); western Mexico to Guatelmala and El Salvador (Fig. 110) ������������������������������O. schellae Drake – Smaller species, body length 4.90–5.80; male right paramere with a slight inflection on inner margin of distal arm, lacking a low flange on outer margin (Figs 121, 122); northeastern Mexico (Fig. 125) �������O. pseudorotundus n. sp.
Ochterus acutangulus (Champion) Figs 16–22 Fig. 16. Ochterus acutangulus (Champion), dorsal hab- Pelogonus acutangulus Champion 1901: 346. itus. Male specimen from Mexico, Chiapas, 7 mi. N of Ochterus acutangulus: Schell 1943: 38. Arriaga, CL 1072. Downloaded from Brill.com10/08/2021 06:09:55PM via free access
Fig. 17. Ochterus acutangulus (Champion), head, anterior view. Female specimen from Belize, Cayo, Rio Maya, CL 643.
bluish pruinose patches along transverse midline behind calli; mesoscutum and scutellum black, with bluish pruinose patches at basal angles, as a small roughly triangular patch medially on anterior mar- gin, and to either side of posterior apex, posterior apical angle sordid white; hemelytra dark brown to rusty brown, costal margins broadly orange-brown, marked with greyish to lavender pruinose patches on Figs 18–21. Ochterus acutangulus (Champion), male each hemelytron consisting of a large lavender pru- genitalic structures – 18, right paramere; 19, detail of inose patch centrally on clavus, 4 irregular lavender terminal section of right paramere; 20, terminal section pruinose patches scattered on the basal corium, 4 of right paramere in alternate view; 21, left paramere. larger irregular greyish pruinose patches scattered on Specimen from Mexico, Chiapas, 7 mi. N of Arriaga, posterior corium, and 4 irregular lavender pruinose CL 1072. patches on the embolium inward of margin, fore- wing membrane black to dark brown, mottled with greyish pruinosity, veins rusty brown (Fig. 16). Frons Comparative notes. Ochterus acutangulus is most sim- with coarse transverse striations, with broad, shallow ilar to O. shepardi from Costa Rica, with both taxa channels parallel to inner eye margins, these channels possessing sharply angular posterolateral angles on bearing transverse striations along their lengths, the pronotum, and lacking short, pale setae on the upper portion of frons with an incipient raised carina dorsum, character states that easily separate them along longitudinal midline (Fig. 17). Dorsal surface from other members of the Mesoamerican ochterid lacking obvious setae (Fig. 16); posterior pronotum, assemblage (Figs 16, 140). Ochterus acutangulus may clavus and corium bearing numerous coarse, black be separated from O. shepardi by the more massive punctations (Fig. 16). distal structure on the stout male right paramere, Male right paramere short and stout, distal arm which comes to a broad, roughly triangular apex short and thick, apical section expanded, with pro- (compare Figs 18–20 and 142, 143), and by the jecting angle on outer margin, tip acutely rounded more extensive lavender pruinose coloration on (Figs 18–20); left paramere small, expanded basally the hemelytra (compare Figs 16 and 140). For with stepped shoulder, distal arm slender, apex further comparative notes see the discussion under rounded (Fig. 21); pygophore weakly produced, O. shepardi. apex truncate; aedeagus of moderate length, distal The second author examined the holotype section not recurved, tip acuminate, lying within female of this species, held in the Natural History inflated phallotheca. Museum, London,Downloaded UK. The from specimen Brill.com10/08/2021 is in 06:09:55PMgood via free access
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Fig. 22. Distribution of Ochterus acutangulus in Mesoamerica. condition, but slightly teneral. It bears the following O. acutangulus and provided an illustration of the labels: “Type (round label)/Rio Naranjo, 450 ft., distinctively acute and pointed lateral pronotal Champion/B. C. A. Rhyn. II, Pelogonus viridifrons, angles that characterize this species. We now add male, Ch./Sp. Figured.” new records for O. acutangulus from Mexico Ecological notes. Ochterus acutangulus is a hygropet- (Chiapas) as well as Belize. Based on current intrepre- ric species associated with rock faces adjacent to tations, this species is confined to uplands at the base waterfalls, or large rocks in streams. As such, it is of the Yucatan Peninsula in Belize, Guatemala, and not frequently collected. The field notes of the sec- extreme southeastern Mexico (Fig. 22). Previous list- ond author describe the ecological preferences of ing of this species from Cuba could not be recon- this species at CL 643 on the Rio Maya in Belize: firmed by Naranjoet al. (2010), and its presence on “This is a big stream in a big valley. It can be waded that island is considered doubtful. in places. I got a series of a fine ochterid from coarse, bristly vegetation on rocks in the swift stream part, Ochterus aeneifrons aeneifrons (Champion) where I also took Mesoveloidea…The ochterids Figs 23–30 were taken by holding a net downstream from the rocks and splashing or rubbing the vegetation. Pelogonus aeneifrons Champion 1901: 345. The bugs were quick to fly, and tried to fly from Ochterus aenifrons [sic]: Schell 1943: 43. the net.” Ochterus aeneifrons: Froeschner 1981: 61. Distribution. This species was originally described from a single male specimen taken at Rio Naranjo, Material examined. Belize, Stann Creek Dist.: 4 ♂, 2 ♀, Guatemala (Champion, 1901). In her subsequent stream with waterfalls at Mile 31 on Hummingbird Hwy., monograph, Schell (1943) misinterpreted this spe- 31 mi. W of Stann Creek, 27 December 1973, CL 638, cies, and provided records of O. acutangulus from the J. T. Polhemus (USNM ex JTPC); 1 ♂, Blue Hole, 15 mi. state of Mexico (Temescaltepc) based on misidenti- SE of Belmopan, on wet bedrock walls next to stream fied specimens. The Mexican species involved was entering cave [17°08'48"N, 88°40'32"W, elev. ~140 m], subsequently described as O. schellae by Drake 28 December 1973, CL 632, J. T. Polhemus (USNM ex (1952), who clarified the species concept for JTPC). Costa Rica, Cartago: 1 ♀, La Suiza, 17 June 1967, Downloaded from Brill.com10/08/2021 06:09:55PM via free access
Fig. 23. Ochterus aeneifrons aeneifrons (Champion), Fig. 24. Ochterus aeneifrons aeneifrons (Champion), dorsal habitus. Female specimen from Mexico, Oaxaca, dorsal habitus. Female specimen from Mexico, Guer- El Carmen, CL 1068. rero, Los Terreros, CL 1896.
Flint and Ortiz (USNM). Puntarenas: 1 ♂, San Vito, 2 km Soyalo, Rio Blanco, on road to Villahermosa ~ 14 mi. N of S of Finca La Cruces, 4000 ft., 8°42'N, 83°0'W, 14 March Hwy. 190, small swift mountain river with backwater pools 1969, D. C. Rentz (USNM); 8 ♂, 4 ♀, Golfito, 4 August [vic. 16°53'27"N, 92°55'30"W, elev. ~1600 m], 4 May 1957, A. Menke (USNM); 2 ♂, Puntarenas Prov., San 1964, CL 1086, J. T. & M. S. Polhemus (USNM ex Vito, 2 km S, Finca Las Cruces, 4000 ft., 14 March 1969, JTPC); 1 ♂, 7 mi. N of Arriaga, rushing mountain stream 08°42’N, 83°0’W, D. C. Rentz (USNM). El Salvador, with waterfalls and mossy rocks [vic. 16°19'10"N, Chalatenango: 1 ♂, 3 ♀, roadside spring with small pools 93°52'18", elev. ~370 m], 1 May 1964, CL 1072, J. T. & nr. La Majadita, 21 December 1969, CL 1255, M. S. Polhemus (USNM ex JTPC); 1 ♂, 1 mi. SW of J.. T Polhemus (USNM ex JTPC). Guatemala, Alta Ixhuatan, mountain stream with waterfalls and pools [vic. Verapaz: 1 ♂, 1 ♀, 57.6 km N of El Rancho on new Coban 17°16'39"N, 93°01'14"W, elev. ~590 m], 5 May 1964, CL highway, 1463 m, 30 May 1973, Erwin & Hevel Central 1098, J. T. & M. S. Polhemus (USNM ex JTPC); 1 ♂, 2 ♀, American Expedition (USNM). Izabal: 1 ♂, 1 ♀, swift 19 mi. N of Arriaga, small stream with sandy bed [vic. stream E of Quirigua, 11 Janauary 1970, CL 1317, J. T. 16°26'59"N, 93°49'27"W, elev. ~640 m], 1 May 1964, CL Polhemus (USNM). Mexico, Chiapas: 2 ♂, 1 ♀, 12 mi. N 1076, J. T. & M. S. Polhemus (USNM ex JTPC); 12 ♂, of Bochil, tiny shaded mountain stream with pools [vic. 4 ♀, roadside seep and small stream 2 mi. N of Tapiula [vic. 17°04'24"N, 92°51'51"W, elev. ~ 1650 m], 4 May 1964, 17°16'07"N, 93°01'25"W, ~ 610 m elev.], 5 May 1964, CL 1091, J. T. & M. S. Polhemus (USNM ex JTPC); 1 ♂, CL 1096, J. T. & M. S. Polhemus (USNM ex JTPC); 1 ♂, 7 mi. S of Pichucalco, small smoothly flowing river with 1 ♀, San Quintin, Chiquita Miramar, 700 ft.. 16°24'N, vertical banks [Rio Ixtacomitan, 17°25'21"N, 93°06'13"W, 91°20'W, 5–20 February 1966, G. E. Ball and D. R. elev. 180 m], 5 May 1964, CL 1103, J. T. & M. S. Whitehead (USNM). Colima: 1 ♂, 1 ♀, Mt. Colima, SE Polhemus (USNM ex JTPC); 2 ♂, 1 ♀, 7 mi. N of Santa slope, pine zone, December 1948, E. S. Ross (USNM ex Fe, rushing river with large pools and mossy banks JTPC). Guerrero: 1 ♂, Puente Terreros at Km 21 on [17°25'56"N, 93°02'05"W, elev. ~150 m], 5 May 1964, Mexico highway 134, NE of Ixtapa, 395 m [1290 ft., vic. CL 1102, J. T. & M. S. Polhemus (USNM ex JTPC); 1 ♂, 17°47'44"N, 101°27"W], 29 January 1985, CL 1896, Downloaded from Brill.com10/08/2021 06:09:55PM via free access
Fig. 25. Ochterus aeneifrons aeneifrons (Champion), head, anterior view. Female specimen from Mexico, Oaxaca, El Carmen, CL 1068.
J. T. Polhemus (USNM ex JTPC). Jalisco: 1 ♀, Puente El Naranjo, 13.1 mi. SW of Pihuamo, Route 110, 1500 ft., 19 January 1966, G. E. Ball and D. R. Whitehead (USNM). Mexico: 2 ♂, 1 ♀, Tejupilco Dist., Temascaltepec, 1340 m, June–July 1933, H. E. Hinton and R. L. Usinger (USNM ex JTPC). Nayarit: 1 ♀, 32 mi. S of Acaponeta, 24 November 1948, H. B. Leech (USNM ex JTPC). Nuevo Leon: 5 ♂, 1 ♀, 7.5 mi. S of Monterrey, 18 April 1974, J. C. Schaffner (TAMU). Oaxaca: 5 ♂, El Carmen, sandy bottomed tropical river with rock outcrops, 30 April 1964, CL 1068, J. T. Polhemus (USNM ex JTPC); 2 ♀, 8 mi. NE of Oaxaca, small rocky mountain stream [vic. 17°06'21"N, Figs 26–29. Ochterus aeneifrons aeneifrons (Cham- 96°38'32"W, elev. ~1705 m], 29 April 1964, CL 1060, pion), male genitalic structures – 26, right paramere; J. T. & M. S. Polhemus (USNM ex JTPC); 3 ♂, 3 ♀, rocky 27, detail of terminal section of right paramere; 28, left stream on Hwy. 190 10 mi. W of Rio Hondo bridge [vic. paramere; 29, apex of pygophore. Specimen from USA, 16°30'24"N, 94°54'27"W, elev. ~825 m], 30 April 1964, Illinois, Lake Co., Dead River. CL 1064, J. T. & M. S. Polhemus (USNM ex JTPC); 1 ♀, 3 mi. NW of Tapanatepec, 31 December 1955, 27–28 May 1966, G. E. Ball and D. R. Whitehead J. C. Schaffner (USNM ex JTPC). Puebla: 2 ♂, 2 ♀, 20 mi. (USNM); 1 ♂, Est. Biol. Los Tuxtlas, vic. Laguna SE of Acatlan (10 mi. NW of Puebla), stream and small Escondida, 16 September 1987, Malaise trap, L. Carroll seeps in deep ravine [vic. 17°59'54"N, 97°53'28"W, elev. (TAMU). Nicaragua: 1 ♀, 8 mi. N of Esteli, small stream ~1600 m], 28 April 1964, CL 1053, J. T. & M. S. Polhemus in bed of volcanic rocks, [vic 13°11'02"N, 86°22'10"W], (USNM ex JTPC). Queretaro: 1 ♀, Escanelilla, Rio Jalpan, 2500 ft. [790 m], 23 December 1969, CL 1262, J. T. 3900 ft., 16 November 1965, G. E. Ball and D. R. Polhemus (USNM ex JTPC). Panama, Coclé: 3 ♂, swift Whitehead (USNM). San Luis Potosi: 2 ♂, 1 ♀, Rio Santa stream with waterfalls in rainforest below El Valle [vic. Maria, S of Santa Maria del Rio [vic. 21°47'40"N, 8°36'06"N, 80°07'48"W, elev. 585 m], 3 January 1970, 100°42'40"W, elev. 1710 m], 7 May 1964, CL 1110, CL 1299, J. T. Polhemus (USNM ex JTPC); 2 ♂, 3 ♀, sand J.. T & M. S. Polhemus (USNM ex JTPC). Tamaulipas: 2 bottomed river at San Carlos, 1 mi. N of El Valle, 4 January ♂, 3 ♀, Bocatoma, 7 km SSE of Gomez Farias, 19–23 May 1970, CL 1300, J. T. Polhemus (USNM ex JTPC). 1979, E. G. Riley (USNM ex JTPC); 2 ♂, 1 ♀, 5 mi. SSE Canal Zone: 3 ♂, 1 ♀, Galeta Island, 9°23'N, 79°52'W, of Gomez Farias, 19–20 July 1970, Murray, Phelps, Hart 15 January 1974, D. Engleman (USNM); 2 ♂, Pipeline & Schaffner (TAMU). Veracruz: 1 ♂, 2 ♀, 17 mi. N of Road, 30 June 1974, D. Engleman (USNM); 1 ♂, Barro Acayucan, 11 June 1965, Burke, Meyer and Schaffner Colorado Island, 10–17 May 1964, W. D. and S. S. (USNM ex JTPC); 1 ♀, Fortin de las Flores, 2900 ft., Duckworth (USNM). Downloaded from Brill.com10/08/2021 06:09:55PM via free access
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Fig. 30. Distribution of Ochterus aeneifrons aeneifrons in Mesoamerica.
Extralimital material examined. Trinidad & Tobago: bluish to lavender flecks on each hemelytron consist- 1 ♂, Trinidad, 26 June 1902, Chipman (USNM ex JTPC). ing of 5 relatively evenly spaced patches along cos- COLOMBIA: 1 ♂, Rio Frio, 15 March 1925, no collector tal margin, an elongate patch at the inner basal data (USNM ex JTPC). margin of the clavus, another small patch centrally on clavus, and 5 irregular patches on the inner corium, Diagnosis. Body form elongate ovate; male body extreme posterior inner corium with a prominent length 4.20–4.50, maximum width (across hemely- pale grey pruinose patch immediately posterior to tra) 2.40–2.45; female body length 4.40–4.90, the dark yellow mark adjacent to claval apex, costal maximum width (across hemelytra) 2.30–2.70. margin dark yellow on basal third, membrane black, General coloration dark brown to reddish brown irregularly maculate with bluish to grey pruinose with scattered yellow-orange markings on prono- patches (Figs 23, 24). Frons with fine transverse stria- tum and hemelytra, dorsal suface flecked with bluish, tions, these striations becoming somewhat coarser lavender and pale grey pruinose patches (Figs 23, 24); centrally, moderately wide channels present along head including frons and vertex shining black with inner eye margins, these channels bearing a few trans- vague dark green overtones, eyes silvery; prono- verse striations centrally (Fig. 25). Basal head, prono- tum black, lateral flanges yellow, posterior pronotal tum, scutellum and hemelytra set with scattered margin yellowish-orange medially, this coloration in short, stout, pale setae (Figs 23, 24); posterior prono- some cases broadly expanded to cover entire posterior tum, clavus and corium coarsely punctate, these pronotal margin, posterior pronotum with scattered punctations bluish to lavender pruinose in con- bluish to lavender pruinose patches, consisting of strast to surr ounding dark brown ground color 4 transverse patches centrally near transverse midline, (Figs 23, 24). a small patch well inside each posterolateral angle, Male genitalia with right paramere U-shaped, dis- and a small patch posteromedially; scutellum black, tal arm very long, slender, rod-like (Fig. 26), apex with bluish pruinose patches at anterolateral and rounded, tip slightly hooked (Fig. 27); left paramere posterior angles, and a large pale grey patch centrally small, broadly Z-shaped (Fig. 28); pygophore with on anterior margin; hemelytra dark rusty reddish medial process produced, apex blunt, rounded brown, marked with a pair (1+1) of dark yellow spots (Fig. 29); aedeagus very elongate, distally recurved, on inner corium behind posterior claval apex, and apex lying within inflated phallotheca. Downloaded from Brill.com10/08/2021 06:09:55PM via free access
Discussion. Ochterus aeneifrons is among the most widespread and commonly encountered Ochterus species in Mesoamerica. The very slender and elon- gate distal arm on the male right paramere, ending in a slightly hooked tip, is diagnostic for this taxon (Figs 26, 27). In addition, the pronotum is relatively narrow, with rounded posterolateral angles, giving the species a distinctively compact and ovate appear- ance. The dorsal coloration exhibits significant regional variation in O. aeneifrons, particularly in regard to the extent of yellowish coloration on the posterior pronotal margin, which usually comprises a broad patch centrally but may cover the entire width of the posterior pronotum in specimens from Costa Rica and Panama; and in the degree of devel- opment of a pair of dark yellowish spots on the inner angles of the corium posterior to the claval apex (Figs 23, 24). Among all these variants, how- ever, a general brownish to yellowish-brown ground color prevails, which is somewhat lighter than that of other species occurring in Mesoamerica. Distribution. Ochterus aeneifrons was originally described from a series of specimens taken at widely separated localities in Mexico (Teapa, Tabasco), Guatemala (San Geronimo; Guatemala City), Panama (Tolé, Peña Blanca, San Feliz), Grenada, and St. Vincent. Schell (1943) listed specimens from Mexico, Guatemala, Honduras, Costa Rica, Panama, Colombia, Ecuador, and the island of Fig. 31. Ochterus americanus (Uhler), dorsal habitus. Trinidad, and Moreira et al. (2011) reported the Female specimen from USA, Missouri, Tucker Prairie. species from Brazil. In the region under study, this taxon is widespread throughout Mexico and Central America, and ranges into the southern- most Caribbean islands. In addition to the syntype- series records from Tabasco, we now provide further Mexican records for the states of Chiapas, Colima, Jalisco, Mexico, Nayarit, Oaxaca, Puebla, Queretaro, San Luis Potosi, Tamaulipas and Veracruz, as well as the first country records for Belize and Nicaragua (Fig. 30). The subspecies O. aeneifrons surinamensis was described from Suriname (Nieser, 1975), and we have seen further specimens from Venezuela. Nieser & Chen (1992) also recorded O. aeneifrons from Ecuador, but did not indicate which subspecies their records pertained to. All records listed in the current work apply to the nominate subspecies. Ecological notes. This species occurs on wet rocks along swift upland streams.
Ochterus americanus (Uhler) Figs 31–37 Fig. 32. Ochterus americanus (Uhler), head, anterior view. Female specimen from USA, Missouri, Tucker Prairie. Pelogonus americanus Uhler, 1876: 335. Pelogonus americatus [sic]: Snow, 1906: 180. Material examined. United States, Florida: 1 ♂, Highlands Ochterus americanus: Torre-Bueno & Brimley, 1907: 433. Co., Lake Placid, Archbold Biological Station, Price Tract, Ochterus americanum [sic]: Barber, 1913: 215. 1 April 1990, Malaise trap, W. W. Worth (USNM); 1 ♀, Downloaded from Brill.com10/08/2021 06:09:55PM via free access
1 ♂, Worcester, Assateague Island, taken at high water mark, 21 September 1988, C. L. and S. L. Staines (USNM). Minnesota: 1 ♂, Mille Lacs Co., 14 May 1938, H. E. Milleron (MSUC). Missouri: 9 ♂, 5 ♀, Callaway Co., Tucker Prairie [vic. 38°57'00"N, 91°59'52"W, elev. 270 m], 11 September 1981, CL 56, C. N McKinnon (USNM ex JTPC). Nebraska: 1 ♂, Cherry Co., Niobrara River at wildlife refuge entrance [42°54'00"N, 100°28'54"W, elev. 720 m], 3.5 mi. NW of Valentine, 11 June 1963, D. R. Smith (USNM ex JTPC); 1 ♀, Valentine, 11 June 1950, Hicks, Slater and Lafoon (USNM). New Jersey: 1 ♀, Lakehurst, 2 June 1918, H. G. Barber collection (USNM). North Carolina: 1 ♂, Wilmington Beach, 6 April 1914, H. G. Barber collection (USNM).
Diagnosis. Body form elongate ovate; male body length 4.20–4.60, maximum width (across hemely- tra) 2.60–2.80; female body length 4.85–5.20, max- imum width (across hemelytra) 2.85–3.10. General coloration black with limited yellowish white markings on anterolateral pronotum and cos- tal margin, hemelytra flecked with dark bluish grey pruinose patches (Fig. 31); frons and labrum shining black; pronotum black, lateral flanges with pale yel- lowish white patches, pronotal disk mottled with dark bluish grey pruinose patches; scutellum black, with a pair (1+1) of pale bluish patches at lateral and posterior angles, broadly at anterior margin flanking longitudinal midline, and as small paired dots cen- trally to either side of midline; hemelytra black, each hemelytron with 5 small, evenly spaced dark yellow marks along length of costal margin, corium and cla- vus mottled with dark bluish grey pruinose patches, membrane black, irregularly maculate with dark blu- ish grey pruinose patches (Fig. 31). Frons with coarse transverse striations, broad channels present along inner margins of eyes, these channels bearing a few transverse striations, upper half of frons with a raised carina along longitudinal midline (Fig. 32). Basal head, pronotum, scutellum and hemelytra set with scattered short, stout, pale setae (Fig. 31); pronotum, Figs 33–36. Ochterus americanus (Uhler), male genital- clavus and corium lacking obvious punctations ic structures – 33, right paramere; 34, detail of terminal section of right paramere; 35, left paramere; 36, apex (Fig. 31). of pygophore. Specimen from USA, Illinois, Lake Co., Male genitalia with right paramere U-shaped, dis- Dead River. tal arm moderately long, stout, bearing a small fin on outer margin, apex tapering, tip rounded (Figs 33, 34); left paramere small, bearing angled lateral projection Alachua Co., 23 December 1973, coll. Matta (USNM). (Fig. 35); pygophore with posterior margin elongate, Georgia: 1 ♀, Echols Co., 11 April 1970, R. J. Beshear roughly triangular, apex acutely pointed (Fig. 36); (USNM). Illinois: 1 ♂, Lake Co., Illinois Beach State Park, aedeagus with distal flagellum moderately short, 4× flooded sedges N of bridge over Dead River, 25 June 1970, length of phallobase, terminal section acuminate, Brigham and Sanderson (USNM ex JTPC). Kansas: 1 ♂, 1 contained within expanded phallotheca. ♀, Cheyenne Co., spring-fed gulch approx. 3.6 mi. N of St. Comparative notes. Ochterus americanus is a relatively Francis [vic. 39°48'26"N, 101°50'37"W, elev. 1025 m], 9 small, dark-colored species with bright white patches July 1987, CL 1919, J. T. Polhemus (USNM ex JTCP). on the anterolateral flanges of the pronotum Louisiana: 5 ♂, 7 ♀, Beauregard Parish, Dry Creek, 24 (Fig. 31), which are a useful spot character for recog- June 1969, H. C. Chapman (USNM ex JTPC). Maryland: nizing this species in the field. The male right Downloaded from Brill.com10/08/2021 06:09:55PM via free access
Ochterus americanus
Fig. 37. Distribution of Ochterus americanus (Uhler) in North America.
paramere is distinctive, bearing a small, rounded fin records from the Appalachian Mountains and the on the outer margin of the distal arm (Figs 33, 34). Ozark Plateau, a pattern also seen in O. banksi, In terms of pronotal morphology and coloration, which occupies a largely congruent range in eastern this species possesses similarities to O. explanatus North America (compare Figs 37 and 46). In the (Figs 2, 71) and O. obscurus (Fig. 92), two other spe- western portion of its range, populations of O. amer- cies that are also associated with moist soil and leaf icanus are strongly correlated with spring outflows litter rather than stream banks, sand bars or vertical from the Ogallala Aquifer at widely scattered points rock seeps. The elongate, roughly triangular and api- across the Great Plains, including Valentine, Nebraska cally acute posterior apex of the pygophore is also and St. Francis, Kansas. diagnostic, and similar to the character state seen in Ecological notes. Ochterus americanus occurs in sparse O. banksi (Fig. 45). vegetation adjacent to spring outflows, and is not an Distribution. This species is widespread in the central inhabitat of open sand bars. Uhler (1884) notes that and eastern United States, with previously confirmed the species “…lives among the grass and weeds on records from the states of Illinois, Kansas, Louisiana, the margins of brooks and ponds…” Massachusetts, Maryland, Mississippi, Missouri, North Carolina, New Jersey, New York, Nebraska, Ochterus banksi Barber Pennsylvania, Texas and Viriginia (D. Polhemus & Figs 3, 8, 38–46 J. Polhemus 1988), to which we now add Georgia, Minnesota and North Carolina (Fig. 37). Despite its Ochterus banksi Barber, 1913: 214. wide geographic distribution, this species is in fact Ochterus flaviclavus Barber, 1913: 215. n. syn. rather uncommon in major collections, with Schell (1943) having been able to examine only two speci- Material examined. United States, Connecticut: 2 ♂, 1 ♀, mens while preparing her monograph of Western Tolland Co., Storrs, 17 June 1970, Wharton (TAMU). Hemisphere Ochteridae. Although our current base Florida: 2 ♂, 1 ♀, Big Pine Key, margins of dry drainage of material examined is larger, it is still surprisingly ditches, 4 February 1968, CL 404, J. T. Polhemus (USNM limited given the broad geographic range involved ex JTPC); 1 ♀, 6 mi. S of Edgewater, margins of drainage within an intensively surveyed country such as the ditches, 6 February 1968, CL 414, J. T. Polhemus (USNM United States. There is a conspicuous absence of ex JTPC); 1 ♀, 6 mi. N of Flamingo, drainage ditch Downloaded from Brill.com10/08/2021 06:09:55PM via free access
Fig. 38. Ochterus banksi Barber, dorsal habitus. F emale Fig. 39. Ochterus banksi Barber, dorsal habitus. Female specimen of macropterous form from USA, Texas, specimen of brachypterous form from USA, Texas, Ingleside. Ingleside.
margins, 4 February 1968, CL 409, J. T. Polhemus (USNM 1966, CL 369, J. T. Polhemus (USNM); 2 ♂, 2 ♀, Brazos ex JTPC); 1 ♂, DeLand, 24–26 March 1939, F. E. Lutz Co., Mintner Springs, 23–25 July 1973, R. Burton (USNM ex JTPC); 1 ♂, Ormond, ex. Barber Collection (USNM ex JTPC); 2 ♂, 3 ♀, San Patricio Co., Ingleside (holotype of O. flaviclavus, USNM); 1 ♀, Orlando, [vic. 27°50'36"N, 97°11'13"W, elev. 1 m], 25–26 May 14 March 1952, H. C. Chapman (USNM); 1 ♂, Lakeland, 1973, R. G. Burton (USNM ex JTPC); 3 ♀, same locality 21 March 1950, J. E. Burgess (USNM). Kansas: 8 ♂, 22 ♀, as preceding except 10 August 1974 (USNM ex JTPC). Meade Co., Meade County State Park [vic. 37°10'08"N, Virginia: 1 ♀, Vienna, 20 July 1913, H. G. Barber collec- 100°26'19"W, elev. 745 m], 6 July 1985, C. McKinnon tion (USNM); 1 ♀, Louisa Co., 4 mi. S of Cuckoo, 19–29 (USNM ex JTPC). Louisiana: 1 ♂, Burns, 21 June 1948, June 1988, in Malaise trap, J. Kloke and D. R. Smith H. W. Crowder (USNM ex JTPC). Maryland: 1 ♂, 3 ♀, (USNM). West Virginia: 1 ♀, Putnam Co., 8 June 1990, Beltsville, 19 June 1915, W. L. McAtee (USNM); 1 ♂, 1 ♀, Kovarik & Wells (TAMU). Odenton, 20 June 1915, W. L. McAtee (USNM); 2 ♂, 4 ♀, Prince Georges County, Beltsville, BARC-East, 6 June Diagnosis. Body form elongate ovate (macropterous 1986, T. J. Henry (USNM). Michigan: 4 ♂, 4 ♀, Clinton form) to broadly ovate (brachypterous form); male Co., Rose Lake W. E. S., 14–20 June 1966, D. Bixler and body length 4.10–4.60, maximum width (across T. Hlavac (MSUC). Mississippi: 4 ♂, Ocean Springs, hemelytra) 2.25–2.50; female body length 4.35–5.20, 23 May 1950, J. T. Polhemus (USNM). Ohio: 2 ♀, maximum width (across hemelytra) 2.50–2.90. Washington Co., Belpre, Veto Lake State Park [vic. General coloration of typical form reddish brown 39°20'34"N, 81°39'22"W, elev. 205 m], 19 May 1977, with limited yellowish white markings on anterolat- C. N. McKinnon (USNM ex JTPC). Pennsylvania: 2 ♂, eral pronotum, hemelytra flecked with bluish grey 5 ♀, Centre Co., Scotia Barrens, at pond’s edge, 26 June pruinose patches (Figs 38, 39), but coloration of 1980, P. H. Adler (USNM). Texas: 1 ♂, Colorado Co., clavus variable, sometimes with yellowish patch cen- Columbus, 25 May 1966, J. T. Polhemus (USNM); 1 ♂, trally (Fig. 40), or entirely yellowish (‘flaviclavus’ 1 ♀, Chambers Co., 3 mi. N of Seabrook, 5 December form – Fig. 41). Frons shining black, labrum dark Downloaded from Brill.com10/08/2021 06:09:55PM via free access
Fig. 40. Ochterus banksi Barber, dorsal habitus. Fig. 41. Ochterus banksi Barber, dorsal habitus. Female specimen of ‘intermediate’ form from USA, Female specimen of ‘flaviclavus’ form from USA, Texas, Texas, Ingleside. Ingleside. brown; pronotum reddish brown, lateral flanges with and corium lacking coarse punctations (Figs 38–41); pale yellowish white patches at anterolateral angles, pronotum bearing lateral tubercles (Fig. 3). Brachyp pronotal disk mottled with dark bluish grey pruinose terous forms similar to above but with forewing patches, calli blackish in some individuals; scutellum membrane variably reduced (Figs 39–41), bearing dark brown to black, with pairs (1+1) of pale bluish 0–7 evident cells, with or without bluish grey pru- patches at basolateral angles, flanking midline at inose maculations. center of anterior margin, and surrounding small, Male genitalia with right paramere U-shaped, dis- triangular preapical projections on posterolateral tal arm long, slender, bearing prominent fin on outer margins ahead of posterior apex; hemelytra reddish margin, apex weakly crescentic (Figs 43, 44); left brown, each hemelytron with 4 small, evenly spaced paramere small, moderately stout, weakly S-shaped; bluish grey pruinose marks along length of costal pygophore with medial process produced, apex margin between base of embolium to immediately roughly triangular (Fig. 45); aedeagus with distal fla- behind posterior embolar margin, clavus and corium gellum elongate, over 3.5x the length of phallobase, mottled with numerous small bluish grey pruinose distal phallotheca narrow; proctiger with distal sec- patches, membrane reddish brown, bearing 7 evi- tion elongate, basal lobes laterally expanded (Fig. 8). dent cells, irregularly maculate with bluish grey pru- Comparative notes. Ochterus banksi is a widespread inose patches (Fig. 38). Frons with coarse transverse species in the eastern half of the United States, and striations, channels present along inner margins of exhibits a significant degree of polymorphism in eyes, these channels bearing a few transverse stria- regard to forewing development and dorsal color tions, upper half of frons with distinct raised carina pattern throughout its range. Both fully macropter- along longitudinal midline (Fig. 42). Basal head, ous and brachypterous forms occur in this species pronotum, scutellum and hemelytra set with scat- (Figs 38–41); in some of the latter the forewings tered short, stiff, golden setae arising from small, have the membrane greatly reduced, appearing cole- slightly depressed foveae (Fig. 38); pronotum, clavus opteriform and giving the impression of a shorter Downloaded from Brill.com10/08/2021 06:09:55PM via free access
Fig. 42. Ochterus banksi Barber, head, anterior view. Female specimen of from USA, Texas, Ingleside. and broader body form, the hind wings are not developed, and the hemelytral and posterior prono- tal punctation is more pronounced. Although the macropterous form is more common within the material examined for this study, both morphs may be present at the same locality, as is the case with the series from Ocean Springs, Mississippi listed above, which consists of two macropterous ♂ and two brachypterous ♂, all taken together. The venation of the forewing membrane is also highly variable across the material examined, with certain brachypterous specimens exhibiting no evident venation at all, and others having up to 7 clearly defined closed cells (Figs 38–41). The coloration of O. banksi is also highly vari- able. The most typical form has a dark reddish brown ground color with scattered lavender mark- Figs 43–45. Ochterus banksi Barber, male genitalic ings (Figs 38, 39). In the Pennsylvania series listed structures – 43, right paramere; 44, detail of terminal above the specimens have strongly developed laven- section of right paramere; 45, apex of pygophore. Speci- der markings, in contrast to the Maryland series men from USA, Texas, Colorado Co., Columbus. where these markings are more obscure or in some individuals nearly lacking. In addition, specimens from the southeastern United States may possess circular creamy white patch subtending the apex pale, creamy white markings to varying degrees on of the clavus and adjacent inner portions of each the central hemelytra (Figs 40, 41). Notable in this corium (Fig. 40). Another example of this same regard is a series from Ingleside, Texas which con- color morph is also at hand from DeLand, Florida. tains individuals that intergrade from the com- Given the syntopic occurrence of specimens con- pletely dark ‘normal’ color form of O. banksi to forming to the description of both O. banksi having the clavus almost entirely pale creamy yellow and O. flaviclavus, and the fact that the geo- (Fig. 41), the latter form having been described as a graphic ranges of both species completely overlap, separate species, O. flaviclavus, by Barber (1913). we have concluded that O. flaviclavus is simply an The latter is known from occasional rare specimens extreme (although visually distinctive) color form taken in Florida, Loiuisiana, and Texas. In addition, of O. banksi. the Ingleside series contains certain intermediate We have also examined the holotype of O. flavi- color forms consisting of specimens with a roughly clavus, a male held in the Smithsonian Institution Downloaded from Brill.com10/08/2021 06:09:55PM via free access
Ochterus banksi
Fig. 46. Distribution of Ochterus banksi Barber in North America. and bearing the following labels: “ORMOND, lobase, and has a closely appressed phallotheca on its FLA./Type (on red paper)/Type No. 64726 U. S. N. terminal portion, similar in form to that seen in M./Ochterus flaviclavus Barber Type (red margin on O. barberi from the southwestern United States. label)”. The specimen is mounted on a point and is Distribution. Originally described from Virginia in generally good condition, with the forewing (Glencarlyn, Nathan Banks), this species is wide- membranes torn, and both hind legs plus the left spread in the eastern United States, with confirmed foreleg missing. The genitalia have been dissected records from the states of Florida, Indiana, Kansas, and are in a vial below the specimen. A comparison Louisiana, Massachusetts, Mississippi, Missouri, of the male genitalia to those of O. banksi validates New Jersey, New York, South Carolina, Texas and the observation of Schell (1943) that there are no Viriginia (Polhemus & Polhemus 1988). To this we structural differences between these two species in now add records from Connecticut, Ohio, Michigan regard to this character system, with the only differ- and West Virginia (Fig. 46). As is the case with the ences instead lying in dorsal color pattern. Based on previously discussed O. americanus, this species this, and given the presence of intergrading color seems to be absent from the Appalachian Mountains morphs as noted above, we have synonymized and Ozark Plateau, although it occurs in lowland O. flaviclavus under O. banksi. areas surrounding these uplifts. The male right paramere of O. banksi is similar to Ecological notes. This is a species of horizontal sub- that of O. perbosci, with both having a large, arcuate, strates, and is not associated with rocks or vertical projecting fin on the outer margin of the distal arm, seeping rheocrenes. although this fin is larger in O. banksi (compare Figs 43, 44 to 113, 114). This paramere morphology Ochterus barberi Schell is not seen elsewhere in the Mesoamerican ochterid Figs 47–53 biota, although the male right paramere of O. ameri- canus incipiently approaches this character state Ochterus barberi Schell, 1943: 41. (Figs 33, 34). Ochterus banksi also shares with O. americanus an angular and distally acute posterior Material examined. Mexico, Sonora: 1 ♂, Guaymas [can- lobe on the pygophore (compare Figs 36 and 45). yon with palms ~8 mi. W of San Carlos Bay], 8 June 1966, The distal flagellum of the aedeagus in O. banski is CL 1207, J. T. Polhemus (USNM ex JTPC). United elongate, being over 3.5 times the length of the phal- States, Arizona: 1 ♂, 2 ♀, springs in Aravaipa Canyon,
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Fig. 48. Ochterus barberi Schell, head, anterior view. Female specimen from USA, California, Bard.
gravel bars, 31 May 1972, CL 551, J. T. & M. S. Polhemus (USNM ex JTPC). California: 1 ♂, Bard [vic. 32°46'31"N, 114°33'53"W, elev. ~40 m], 14 November 1951, R. L. Usinger (USNM ex JTPC); 1 ♀, Lake Houghtelin, nr. Bard, 13 November 1951, R. L. Usinger (USNM). Fig. 47. Ochterus barberi Schell, dorsal habitus. Female Colorado: 1 ♀, Eagle Co., 1.5 mi. N of Eagle, along Eagle specimen from USA, California, Bard. River [vic. 39°39'56"N, 106°48'14"W, elev. ~2020 m], 7 April 1989, B. Kondratieff (USNM ex JTPC). New Mexico: 2 ♂, seeping bank along [West Fork] Gila River nr. 8 October 1964, CL 315, J. T. Polhemus (USNM Gila [Cliff Dwellings] National Monument [33°13'33"N, ex JTPC); 1 ♀, Indian Hot Springs [32°59'56"N, 108°15'05"W, elev. ~1695 m], 17 April 1965, CL 323, 109°54'90"W, elev. 850 m], CL 314, 9 October 1964 J. T. Polhemus (USNM ex JTPC); 1 ♀, Sherman, Mimbres J.. T Polhemus (USNM); 2 ♀, small dam and seeps 5 mi. N Hot Springs [32°44'55"N, 107°50'26"W, elev. 1735 m], of Castle Hot Springs, 7 October 1964, CL 312, 17 April 1965, CL 322, J. T. Polhemus (USNM ex J.. T Polhemus (USNM ex JTPC); 1 ♀, Castle Hot Springs JTPC); 2 ♀, Las Cruces [vic. 32°18'24"N, 106°49'33"W, [33°58'56"N, 112°21'45"W, elev. 600 m], 4 August 1941, elev. ~1185 m], 12 June 1950, R. H. Beamer (SEMK). R. H. Beamer (USNM ex JTPC); 1 ♂, springs and seeps along fish hatchery road above White River, 15 April 1965, Diagnosis. Body form elongate ovate; male body CL 317, J. T. Polhemus (USNM ex JTPC); 1 ♂, Slaughter length 4.60–4.80, maximum width (across hemely- Ranch, E of Douglas, margins of spring-fed lake by ranch tra) 2.20–2.60; female body length 4.85–5.10, house [31°20'12"N, 109°16'45"W, elev. 1155 m], 1 June maximum width (across hemelytra) 2.60–2.80. 1975, CL 702, J. T. Polhemus (USNM ex JTPC); 4 ♂, 2 General coloration rusty brown with head, prono- ♀, Mojave Co., springs above Beaverdam Wash at I-15 tum and scutellum blackish, pronotum and hemely- bridge [36°53'38"N, 113°55'51"W], 1800 ft. [550 m], tra with scattered yellow-orange markings, dorsal water temp. 77° F., 21 September 1997, CL 3024, J. T. & body overall flecked with bluish, lavender or grey D. A. Polhemus (USNM ex JTPC); 2 ♂, 2 ♀, Coconino pruinose patches (Fig. 47); frons black with greenish Co., seeps on bedrock in middle section of Buck Farm overtones, eyes golden with overtones of green, red- Canyon, trib. to Colorado River in Marble Canyon, Mile dish, and honey yellow, remainder of head black 41 below Lee’s Ferry, [36°23'52"N, 111°53'80"W, elev. (Fig. 48); pronotum black, lateral flanges dark yellow 930 m], 28 May 1972, CL 546, J. T. and M. S. Polhemus with dark brown patches centrally on outer margins, (USNM ex JTPC); 3 ♂, 1 ♀, Grand Canyon, Stone Creek posterior margin of pronotum yellowish orange in at confluence with Colorado River, Mile 132 from Lee’s central section, pronotal disk with scattered bluish or Ferry [vic. 36°20'46"N, 112°27'15"W, elev. ~610 m], on grey pruinose patches, consisting of a pair (1+1) of
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scutellum black, with bluish pruinose patches at anterolateral and posterior angles, a pair (1+1) of patches at anterior margin to either side of longi- tudinal midline, and another pair of similar patches immediately posterior to the preceding; hemelytra dark brown, clavus rusty brown, inner corium marked with a pair (1+1) of dark yellow spots behind posterior claval apex, each hemelytron with addi- tional grey to lavender pruinose patches consisting of a narrow patch at the inner basal margin of the clavus, 6 diffuse and irregular patches on the inner corium, and a prominent grey pruinose patch on posterior corium adjacent to central inner margin of membrane, each costal margin bearing 5 dark yellow markings, with the central 3 of these markings being more distinct, and the costal margin basal to the most anterior of these marks remaining nar- rowly yellow, membrane dark brown, irregularly maculate with bluish to lavender pruinose patches, veins orange-brown (Fig. 47). Frons with moderately coarse transverse striations, with ventrally angling striae on ventral half, and dorsally angling striae on upper half, separated by a set of transverse stria- tions centrally, broad channels present to inner eye margins (Fig. 48). Basal head, pronotum, scutel- lum and hemelytra set with a few scattered short, stout, pale setae (Fig. 47); vertex not sunken below eyes when viewed head on; posterior prono- tum, clavus and corium lacking obvious coarse punctations (Fig. 47), pronotum lacking lateral tubercles. Male genitalia with right paramere U-shaped, distal arm elongate, a small, angular projection pres- ent on inner margin of bend at two-thirds distance to apex (Fig. 49), tip forming a small, transverse head (Fig. 50); left paramere small, elongate, with small angle centrally (Fig. 51); pygophore with medial process moderately produced, apex rounded with a slight lateral indentation (Fig. 52); aedeagus with distal flagellum extremely long, over 5× the length of the phallobase, distal phallotheca narrow. Discussion. Ochterus barberi is found as scattered populations associated with springs and rivers in the deserts of the southwestern United States. The struc- ture of the male left paramere, with a projecting angle on the inner margin of the bend and a small, transverse cap at the apex is diagnostic for this species Figs 49–52. Ochterus barberi Schell, male genitalic (Figs 49, 50), although similar in some respects to structures – 49, right paramere; 50, detail of terminal the shape seen in O. mexicanus (Figs 85–87). In addi- section of right paramere; 51, left paramere; 52, apex tion, the distal flagellum of the aedeagus is extremely of pygophore. Specimen from Mexico, Sonora, near long, being nearly 6 times the length of the phallo- Guaymas, CL 1207. base, and coils back upon itself in a large loop when at rest within the pygophore. Externally, the antero- patches anteriorly behind eyes, another pair (1+1) of lateral flanges of the pronotum with broad yellow elongate, sinuate patches centrally around calli, small margins surrounding a small interior dark patch patches posterolaterally set well inside each postero- (Fig. 47), combined with a large, roughly cir cular, lateral angle, and a small patch posteromedially; greyish pruinose patch on the inner posterior corium Downloaded from Brill.com10/08/2021 06:09:55PM via free access
Ochterus barberi
Fig. 53. Distribution of Ochterus barberi Schell in North America. behind the claval apex (Fig. 47), are also characteris- the Mexican state of Veracruz, but these specimens tic of this species. have proven on closer examination to represent We have examined the holotype, a male specimen females of O. aeneifrons, and therefore are not held in the Smithsonian Institution and bearing included on the distribution map presented (Fig. 53). the following labels: “Col Canyon, 13–7. Ar/H. Ecological notes. This species inhabits damp, horizon- S. Barber Collector (on white paper)/Ochterus tal, littoral substrates, such as river sand bars, at barberi, D. Viets (on red paper)/Type No. 55607 scattered sites throughout the northwestern Mexico U. S. N. M. (on red paper)”. The specimen is in good and the southwestern United States. It is a wary condition and retains all its appendages except the insect that is quick to take flight, and therefore left rear tibia and tarsi. The male genitalia have been probably more widespread than current records dissected and are in a vial below the specimen. indicate. Distribution. Ochterus barberi was originally described from a holotype taken in Arizona at “Colorado Ochterus chiapensis n. sp. Canyon” (ie., the Grand Canyon of the Colorado Figs 11, 54–62 River), with an allotype from Hot Springs, Arizona (probably Castle Hot Springs). This species occurs in Type material. Holotype, male: Mexico, Chiapas, seeping the Colorado River Basin of the southwestern United roadside rock faces near Ixhuatan [2.5 mi. S of Ixhuatan, States and adjacent desert regions, with confirmed vic. 17°16'04"N, 93°01'26"W, elev. ~640 m], 6 January records from the states of Arizona, California 1973, CL 672, J. T. Polhemus (USNM ex JTPC). Paratypes: and New Mexico (Polhemus & Polhemus 1988), as Mexico, Chiapas: 12 macropterous ♂, 5 macropterous ♀, well as the Mexican state of Sonora. To this we can same data as holotype (USNM ex JTPC, BPBM); 1 ♂, add a new record from western Colorado, which sug- swift mountain river in rain forest near Santa Fe [vic. gests this species may occur further upstream along 17°20'59"N, 93°00'22", elev. ~290 m], 5 May 1964, CL the tributary rivers of the Colorado River basin than 1101, J. T. & M. S. Polhemus (USNM ex JTPC). is currently realized, at least during the summer months, and is almost certainly present in southeast- Description ern Utah. In the original description of this species, Male. General form elongate ovate, body length 3.30– Schell (1943) also listed paratypes from Orizaba, in 3.35, maximum width (across hemelytra) 1.80–2.00. Downloaded from Brill.com10/08/2021 06:09:55PM via free access
Fig. 55. Ochterus chiapensis n. sp., head, anterior view. Female specimen from Mexico, Chiapas, near Ixhuatan, CL 672.
Fig. 54. Ochterus chiapensis n. sp., dorsal habitus. Structural characters. Head — Length (along midline Female specimen from Mexico, Chiapas, near Ixhua- as measured from directly above)/width (across eyes) tan, CL 672. = 0.30/1.20, angled downward at greater than 45° when viewed laterally; frons with coarse trans- verse striations, upper half of frons with prominent Color. General coloration dull dark blackish brown longitudinal medial carina, inner margins of eyes flecked with small pale bluish pruinose patches bordered by narrow channels, these channels bearing (Fig. 54); head with vertex dull dark blackish brown numerous transverse striations (Fig. 55); eyes large, with small bluish pruinose patches at each postero- protruding, projecting posterolaterally slightly lateral angle, eyes reddish brown, frons brilliant beyond vertex, upper margins not rising above level metallic dark green, tylus bronzy reddish metallic, of vertex when viewed anteriorly; antennae with seg- clypeus and labrum dark yellow (Fig. 55), rostrum ments I and II short, globose, segments III and IV pale yellowish brown, antennae pale brown; prono- slender, filiform, lengths of segments I–IV = 0.05, tum dull dark blackish brown, narrowly dark yellow 0.10, 0.25, 0.30; rostrum long, length = 2.00, along posterior margin, lateral flanges yellow nar- exceeding hind coxae and extending onto base of rowly margined with black, paired small pale, bluish abdominal venter. pruinose patches present at anterolateral angles, Pronotum — Length (midline)/width = 0.80/1.80, along lateral margins adjacent to lateral flanges, and anterior lobe lacking lateral tubercles, entire surface centrally to either side of midline; mesoscutum and set with scattered short, recumbent, golden setae, scutellum dull dark blackish brown, trending to red- bearing numerous small shallow punctations; ante- dish brown centrally, posterior apex with small, pale rior collar weakly demarcated; anterolateral margins bluish pruinose patch; hemelytra dull dark reddish explanate, well demarcated from disk; calli barely brown with metallic red overtones, costal margins tumescent; posterior margin multisinuate, posteri- dark orange to yellow, bearing scattered pale bluish orly concave centrally above base of scutellum. pruinose markings as follows: large, irregular patch Scutellum — Triangular, length/width = 0.50/0.90, on basal half of clavus, 6–8 scattered flecks on corium bearing scattered short, recumbent, golden setae, and wing membrane, 6 moderate-sized, roughly cir- mesoscutum transversely raised, remainder of scutel- cular to ovate patches along each wing margin lum only weakly tumescent. (Fig. 54); thoracic venter dull black, acetabulae pale Hemelytra — Long, attaining tip of abdomen, brown, abdominal venter dark brown; legs with fem- with corium, clavus and embolium well defined, ora straw yellow, femoral apices plus all tibiae and surfaces set with scattered short, recumbent, golden tarsi brown. setae; clavus, adjacent inner corium and embolar
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length, upright, slender, pale setae; hind femur straight, not bowed downward when viewed later- ally; all tibiae bearing numerous short, bristly golden setae, these setae becoming more numerous distally, interspersed with scattered short, stout, golden spines; claws golden, gently curving, arolia long, slender, length subequal to that of claws. Lengths of leg segments as follows: fore femur/tibia/tarsal 1/ tarsal 2 = 1.00/1.00/0.05/0.15; middle femur/tibia/ tarsal 1/tarsal 2 = 1.00/1.00/0.05/0.20; hind femur/ tibia/tarsal 1/tarsal 2/tarsal 3 = 1.10/1.60/0.05/ 0.15/0.15. Male genitalia — Right paramere U-shaped, mod- erately short, distal section strongly expanded, with a backwardly directed angular spur on inner margin, apex broadly rounded (Figs 56–58); left paramere slender, with small rounded lateral projection near midpoint (Fig. 59); pygophore with caudal tip curved, finger-like, and with apex rounded in lateral view (Fig. 60), broad and blunt in ventral view (Fig. 61); aedeagus moderately elongate, not distally recurved, apex lying within inflated phallotheca (Fig. 11). Female. Similar to male in general structure and col- oration, body length 3.80–3.90, maximum width (across hemelytra) 2.00–2.25. Similar to male in gen- eral structure and coloration, and body proportions. Etymology. The name ‘chiapensis’ refers to the Mexican state of Chiapas, from which this species was described. Comparative notes. Ochterus chiapensis occurs on seeps in southern Mexico, and is the smallest Ochterus species occurring in Mesoamerica, with a Figs 56–61. Ochterus chiapensis n. sp., male genitalic body length ranging from 3.30–3.90 mm. The male structures – 56, right paramere, inner view of apex; 57, right paramere; 58, detail of terminal section of right right paramere is distinctive, with the distal section paramere; 59, left paramere; 60, apex of pygophore, expanded to a broadly rounded apex, and a project- lateral view; 61, apex of pygophore, ventral view. Speci- ing angular spur present on the inner margin of the men from Mexico, Chiapas, Santa Fe, CL 1101. distal arm (Figs 56–58). The finger-like projection on the posterior margin of the pygophore in lateral view (Fig. 60) is also diagnostic. margin bearing shallow punctations; anterolateral Distribution: Mexico (Chiapas) (Fig. 62). embolar margin only very narrowly explanate; length Ecological notes: The field notes of the second of clavus along outside margin 1.30; membrane author provide the following information on the venation weakly defined, with 3 evident closed cells ecological preferences of this species (under the basally. manuscript name Ochterus chiapasana) at CL 672 Ventral surface — Ventral thorax smooth and dull, near Ixhuatan: “Near Santa Fe, Chiapas, where one bearing scattered extremely short, pale, recumbent reaches the Rio Teapa after dropping down from setae; scattered tiny, dark punctations present on pro-, Tapilula, there are many tiny falls and seeps on the meso-, and metepisterna; abdominal venter covered vertical rock faces along the road. I stopped at one with very short, fine, appressed pale setae, this setal to look for Ochterus chiapisana (n. spp.) that we covering interrupted by ovate glabrous patches sur- found as one specimen in 1964. Sure enough, it was rounding spiracles laterally on paratergites. on the wet algae-moss covered rock faces, where the Legs — All segments covered by short, fine, face was quite wet but not running…The ochterids recumbent pale setae; fore and middle trochanters preferred the overhangs and dark places, and would and basal portions of fore and middle femora with settle themselves into depressions in the soft surface very long, semi-erect, slender, pale setae ventrally, so as to look like a spot of dirt. They were not remainder of ventral margins bearing moderate quick to move, and even slower to jump. I collected
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Ochterus chiapensis
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Ochterus costaricensis 5 5 SCALE 0 100 200 300 400 500 600 MILES 0 200 400 600 800 KILOMETERS CONIC PROJECTION
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Fig. 62. Distribution of Ochterus species in Mesoamerica. ● O. chiapensis n. sp.; ■︎ O. costaricensis n. sp.
them with a vial and had a chore getting them to del Toro: 40 ♂, 33 ♀, seeping roadside bedrock face ~1 km jump into the bottle. Weather was warm, cloudy off road to Chiriqui Grande on Continental Divide Road and humid.” [vic. 8°46'49"N, 82°12'26"W], ~1100 m, 13 January 1993, CL 2821, J. T. Polhemus & A. R. Gillogly (USNM, Ochterus costaricensis n. sp. BPBM, DAPC). Canal Zone: 1 ♂, 1 ♀, Canal Zone, Barro Figs 63–70 Colorado Island, 6–8 February 1939, C. J. Drake (USNM). Chiriqui: 4 ♂, 10 ♀, Balneareo, 13 January 1993, CL Type material. Holotype, male: Costa Rica, Puntarenas, 2819, J. T. Polhemus (DAPC); 4 ♂, 2 ♀, 3 km N of David, roadside waterfall 15 mi. N of Palmar Norte [8°58'14"N, Rio Majagua at Balneario Majagua, spring seepage at 83°18'15"W, elev. 70 m], 6 January 1970, CL 1302, waterfall, 31 May 1983, P. J. Spanger, R. A. Faitoute and J.. T Polhemus (USNM ex JTPC). Paratypes: Costa Rica, W. E Steiner (USNM). Coclé: 2 ♂, 1 ♀, swift stream with Puntarenas: 2 ♂, 6 ♀, same data as holotype (USNM ex waterfalls in rainforest below El Valle [vic. 8°36'06"N, JTPC, BPBM); 3 ♂, 3 ♀, Puntarenas, 11 km E of Esparta, 80°07'48"W, elev. ~585 m], 3 January 1970, CL 1299, 9 June 1963, C. L. Hogue (USNM ex JTPC); 3 ♀, road- J. T. Polhemus (USNM). side waterfall 14 mi. N of Palmar Norte [8°56'34"N, 83°20'36"W, elev. 60 m], 28 December 1969, CL 1280, J. T. Polhemus (USNM ex JTPC); 1 ♂, 1 ♀, Puntarenas, Description Monteverde area, 1400–1700 m, 6–14 June 1973, Erwin Male. General form elongate ovate, body length & Hevel Central American Expedition 1973 (USNM). 4.50–4.65, maximum width (across hemelytra) Cartago: 3 ♂, 4 ♀, Rio Catarat, Hwy. 2, 228 m, 16 June 2.40–2.55. 1973, Erwin & Hevel Central American Expedition Color. General coloration dark orange-brown flecked (USNM). San José: 3 ♂, 4 ♀, San Jose, 30 June 1957, with small pale bluish pruinose patches, scutellum and Truxal and Menke (USNM). El Salvador, La Libertad: anterior pronotum dark brown, head black (Figs 63, 2 ♂, 5 ♀, El Chorro [Los Chorros], W of Santa Tecla, from 64); head with vertex dull dark blackish brown with seeping rock faces next to waterfalls in shaded canyon [vic. small bluish pruinose patches at each posterolateral 13°41'43"N, 89°19'10"W, elev. 790 m], 20 December angle, eyes silvery, frons brilliant metallic dark green 1969, CL 1253, J. T. Polhemus (USNM). Panama, Bocas above, metallic bronzy reddish below, clypeus dark Downloaded from Brill.com10/08/2021 06:09:55PM via free access
Fig. 63. Ochterus costaricensis n. sp., dorsal habitus. Fig. 64. Ochterus costaricensis n. sp., dorsal habitus. Female specimen from Costa Rica, Guanacaste, 15 km Male specimen showing dark color form, from Costa N of Palmar Norte, CL 1302. Rica, Guanacaste, 15 km N of Palmar Norte, CL 1302. yellow, labrum dark brown (Fig. 65), rostrum pale in the closed cells, extensively mottled with bluish yellowish brown, antennae with segments I and II pruinose posterior to the closed cells (Figs 63, 64); pale yellowish brown, segments III and IV medium thoracic venter dull black extensively overlain by brown. Pronotum with anterior lobe dull dark black- blue-grey pruinosity, acetabulae pale yellowish ish brown, posterior lobe dull dark orange-brown, brown, abdominal venter medium brown; legs with narrowly lighter orange-brown along central section femora medium brown dorsally, straw yellow ven- of posterior margin, lateral flanges yellow narrowly trally, all tibiae and tarsi yellowish brown, terminal margined with black, paired small, pale, bluish pru- tarsal segments darker brown. inose patches present at inner angles of lateral flanges Structural characters. Head — Length (along midline adjoining lateral tubercles. Scutellum including as measured from directly above)/width (across mesoscutum dull dark blackish brown, extreme apex eyes) = 0.50/1.50, angled downward at greater than shining pale yellow, a short bluish pruinose stripe 45° when viewed laterally; frons with coarse stria- present along longitudinal midline on mesoscutum. tions on lower section, upper portion of frons with a Hemelytra dull dark orange-brown with metallic pair (1+1) of roughly circular depressions to either red overtones; embolium and costal margins orange- side of a raised longitudinal medial carina, upper brown, embolium bearing 3 evenly spaced small frons above depressions and vertex lacking striations, bluish pruinose patches along its length; clavus with inner margins of eyes bordered by narrow channels, 2 small bluish pruinose patches on basal third, con- these channels bearing numerous transverse stria- sisting of a narrow stripe basally followed by a smaller tions (Fig. 65); eyes large, protruding, projecting bluish pruinose patch; basal corium with a single posterolaterally very slightly beyond vertex, upper small bluish pruinose patch along inner embolar margins equal with vertex when viewed anteriorly; suture, distal corium with 2 larger, irregular bluish antennae with segments I and II short, globose, pruinose patches at base of wing membrane; wing segments III and IV slender, filiform, lengths of membrane with a few small flecks of bluish pruinose segments I–IV = 0.12, 0.12, 0.30, 0.30; rostrum Downloaded from Brill.com10/08/2021 06:09:55PM via free access
by ovate glabrous patches surrounding spiracles lat- erally on paratergites. Legs — All segments covered by short, fine, recumbent pale setae; fore and middle trochanters and basal portions of fore and middle femora with very long, semi-erect, slender, pale setae ventrally, remainder of ventral margins bearing moderate length, upright, slender, pale setae; hind femur bowed slightly downward when viewed laterally; all tibiae bearing numerous short, bristly golden setae, these setae becoming more numerous distally, inter- spersed with scattered short, stout, golden spines; hind tibia bearing fringe of very long, slender, golden setae along posterior margin on basal half; claws golden, gently curving, arolia long, slender, length subequal to length of claws. Lengths of leg segments as follows: fore femur/tibia/tarsal 1/tarsal 2 = 1.50/1.15/0.06/0.25; middle femur/tibia/tarsal 1/ tarsal 2 = 1.40/1.35/0.10/0.30; hind femur/tibia/tar- Fig. 65. Ochterus costaricensis n. sp., head, anterior sal 1/tarsal 2/tarsal 3 = 1.60/2.30/0.10/0.20/ 0.25. view. Female specimen from Costa Rica, Guanacaste, Male genitalia — Right paramere U-shaped, distal 15 km N of Palmar Norte, CL 1302. arm relatively long, slender, apical section expanded, with a projecting angular spur on inner margin and rounded projection on outer margin, apex broadly long, length = 2.60, exceeding hind coxae and rounded (Figs 66–68); left paramere small, slender, extending onto base of abdominal venter. finger-like, slightly curved (Fig. 69); pygophore with Pronotum — Length (midline)/width = 0.95/2.50, medial process moderately produced, apex rounded anterior lobe bearing lateral tubercles, entire surface (Fig. 70); aedeagus with distal flagellum relatively set with scattered very short, obscure, recumbent, short, tip slender and acuminate, contained within silvery setae, bearing numerous small, prominent, an elongate phallotheca. shallow punctations; anterior collar weakly demar- Female. Similar to male in general structure and col- cated; anterolateral margins explanate, narrow, well oration but slightly larger in over size, body length demarcated from disk; calli weakly tumescent; poste- 4.90–5.30, maximum width (across hemelytra) rior margin multisinuate, posteriorly concave cen- 2.60–3.00. trally above base of scutellum. Etymology. The name ‘costaricensis’ refers to the Scutellum — Large, triangular, length/width = country of Costa Rica, the type locality. 1.00/1.40, bearing scattered very short, obscure, Comparative notes. Ochterus costaricensis is a moder- recumbent, silvery setae, mesoscutum not raised; ately large, macropterous species occurring on seeps basal portion of posterior portion of scutellum in Central America. It can be easily recognized by the behind mesoscutum transversely raised; scutellar form of the male right paramere, which forms a dis- apex acute, flattened, shining. tinct head terminally, the base of which projects out Hemelytra — Long, attaining tip of abdomen, from both the inner and outer margins of the distal with corium, clavus and embolium well defined, sur- arm (Figs 66–68). The frons is blackish-green, with faces set with scattered very short, obscure, recum- reddish overtones on the lower half (Fig. 65), in con- bent, silvery setae; clavus, adjacent inner corium and trast to other regional congenors in lower Central embolium bearing numerous shallow punctations; America, such as O. viridifrons and O. panamensis, in anterolateral embolar margin only very narrowly which the frons is a more brilliant metallic green. explanate; length of clavus along outside margin The hemelytral markings in females often form a 1.80; membrane venation weakly defined, with 8 rough inward-curving crescent extending backward evident closed cells basally. from the base of the clavus onto the inner corium, Ventral surface — Ventral thorax smooth and dull, which can be a useful spot character (Fig. 63). By bearing scattered extremely short, pale, recumbent contrast, the pruinose markings on the male fore- setae; scattered tiny, dark punctations present on wings are more variable, in some cases being greatly pro-, meso-, and metepisterna, these punctations reduced in comparison to the female (Fig. 64). absent on ventral surfaces of pronotal flanges; Distribution. Costa Rica, Panama and El Salvador abdominal venter covered with very short, fine, (Fig. 62). Given the current distribution, this species appressed pale setae, this setal covering interrupted is likely to occur in Honduras and Nicaragua as well. Downloaded from Brill.com10/08/2021 06:09:55PM via free access
Fig. 71. Ochterus explanatus n. sp., dorsal habitus. Figs 66–70. Ochterus costaricensis n. sp., male genitalic Female specimen from Mexico, Durango, 7 mi. W of structures – 66, right paramere, inner view of apex; 67, Los Bancos. right paramere; 68, detail of terminal section of right paramere; 69, male left paramere; 70, apex of pygo- phore. Specimen from Costa Rica, San Jose. Color. General coloration dark reddish brown flecked with small pale bluish pruinose patches, scutellum Ecological notes. This species inhabits rheocrenes, and central pronotum dark brown, head black, and is often found in association with waterfalls, anterolateral spots on pronotum pale yellow (Fig. 71); where it occurs on vertical rock faces in the periph- head with vertex dull black with each posterolateral eral splash zone. angle bluish grey, eyes silvery-red; frons shining black, clypeus, tylus and rostrum orange-brown (Fig. 72), Ochterus explanatus n. sp. antennae with segment I pale whitish yellow, segment Figs 2, 71–73 II whitish yellow clouded with pale brown, segments III and IV pale brown; pronotum dull dark brown, Type material. Holotype, female: Mexico, Durango lateral flanges medium brown, with large, transversely [incorrectly indicated as Sonora], 7 mi. W of Los ovate pale yellow patches centrally, paired small pale, Bancos [vic. 23°38'54"N, 105°47'56"W, elev. ~2320 bluish pruinose patches present to either side of m], 26 April 1974, M. S. Polhemus (USNM ex longitudinal midline on anterior margin behind eyes, JTPC). Paratypes: Mexico, Oaxaca: 1 ♀, 10.9 km N adjacent to inner margins of lateral flanges at bases of of Candalaria [Candelaria Loxicha], 991 m, ex. tubercles, and centrally behind calli; mesoscutum and cloud forest, 12 July 1987, R. Anderson (USNM ex scutellum dull black, small pale blue pruinose patches JTPC); 1 ♀, Portillo del Rayo, 5 km S of Jalatengo present centrally on anterior margin and at posterior [vic. 15°58'49"N, 96°31'02' W, elev. 1510 m], apex; hemelytra dull dark brown, costal margin with 21 October 1982, H. Brailovsky (USNM ex JTPC). 4 similarly sized, evenly spaced dark yellow patches along its length, clavus with pale bluish pruinose Description patch basally and additional small, irregular bluish Female. General form broadly ovate, body length pruinose patches posteriorly, inner corium with 4.65, maximum width (across hemelytra) 2.80. narrow band of dense pale bluish pruinose mottling Downloaded from Brill.com10/08/2021 06:09:55PM via free access
Pronotum — Length (midline)/width = 1.10/2.55, anterior lobe with small lateral tubercles, entire sur- face set with scattered, moderately short, recumbent, golden setae, posterior lobe bearing numerous very small, shallow, indistinct punctations; anterior collar moderately demarcated; anterolateral margins highly explanate, broadly curving, well demarcated from disk, lateral angles obtuse, pronotal tubercles present as in Fig. 2; calli weakly tumescent; posterior margin multisinuate, posteriorly concave centrally above base of scutellum. Scutellum — Triangular, length/width = 0.80/1.30, moderately tumescent, bearing scattered short, recum- bent, golden setae, mesoscutum largely hidden under posterior pronotum, separated from remainder of scutellum by prominent transverse furr ow, posterior portion of scutellum with raised transverse tumes- cence on basal section, and a pair (1+1) of raised tumescences on posterior section separated by a shal- Fig. 72. Ochterus explanatus n. sp., head, anterior view. low depression along the longitudinal midline. Female specimen from Mexico, Durango, 7 mi. W of Hemelytra — Macropterous, exceeding tip of Los Bancos. abdomen, clavus and embolium well defined, surface set with scattered moderately short, recumbent, golden setae; basal clavus, adjacent inner corium, adjacent to claval suture, lateral and distal margins of and basal embolium bearing shallow punctations; corium with scattered pale bluish pruinose patches, anterolateral embolar margin thinned and folded small J-shaped bluish pruinose patch present on cen- slightly upward; length of clavus along outside mar- tral corium; wing membrane extensively mottled gin 1.80; membrane well developed, with 6 evident with small, irregular pale bluish pruinose patches closed cells basally. (Fig. 71); thoracic venter dull brownish black cen- Ventral surface — Ventral thorax smooth and dull, trally, orange-brown laterally, small dark puncta- bearing scattered extremely short, pale, recumbent tions, overlain with faint bluish pruinosity, setae; scattered tiny, dark punctations present on pro-, undersides of pronotal flanges each with pale yellow meso-, and metepisterna, and pro- and mesoacetabu- patch, ventral embolium orange-brown, pro-, meso- lae; abdominal venter covered with very short, fine, and metasternum dark blackish brown, acetabulae appressed pale setae, this setal covering interrupted and posterior sections of pro- and metapleurae by ovate glabrous patches surrounding spiracles lat- orange-brown; abdominal venter including genital erally on paratergites. segments orange brown; legs yellowish brown, all Legs — All segments covered by short, fine, femoral apices narrowly brown. recumbent gold setae; fore and middle trochanters Structural characters. Head — Length (along midline and basal portions of fore and middle femora with as measured from directly above)/width (across eyes) slightly longer pale setae ventrally; hind femur bowed = 0.50/1.35, angled downward at greater than very slightly downward when viewed laterally; fore 45° when viewed laterally; frons and clypeus with and middle tibiae bearing numerous short, bristly coarse transverse striations, upper half of frons with golden setae, these setae becoming more numerous prominent longitudinal medial carina, inner margins distally; all tibiae bearing scattered short, stout, of eyes bordered by narrow channels with strongly golden spines, length of these spines on middle and raised inner margins forming prominent carinae par- hind tibiae subequal to the widths of these segments; allel to inner margins of eyes, these channels bearing claws golden, gently curving, arolia long, equal to or numerous transverse striations (Fig. 72); eyes large, exceeding length of claws. Lengths of leg segments protruding, projecting posterolaterally slightly beyond as follows: fore femur/tibia/tarsal 1/tarsal 2 = posterior margin of vertex, upper margins not rising 1.40/1.15/0.05/0.15; middle femur/tibia/tarsal 1/ above level of ocelli when viewed anteriorly; anten- tarsal 2 = 1.30/1.25/0.10/0.15; hind femur/tibia/tar- nae with segments I and II short, globose, segments sal 1/tarsal 2/tarsal 3 = 1.45/2.00/0.05/0.20/0.15. III and IV slender, filiform, lengths of segments Male. Unknown. I–IV = 0.10, 0.15, 0.30, 0.40; rostrum long, length Comparative notes. Ochterus explanatus is a moder- = 2.30, exceeding hind coxae and extending onto ately small and robust species, which may be recog- base of abdominal venter. nized by the highly explanate and broadly curving Downloaded from Brill.com10/08/2021 06:09:55PM via free access
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Ochterus obscurus 5 5 SCALE 0 100 200 300 400 500 600 MILES 0 200 400 600 800 KILOMETERS CONIC PROJECTION
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Fig. 73. Distribution of Ochterus species in Mesoamerica. ● O. explanatus n. sp.; ■︎ O. obscurus n. sp. anterolateral flanges of the pronotum (Fig. 2), Ochterus manni Hungerford which are well demarcated from disk and bear large, Figs 4, 14, 74–82 pale yellow patches (Fig. 71). It may be further rec- ognized by the strongly sculptured frons, with a Ochterus manni Hungerford 1927: 189. prominent vertical medial carina on the upper half, flanked by a pair (one to either side laterally) of Material examined. Costa Rica, Guanacaste: 1 ♂, sharply raised vertical carinae that parallel the inner Guanacaste, 16 July 1957, D. R. Lauck (USNM ex JTPC). margins of the eyes for the entire length of the Puntarenas: 1 ♂, Rincon de Osa, Playa Blanca road on Rio frons, setting off deep, transversely striate channels Riyita, 9 March 1969, D. C. Rentz (USNM ex JTPC); 7 along the inner margins of the eyes (Fig. 72). The ♂, 3 ♀, roadside waterfall, 15 mi. N of Palmar Norte male is currently unknown, precluding ev aluation of [8°58'14"N, 83°18'15"W, elev. 70 m], 6 January 1970, CL the character states in the male genitalia, but the 1302, J. T. Polhemus (USNM ex JTPC); 1 ♂, 1 ♀, Boca de above character states should permit its future Barranca, 7 January 1970, CL 1306, J. T. Polhemus recognition. (USNM ex JTPC). Panama, Bocas del Toro: 2 ♂, crashing The locality label on the holotype specimen from stream in culvert, 3.2 km N of Continental Divide along Los Bancos, Mexico contains an error, since this seep road to Chiriqui Grande, 800 m, 13 January 1993, CL locality actually lies in the Mexican state of Durango, 2823, J. T. Polhemus & A. R. Gillogly (DAPC). Canal rather than Sonora (see subsequent discussion under Zone: 15 ♂, 11 ♀, small waterfall and stream at La Cascada O. rotundus). (trib. to Rio Pedro Miguel), on Madden Hwy., 4 km S of Distribution. Known only from Mexico (Oaxaca and Maria Eugenia, 100 m, water temp. 24° C., 31 December Durango) (Fig. 73). The two specimens from Oaxaca 1992, CL 2771, J. T. Polhemus (DAPC); 3 ♂, 7 ♀, Pipeline originate from localities in close proximity, along Road, km 17, Rio Agua Salud, sea level, 5 January 1993, Mexico Hwy. 175 between Candelaria Loxicha and CL 2785, J. T. Polhemus (DAPC); 3 ♂, Barro Colorado Santa Maria Jalatengo. Island, 2 January 1924, C. H. Curran (USNM ex JTPC); Ecological notes. Based on the limited number of Coclé: 2 ♂, 2 ♀, small stream 4.1 km W of El Valle, specimens at hand, this seems to be a cryptic species 17 January 1993, CL 2839, J. T. Polhemus (DAPC); 2 ♀, that occurs on seeps or in forest leaf litter. Cocle, El Valle, 829 m, 26 May 1983, P. J. Spangler, R. A. Downloaded from Brill.com10/08/2021 06:09:55PM via free access
Fig. 74. Ochterus manni Hungerford, dorsal habitus. Fig. 75. Ochterus manni Hungerford, dorsal habitus. Male specimen from Costa Rica, Guanacaste, 15 km N Male specimen from Panama, San Carlos. of Palmar Norte, CL 1970.
Faitoute & W. E. Steiner (USNM); 2 ♀, Galeta Island, General coloration dark brown to reddish brown 9°23'N, 79°52'W, 15 January 1974, D. Englemann with scattered yellow-orange markings, hemelytra (USNM ex JTPC). Panama: 4 ♂, 3 ♀, stream approx. flecked with bluish to lavender pruinose patches 14 km from Hwy. 1 on Nusagandi road, 350 m, 3 January (Figs 74, 75); frons black with reddish overtones 1993, CL 2776, J. T. Polhemus & A. R. Gillogly (DAPC). centrally and green laterally (Fig. 76), eyes also with San Blas: 18 ♂, 11 ♀, Nusagandi River, 200 m, water temp. green overtones, remainder of head black; pronotum 24.5° C, 1 January 1993, CL 2772, J. T. Polhemus & dark brown, lateral flanges yellow, posterior edge of A. R. Gillogly (DAPC). pronotum margined with yellowish orange, central section with scattered bluish to lavender pruinose Extralimital material examined. Bolivia, Huanchi: 1 ♂, patches, consisting of a pair (1+1) of small patches Rio Beni, Sept., Mulford Bio. Expl. 1921–1922 (holotype, on anterior margin behind the eyes, 4 transverse USNM). patches centrally near transverse midline, a small Brazil, Pará: 1 ♂, Belem, 22 October 1963, Oliviera and patch inside each posterolateral angle, and a small Wygodzinsky (USNM). Ecuador, Guayas: 2 ♂, 10 mi. N patch posteromedially; scutellum dark brown, with of Manglar Alto, Guayas, 30 January 1955, E. I. Schlinger bluish pruinose patches at anterolateral and poste- and E. S. Ross (USNM). Peru, San Martin: 2 ♂, 8 km rior angles, and a pair (1+1) of patches at anterior ENE of Tarapoto, 550 m, 9 January 1974, R. T. Schuh margin to either side of longitudinal midline; hem- (USNM). elytra dark rusty reddish brown, marked with a pair (1+1) of dark yellow spots on inner corium behind Diagnosis. Body form elongate ovate; male body posterior claval apex, and bluish to lavender flecks length 4.10–4.30, maximum width (across hemely- on each hemelytron consisting of 5 relatively evenly tra) 2.10–2.20; female body length 4.20–4.45, max- spaced patches along costal margin, a patch at the imum width (across hemelytra) 2.20–2.30. inner basal margin of the clavus, and 4 irregular Downloaded from Brill.com10/08/2021 06:09:55PM via free access
Fig. 76. Ochterus manni Hungerford, head, anterior view. Male specimen from Costa Rica, Guanacaste, 15 km N of Palmar Norte, CL 1970. patches on the inner corium, costal margin dark yel- low along entire length, membrane dark brown, irregularly maculate with bluish to lavender pru- inose patches (Figs 74, 75). Frons with fine trans- verse striations, with only narrow channels parallel to inner eye margins, upper portion of frons with an obvious raised striation along longitudinal midline (Fig. 76). Basal head, pronotum, scutellum and hemelytra set with scattered short, stout, pale setae (Figs 74, 75); posterior pronotum, clavus and corium coarsely punctate, these punctations bluish pruinose in constrast to surrounding dark brown ground color (Figs 74, 75). Male genitalia with right paramere U-shaped, distal arm long, slender, tapering (Fig. 77), apex slightly expanded, bearing a small, papillate, bulbous process at tip (Fig. 78); left paramere small, expanded basally, distal section crescent-shaped (Figs 79, 80); pygophore with medial process produced, apex roughly triangular (Fig. 81); aedeagus with distal flagellum very long, over 6× length of phallobase, terminal section enclosed Figs 77–81. Ochterus manni Hungerford, male geni- within narrow, appressed phallotheca (Fig. 14). talic structures – 77, right paramere; 78, detail of termi- Comparative notes. Ochterus manni possesses a very nal section of right paramere; 79, left paramere; 80, left distinctive male right paramere, which is slightly paramere, alternate view; 81, apex of pygophore. Speci- men from Costa Rica, Puntarenas, Boca de B arranca, expanded on the apical portion of the distal arm, with CL 1306. a small, bulb-like tip bearing tiny papillate structures (Figs 77, 78). The distal flagellum of the male aedeagus is extremely long, being over six times the yellow-orange patches on the inner angles of the length of the phallobase (Fig. 14), and is curved back corium behind the claval apex, although the degree upon itself in a large loop when at rest within the of development of these patches is intraspecifically pygophore. External characters useful in recognizing variable (Figs 74, 75). this species include the narrow yellowish-orange mar- We have examined the holotype of this species, gining on the posterior border of the pronotum, and a male held in the Smithsonian Institution. The the presence of a pair (1+1) of roughly circular specimen bears the labels: “Huachi, Rio Beni, Boliv., Downloaded from Brill.com10/08/2021 06:09:55PM via free access
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Ochterus ovatus
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Ochterus manni 5 5 SCALE 0 100 200 300 400 500 600 MILES 0 200 400 600 800 KILOMETERS CONIC PROJECTION
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Fig. 82. Distribution of Ochterus species in Mesoamerica. ● O. ovatus n. sp.; ■︎ O. manni Hungerford.
W. M. Mann (on white paper)/Sept. (on white O. manni, because his Figs 1 and 4 closely resemble paper)/Mulford, BioExpl, 1921–22/Europe Trip our Panamanian material. Nieser & Chen (1992) 1928 No., H. B. Hungerford (on white paper)/ made no reference to O. manni in their description Holotype, O. manni, H. B. Hungerford (on red of O. unidentatus, and thus may have overlooked paper)/Type No. 58598 U. S. N. M. (on red paper)”. these similarities. Confirmation of this potential syn- The specimen is in good condition and retains all its onymy will require comparison of the holotypes of appendages; the genitalia have been dissected and are the two species involved. in a vial below the specimen on the same pin. Ecological notes. This species occurs on wet rocks Distribution. This species was originally described along stream margins. It is the most commonly from Bolivia (Huachi, Rio Beni). It seems to range encountered Ochterus species in Panama, where it north into Mesoamerica only as far as Costa Rica has been taken at elevations from sea level to slightly (Fig. 82). Schell (1943) noted that she had examined over 800 m. additional specimens from Peru and Costa Rica, and Cianferoni (2012) provided records for Ecuador. Ochterus mexicanus n. sp. Additional extralimital records are provided herein Figs 83–88, 90, 91 from the latter country, as well as Panama, Costa Rica and Brazil. Type material. Holotype, male: Mexico, Sinaloa, Rio Nieser & Chen (1992) described O. unidentatus Quelite at El Quelite [23°32'11"N, 106°28'46"W, elev. from Ecuador, comparing it to O. bidentatus Schell. 25 m], 29 November 1968, CL 1235, J. T. Polhemus Cianferoni (2012) subsequently provided color (USNM ex JTPC). Paratypes: Mexico, Colima: 1 ♀, Rio illustrations of dorsal color pattern variation in this Armeria E of Armeria [18°56'25"N, 105°50'16"W, elev. taxon, and of the male right paramere in two differ- 25 m], 26 November 1968, CL 1229, J. T. Polhemus ent orientations. Although we have not examined (USNM ex JTPC). Guerrero: 12 ♂, 8 ♀, seeping roadside specimens of O. unidentatus, which occurs outside rock face 35 mi. S of Chilpancingo, 26 April 1964, the geographic area covered by the present work, we CL 1044, J. T. Polhemus (USNM ex JTPC); 2 ♀, rocky strongly suspect based on Cianferoni’s illustrations upland stream 40 mi. N of Acapulco [17°18'02"N, that this species may in fact be a synonym of 99°30'19"W, elev. 850 m], 26 April 1964, CL 1045, J. T. Downloaded from Brill.com10/08/2021 06:09:55PM via free access
Polhemus (USNM ex JTPC). Jalisco: 1 ♀, Rio Grande de pruinose patches centrally on anterior margin to Santiago at Puente Guadelupe [N of Guadalajara on either side of midline; hemelytra rusty reddish brown, Hwy. 54, 20°50'20"N, 103°19'45"W, elev. 910 m], marked with a pair (1+1) of pale lavender-grey spots 24 November 1968, CL 1224, J. T. Polhemus (USNM ex on inner corium behind posterior claval apex, and JTPC). Morelos: 6 ♂, 4 ♀, Cuautla, small rocky river bluish to lavender flecks on each hemelytron consist- [Rio Cuautla, 18°50'36"N, 98°56'06"W, elev. 1375 m], ing of 5 relatively evenly spaced patches along costal 25 April 1964, CL 1035, J. T. Polhemus (USNM ex margin, an elongate patch at the inner basal margin JTPC); 1 ♂, 1 ♀, sulphur springs 2 mi. S of Cuautla [vic. of the clavus, another small patch on basal third of 18°48'56"N, 98°56'48"W, elev. ~1315 m], 25 April 1964, clavus, and 5 irregular patches on the inner corium, CL 1036, J. T. Polhemus (USNM ex JTPC); 1 ♂, 8 mi. costal margin with 3 dark yellow marks on basal half, E of Zacatepec, swift river with gravel bed [SE of Zacatepec these markings lying immediately lateral to corre- de Hidalgo, 18°33'40"N, 99°06'00"W, elev. 860 m], sponding lavender pruinose patches on outer corium, 25 April 1964, CL 1037, J. T. Polhemus (USNM ex membrane dark brown, irregularly maculate with JTPC); 12 ♂, 9 ♀, 20 mi. S of Cuernavaca, swift rocky bluish to grey pruinose patches (Fig. 83); thoracic upland stream [vic. 18°38'49"N, 99°18'07"W, elev. ~920 venter dull brownish black, acetabulae and posterior m], 27 April 1964, CL 1047, J. T. Polhemus (USNM ex sections of pro-, meso-, and metapleurae yellowish JTPC); 6 ♂, 6 ♀, Oaxtepec, heavily shaded spring fed brown; abdominal venter medium brown, genital stream [vic. 18°54'07"N, 98°57'19"W, elev. ~1370 m], segments yellowish brown; legs with femora straw 27 April 1964, CL 1048, J. T. Polhemus (USNM ex yellow, all femoral apices narrowly brown, fore and JTPC). Oaxaca: 5 ♂, 3 ♀, 8 mi. S of Oaxaca, small rocky middle tibiae with broad, indistinct brown annula- mountain stream [vic. 17°06'21"N, 96°38'32"W, elev. tions centrally. ~1705 m], 29 April 1964, CL 1060, J. T. Polhemus Structural characters. Head — Length (along midline (USNM ex JTPC). Puebla: 2 ♂, 8 mi. W of Izucar de as measured from directly above)/width (across eyes) Matamoros on road from Cuautla, swift rocky stream = 0.35/1.20, angled downward at greater than 45° bordered by large trees [vic. 18°37'38"N, 98°34'22"W, when viewed laterally; frons with coarse transverse elev. ~1275 m], 27 April 1964, CL 1051, J. T. Polhemus (USNM ex JTPC).
Description Male. General form elongate ovate, body length 3.50–3.70, maximum width (across hemelytra) 2.00–2.10. Color. General coloration dull black to reddish brown flecked with small pale bluish pruinose patches, posterior pronotum and posterior half of clavus with scattered coarse punctations, these punc- tations bluish to lavender pruinose in constrast to surrounding dark brown ground color (Fig. 83); head with vertex dull dark blackish brown with small bluish to lavender pruinose patches, eyes silvery, frons, tylus, clypeus, labrum, basal joint of rostrum shining black (Fig. 84), distal joints of rostrum yel- lowish brown, antennae with segment I whitish yel- low, segment II pale whitish yellow clouded with grey, segments III and IV brown; head including frons and vertex shining black with vague dark green overtones, eyes silvery; pronotum black, lateral flanges dark yellow surrounding with a brown mark along outer margins, posterior pronotal margin yel- lowish orange medially, posterior pronotum with scat- tered bluish to lavender pruinose patches, consisting of 2 small patches on anterior margin behind eyes, 4 transverse patches centrally near transverse midline, a small patch well inside each posterolateral angle, and a small patch posteromedially; scutellum black, Fig. 83. Ochterus mexicanus n. sp., dorsal habitus. with bluish pruinose patches at anterolateral and Female specimen from Mexico, Morelos, Oaxtepec, posterior angles, and a pair (1+1) of large bluish grey CL 1048. Downloaded from Brill.com10/08/2021 06:09:55PM via free access
Figs 89–90. Head and pronotal morphology of Ochterus species – 89, O. parvus Schell, male holotype specimen from Ecuador, Mera; 90, O mexicanus n. sp., male specimen from Mexico, Morelos, Cuautla, CL Figs 85–88. Ochterus mexicanus n. sp., male genitalic 1035. structures – 85, right paramere, inner view of apex; 86, right paramere; 87, detail of terminal section of right the vertex set well below the top level of the eyes paramere; 88, male left paramere. Specimen from Mex- (Fig. 89), whereas in O. mexicanus the frons is about ico, Morelos, Cuautla, CL 1035. even with the level of the eyes (Fig. 90), and the ver- tex rises very obviously above the tops of the eyes. A male specimen from Montebello Lakes, in the (sixth visible) female abdominal sternite; this charac- Chiapas highlands near the Guatemala border, is ter is lacking in O. aeneifrons. The second is the dis- large for this species (4.5 mm body length) and has crete nature of the light spots along the lateral a weak discontinuity between the pronotal disc and hemelytral margins, which are separated by darker the lateral flange. Specimens of O. aeneifrons have also regions along the extreme costal margins; in O. aene- been seen with similar discontinuities, which when ifrons there is by contrast a distinct tendency for more pronounced form tubercles. The Montebello development of a yellowish-brown border along the Lakes male has not been included in the paratype series, costal margin (compare Figs 23, 24, 83). because more intensive collecting in the Guatemala Within the Neotropical Region as a whole, the highlands is needed to determine whether a separate species most similar to O. mexicanus in terms of taxon exists in this region. overall body size and genitalic morphology is O. par- Distribution. Mexico (west coast and central plateau) vus Schell from Ecuador. Both the late Dr Carl (Fig. 91). Drake and the second author had previously con- Ecological notes. Ochterus mexicanus is widespread fused series of O. mexicanus with O. parvus, but the across central Mexico, occurring on wet rocks along second author was able to examine the holotype of swift upland streams, and is not an inhabitant of the latter species in the University of Kansas collec- open, sandy, littoral substrates. It is occasionally syn- tion and identified significant differences. The male topic with O. aeneifrons, the two species having been genitalic differences, while apparent, are probably collected together at two localities east of Oaxaca, not as great as they appear considering the morpho- although O. aeneifrons has a much broader overall logical plasticity of these species, but the head and range. In general, O. mexicanus seems to be most pronotal structures are clearly different. In particular, prevalent in the central highlands and the central O. parvus has the frons sunken between the eyes, and west coastal lowland regions of Mexico. Downloaded from Brill.com10/08/2021 06:09:55PM via free access
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Ochterus mexicanus 10 10
5 5 SCALE 0 100 200 300 400 500 600 MILES 0 200 400 600 800 KILOMETERS CONIC PROJECTION
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Fig. 91. Distribution of Ochterus mexicanus n. sp. in Mesoamerica.
Ochterus obscurus n. sp. black; tylus and clypeus orange brown; labrum dark Figs 12, 73, 92–97 brown (Fig. 93); rostrum yellowish brown; antennae pale whitish yellow. Pronotum dull dark brown, lat- Type material. Holotype, male: Panama, Chiriqui, 5.6 eral flanges yellowish white broadly margined with km N of Boquete, La Culebra Trail, 1800 m, wet cloud brown, small lavender pruinose patches present on forest litter, 15 June 1996, 96–131B, R. Anderson anterior margin behind each eye, calli blackish (USNM). Paratypes: Panama, Chiriqui: 5 ♂, 3 ♀, same brown maculate with bluish pruinose marks, 4 small data as holotype (USNM, BPBM). Costa Rica, Cartago: 4 bluish pruinose patches present behind calli evenly submacropterous ♂, 1 submacropterous ♀, R. N. Tapanti spaced along transverse pronotal midline, a longitu- [Tapanti nature reserve], tropical montane forest litter, dinally elongate lavender pruinose mark present 1500 m, 97–001, 4 June 1997, R. Anderson (USNM, centrally on posterior margin; mesoscutum and scu- BPBM). Guanacaste: 1 submacropterous ♂, Guanacaste tellum dull black, posterior apex with small, pale Conservation Area, Cacao Field Station, 1400 m, wet bluish pruinose patch; hemelytra dull dark blackish cloud forest litter, 20 February 1996, 024-B, R. Anderson brown trending to medium brown on costal mar- (USNM ex JTPC). gins, each hemelytron bearing scattered pale bluish to lavender pruinose markings as follows: 3 irregular patches on basal half of clavus, 5 small scattered Description patches on basal corium, a large lavender pruinose Male. General form broadly ovate, posteriorly wid- patch on inner posterior corium behind claval apex ened; body length 3.75–3.90, maximum width followed by 3–4 irregular patches posteriorly, a small (across hemelytra) 2.30–2.40. patch on basal embolium, an O-shaped mark on Color. General coloration dark reddish brown central embolium, 4 more small, roughly evenly flecked with small pale bluish pruinose patches, cen- spaced marks on costal margin from embolar frac- tral pronotum dark brown, head black, lateral spots ture to base of wing membrane, small irregular mac- on anterolateral pronotum pale yellow (Fig. 92). ulations on wing membrane (Fig. 92); thoracic Head with vertex dull black with each posterolateral venter dull brownish black centrally, orange-brown angle bluish grey; eyes bright red; frons shining laterally, with small dark punctations, overlain with Downloaded from Brill.com10/08/2021 06:09:55PM via free access
Fig. 93. Ochterus obscurus n. sp., head, anterior view. Female specimen from Costa Rica, Cartago, Tapanti.
= 0.10, 0.11, 0.30, 0.35; rostrum long, length = 2.10, exceeding to hind coxae. Pronotum — Pronotal length (midline)/width = 0.90/2.25, anterior lobe with small lateral tubercles, Fig. 92. Ochterus obscurus n. sp., dorsal habitus. F emale entire surface set with scattered, moderately short, specimen from Costa Rica, Cartago, Tapanti. recumbent, golden setae, posterior lobe bearing numerous very small, shallow, indistinct punctations; anterior collar weakly demarcated; anterolateral mar- faint bluish pruinosity, undersides of pronotal gins highly explanate, broadly curving, well demar- flanges each with diffuse pale yellow patch, ventral cated from disk, lateral angles obtuse, sharply pointed; embolium orange-brown, pro-, meso- and metaster- calli weakly tumescent; posterior margin multisinuate, num dark blackish brown, acetabulae and posterior posteriorly concave centrally above base of scutellum. sections of pro- and metapleurae orange-brown; Scutellum — Triangular, length/width = 0.65/1.05, abdominal venter including genital segments dark moderately tumescent, bearing scattered short, blackish brown except extreme lateral margins recumbent, golden setae, mesoscutum largely hidden orange brown; legs yellowish brown, all femoral api- under posterior pronotum, separated from remain- ces narrowly brown dorsally. der of scutellum by prominent transverse furrow, Structural characters. Head — Length (along midline posterior portion of scutellum with raised transverse as measured from directly above)/width (across eyes) tumescence on basal section, posterior section with = 0.55/1.40, angled downward at greater than apex slightly raised. 45° when viewed laterally; frons with coarse stria- Hemelytra — Submacropterous, strongly raised tions, ventral portion with a longitudinal depression along claval suture when viewed laterally, barely centrally flanked to either side by raised, roughly tri- exceeding tip of abdomen, clavus well defined, angular, striate tumescences, upper half of frons embolar sutures obscure, surface set with scattered coarsely rugulose, bearing prominent longitudinal moderately short, recumbent, golden setae; clavus, medial carina, inner margins of eyes bordered by adjacent inner corium, and posterior portion of broad channels, these channels bearing numerous embolium bearing shallow punctations; length of transverse striations (Fig. 93); eyes large, protruding, clavus along outside margin 1.50; membrane projecting posterolaterally very slightly beyond pos- reduced to a small flap, venation not evident. terior margin of vertex, upper margins not rising Ventral surface — Venter of thorax smooth and above level of vertex when viewed anteriorly; anten- dull, bearing scattered extremely short, pale, recum- nae with segments I and II short, globose, segments bent setae; scattered tiny, dark punctations present III and IV slender, filiform, lengths of segments I–IV on pro-, meso-, and metepisterna, and pro- and Downloaded from Brill.com10/08/2021 06:09:55PM via free access
Ochterus ovatus n. sp. Figs 82, 98–103
Type material. Holotype, male: Mexico, Chiapas, Soyalo, Rio Blanco, on road to Villahermosa ~ 14 mi. N of Hwy. 190, small swift mountain river with backwater pools [vic. 16°53'27"N, 92°55'30"W, elev. ~1200 m], 4 May 1964, CL 1086, J. T. Polhemus (USNM ex JTPC). Paratypes: Mexico, Chiapas: 1 submacropterous ♂, 1 submacropter- ous ♀, same data as holotype (USNM ex JTPC); 1 ♂, 3 ♀, small mountain stream with grassy banks, 5 mi. N of Bochil, 4 May 1964, CL 1090, J. T. & M. S. Polhemus Figs 94–97. Ochterus obscurus n. sp., male genitalic (USNM ex JTPC, BPBM); 1 ♂, small mountain stream structures – 94, right paramere; 95, detail of terminal sec- with pools, 12 mi. N of Bochil [vic. 17°04'24"N, tion of right paramere; 96, left paramere; 97, apex of py- 92°51'51"W, elev. ~1650 m], 4 May 1964, CL 1091, J. T. gophore. Specimen from Costa Rica, Cartago, Tapanti. Polhemus (USNM ex JTPC).
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Description Male. General form broadly ovate, body length 3.35– 3.50, maximum width (across hemelytra) 2.05–2.10. Color. General coloration orange-brown flecked with small pale bluish pruinose patches, head, cen- tral pronotum and scutellum black (Fig. 98); head with vertex dull black with small bluish pruinose patches at each posterolateral angle, eyes silvery, frons shining black with green overtones, tylus, clyp- eus, labrum, rostrum yellowish brown (Fig. 99), antennae with segments I and II pale whitish yellow, segments III and IV brown; pronotum black, poste- rior section orange-brown, lateral flanges yellow nar- rowly margined with orange-brown on outer margins, paired small pale, bluish pruinose patches present along anterior margin behind eyes, 4 trans- verse patches also present centrally near transverse midline; mesoscutum and scutellum black, anterior margin centrally and posterior apex with small, pale bluish pruinose patches; hemelytra orange-brown costal margins dark yellow at basal angles, each hemelytron with pale bluish pruinose markings as follows: large, irregular patch at base of clavus, a sec- ond more elongate patch on inner basal clavus, ~10 scattered flecks and small patches on corium; wing Fig. 98. Ochterus ovatus n. sp., dorsal habitus. Female membrane brown, flecked with 4–5 lavender pru- specimen from Mexico, Chiapas, 5 mi. N of Bochil, inose patches (Fig. 98); thoracic venter dull brown- CL 1090. ish black overlain with bluish pruinosity, undersides of pronotal flanges and embolium pale yellow with small dark punctations, acetabulae and posterior sec- tions of pro- and metapleurae dark yellow; abdomi- nal venter including genital segments medium brown; legs light yellowish brown, all femoral apices narrowly brown, hind tibiae with small brown spots at bases of spines. Structural characters. Head — Length (along midline as measured from directly above)/width (across eyes) = 0.40/1.20, angled downward at greater than 45° when viewed laterally; frons with coarse transverse striations, upper half of frons with prominent longitudinal medial carina, inner margins of eyes bor- dered by channels of moderate width, these channels bearing numerous transverse striations on their ven- tral halves (Fig. 99); eyes large, protruding, project- ing posterolaterally slightly beyond posterior margin of vertex, upper margins not rising above level of ver- tex when viewed anteriorly; antennae with segments I and II short, globose, segments III and IV slender, Fig. 99. Ochterus ovatus n. sp., head, anterior view. filiform, lengths of segments I–IV = 0.05, 0.10, Female specimen from Mexico, Chiapas, 5 mi. N of 0.25, 0.25; rostrum long, length = 1.90, exceeding Bochil, CL 1090. hind coxae and extending onto base of abdominal venter. punctations; anterior collar not strongly demarcated; Pronotum. — Pronotal length (midline)/width = anterolateral margins explanate, well demarcated from 0.75/1.75, anterior lobe lacking lateral tubercles, entire disk, lateral angles obtuse; calli barely tumescent; pos- surface set with scattered short, recumbent, golden terior margin multisinuate, posteriorly concave cen- setae, bearing numerous small, shallow, indistinct trally above base of scutellum.
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Scutellum — Triangular, length/width = 0.60/1.00, bearing scattered short, recumbent, golden setae, mesoscutum largely hidden under posterior prono- tum, separated from weakly tumescent remainder of scutellum by shallow transverse furrow. Hemelytra — Submacropterous, barely exceeding tip of abdomen, clavus well defined, embolar sutures obscure, surface set with scattered short, recumbent, golden setae; clavus, adjacent inner corium and embolar margin bearing shallow punctations; antero- lateral embolar margin not produced or explanate; length of clavus along outside margin 1.50; mem- brane reduced to a small flap, venation not evident. Ventral surface — Venter of thorax smooth and dull, bearing scattered extremely short, pale, recum- bent setae; scattered tiny, dark punctations present on pro-, meso-, and metepisterna, and pro- and mesoacetabulae; abdominal venter covered with very short, fine, appressed pale setae, this setal covering interrupted by ovate glabrous patches surrounding spiracles laterally on paratergites. Legs — All leg segments covered by short, fine, recumbent pale setae; fore and middle trochanters and basal portions of fore and middle femora with very long, semi-erect, slender, pale setae ventrally, remainder of ventral margins bearing moderate length, upright, slender, pale setae; hind femur bowed slightly downward when viewed laterally; fore and middle tibiae bearing numerous short, bristly golden setae, these setae becoming more numerous distally; all tibiae bearing scattered short, stout, golden spines, length of these spines on middle and hind tibiae equal to or exceeding the widths of these segments; claws golden, gently curving, arolia long, equal to or exceeding length of claws. Lengths of Figs 100–103. Ochterus ovatus n. sp., male genitalic leg segments as follows: fore femur/tibia/tarsal structures – 100, right paramere; 101, detail of terminal 1/tarsal 2 = 1.00/0.95/0.08/0.10; middle femur/ section of right paramere; 102, left paramere; 103, apex tibia/tarsal 1/tarsal 2 = 1.00/0.95/0.09/0.10; hind of pygophore. Specimen from Mexico, Chiapas, Soyalo, femur/tibia/tarsal 1/tarsal 2 /tarsal 3 = 1.10/1.45/ CL 1086. 0.05/0.15/ 0.18. Male genitalia — Right paramere U-shaped, dis- tal arm moderately long (Fig. 100), distal section Comparative notes. Ochterus ovatus is a small slightly expanded, twisted, tip slightly hooked species, currently known only from brachypterous (Fig. 101); left paramere small, broadly crescentic forms, that may be easily recognized by its orange- (Fig. 102); pygophore with caudal tip weakly pro- brown coloration with a contrastingly dark head, duced, apex broadly rounded (Fig. 103); aedeagus pronotum and scutellum (Fig. 98). It is distinct with distal arm moderately short, about 4× length of in the Mesoamerican ochterid biota, though phallobase, terminal section enclosed by an expanded reminiscent of some South American forms. In phallotheca. addition to the dorsal coloration, the male right Female. Similar to male in general structure, color- paramere with its relatively slender and twisted ation, and body proportions, but slightly larger in distal arm is diagnostic (Figs 100, 101). Additional overall size; body length 3.65–4.00, maximum width notable characters are the protruding eyes, which (across hemelytra) 2.25–2.45. project posterolaterally slightly beyond posterior Etymology. The name ‘ovatus’ is derived from the margin of vertex, and the long arolia, the lengths Latin ‘ovum’, or egg shaped, and refers to the overall of which are equal to or exceed the lengths of body shape of this species. the claws.
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Distribution. Mexico (Chiapas) (Fig. 82). Ecological notes. This species occurs along the mar- gins of small, first order upland streams.
Ochterus panamensis n. sp. Figs 10, 104–110
Type material: Holotype, male: Panama, Panama, small stream in limestone bed at Serrania de Majé, 3 km S of Ipete, 100 m [vic. 8°57'37"N, 78°31'17"W], 3 January 1993, CL 2779, J. T. Polhemus (USNM ex JTPC). Paratypes: Panama, Panama: 3 ♂, 2 ♀, same data as holo- type (USNM ex JTPC, BPBM); 1 ♂, 1 ♀, Rio Ipete at Ipete, 100 m, 3 January 1993, CL 2777, J. T. Polhemus (USNM ex JTPC).
Description Male. General form elongate ovate, body length 4.00–4.15, maximum width (across hemelytra) 2.10–2.20. Color. General coloration black flecked with small pale bluish pruinose patches, lateral flanges of prono- tum orange-brown; head with vertex dull dark black- ish brown with small bluish pruinose patches at each posterolateral angle (Fig. 104), eyes silvery, frons brilliant metallic green, clypeus yellowish orange, labrum dark brown (Fig. 105), rostrum pale yellow- Fig. 104. Ochterus panamensis n. sp., dorsal habitus. ish brown, antennae with segments I and II pale yel- Female specimen from Panama, Serrania de Mahé, CL lowish brown, segments III and IV dark brown. 2779. Pronotum black, lateral flanges orange-brown nar- rowly margined with black, paired small, pale, bluish pruinose patches present behind eyes, at inner angles of lateral flanges adjoining lateral tubercles, and as a pair (1+1) centrally to either side of longitudinal midline. Scutellum including mesoscutum dull black, with basolateral angles, small mark centrally at middle of anterior margin, and posterior apex pale bluish pruinose. Hemelytra dull black with faint greenish overtones, slightly embrowned along pos- terolateral margins; embolium bearing 3 evenly spaced small bluish pruinose patches along its length, with middle patch larger than others; clavus heavily mottled with bluish pruinose patches on basal half, and with a smaller bluish pruinose patch at apex; basal corium with a few scattered small bluish pru- inose patches, central corium with a roughly circular bluish pruinose patch adjacent to claval apex, distal corium with a single small bluish pruinose patches at base of wing membrane; wing membrane with scattered large pale bluish pruinose patches across Fig. 105. Ochterus panamensis n. sp., head, ante- outer basal cell, extensively mottled with bluish pru- rior view. Female specimen from Panama, Serrania de inose posterior to the closed cells (Fig. 104); thoracic Mahé, CL 2779. venter dull black extensively overlain by blue-grey pruinosity, posterior margins of acetabulae pale yel- Structural characters. Head — Length (along mid- lowish, abdominal venter medium brown; legs yel- line as measured from directly above/width (across lowish brown, with tips of femora, tibiae, and eyes) = 0.40/1.20, angled downward at greater than terminal tarsal segments dark brown. 45° when viewed laterally; frons with coarse striations Downloaded from Brill.com10/08/2021 06:09:55PM via free access
weakly demarcated; anterolateral margins explanate, narrow, well demarcated from disk; calli weakly tumescent; posterior margin multisinuate, posteri- orly concave centrally above base of scutellum. Scutellum — Large, triangular, length/width = 0.70/1.20, bearing scattered very short, obscure, recumbent, gold setae, mesoscutum not raised; basal portion of posterior portion of scutellum behind mesoscutum transversely raised; scutellar apex acute. Hemelytra — Long, attaining tip of abdomen, with corium, clavus and embolium well defined, surfaces set with scattered very short, obscure, recumbent, gold setae; clavus, corium and embolium without evi- dent punctations; anterolateral embolar margin only very narrowly explanate; length of clavus along out- side margin 1.80; membrane venation defined by small gold setae, with 9 evident closed cells basally. Ventral surface — Venter of thorax smooth and dull, bearing scattered extremely short, pale, recum- bent setae; scattered tiny, dark punctations present on pro-, meso-, and metepisterna, these punctations also present on ventral surfaces of pronotal flanges; abdominal venter covered with very short, fine, appressed pale setae, this setal covering interrupted by ovate glabrous patches surrounding spiracles lat- erally on paratergites. Legs — All leg segments covered by short, fine, recumbent pale setae; fore and middle trochanters and basal portions of fore and middle femora with moderately long, semi-erect, slender, pale setae ven- trally, remainder of ventral margins bearing moder- ate length, upright, slender, pale setae; hind femur bowed very slightly downward when viewed later- ally; all tibiae bearing numerous short, bristly golden setae, these setae becoming more numerous distally, interspersed with scattered short, stout, golden spines; Figs 106–109. Ochterus panamensis n. sp., male geni- hind tibia bearing fringe of very long, slender, golden talic structures – 106, right paramere; 107, detail of setae along posterior margin on basal half; claws terminal section of right paramere; 108, left paramer e; golden, gently curving, arolia long, slender, length 109, apex of pygophore. Specimen from Panama, slightly less than length of claws. Lengths of leg seg- Serrania de Mahé, CL 2779. ments as follows: fore femur/tibia/tarsal 1/tarsal 2 = 1.05/1.00/0.06/0.20; middle femur/tibia/tarsal 1/ centrally, inner margins of eyes bordered by narrow tarsal 2 = 1.10/1.10/0.10/0.20; hind femur/tibia/tar- channels, these channels bearing numerous trans- sal 1/tarsal 2/tarsal 3 = 1.45/1.90/0.05/0.18/ 0.20. verse striations (Fig. 105); eyes large, protruding, not Male genitalia — Right paramere U-shaped, distal projecting posterolaterally beyond vertex, upper arm relatively long, slender basally, apical section margins equal with vertex when viewed anteriorly; moderately expanded (Fig. 106), with angled spur antennae with segments I and II short, globose, on inner margin (Fig. 107); left paramere small, pre- segments III and IV slender, filiform, lengths of seg- apically notched, apex slightly expanded, angled ments I–IV = 0.06, 0.10, 0.30, 0.20; rostrum (Fig. 108); pygophore with medial process moder- moderately long, length = 1.90, extending to hind ately produced, apex angled, roughly triangular in coxae. shape (Fig. 109); aedeagus with distal flagellum Pronotum — Pronotal length (midline)/width = moderately long, approximately 4x length of phallo- 0.85/2.20, anterior lobe lacking lateral tubercles, base, terminal portion not recurved, apex acuminate, entire surface set with scattered very short, obscure, not bifurcated, contained within expanded phal- recumbent, gold setae, bearing numerous small, lotheca; proctiger roughly conical in shape, apex prominent, shallow punctations; anterior collar rounded (Fig. 10). Downloaded from Brill.com10/08/2021 06:09:55PM via free access
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Fig. 110. Distribution of Ochterus species in Mesoamerica. ● O. schellae Drake; ■︎ O. panamensis n. sp.
Female. Similar to male in general structure and col- Material examined. Belize, Stann Creek Dist.: 1 ♂, Sittee oration but slightly larger in overall size, body length Point, collected on riverbank at Possum Point Biological 4.60, maximum width (across hemelytra) 2.40. Station, 29 April 1987, P. Spangler and R. Faitoute Etymology. The name ‘panamensis’ refers Panama, (USNM ex JTPC); 1 ♀, Twin Cays, West Bay, Malaise trap, the country of origin for the type series, and the only 17–27 January 1987, W. Mathis and C. Feller (USNM). country from which this species is so far known. Guatemala: 1 ♂, 2 ♀, Izabal, 5.4 mi. S of Puerto Barrios, Comparative notes. Ochterus panamensis is similar to 1 February 1956, T. J. Cantral (UMMZ). Honduras, O. smaragdinus n. sp. in exhibiting green overtones Comayagua: 3 ♂, 3 ♀, Hotel los Remos, S side of Lago over a dark ground color on the pronotum and hem- Yojoa, 15 November 1987, L. Knutson (USNM); 2 ♂, 2 ♀, elytra (Fig. 104). It may be separated from that spe- 17 mi. E of Jicaro Galan, shores of rocky river, 9 January cies by the posteriorly angular apex of the pygophore 1970, CL 1311, J. T. Polhemus (USNM ex JTPC). (Fig. 109), and by the shape of the male right para Mexico, Baja California Sur: 1 ♂, 2 ♀, 8 km W of San Jose mere, which bears a spur distally on the inner margin del Cabo, 11 November 1981, W. E. Steiner (USNM). (Figs 106, 107). This species displays sexual dimor- Campeche: 4 ♂, 3 ♀, Escarcega, in light trap, 3 June 1962, phism in the size of the adults, with ♂ slightly smaller F. Islas S. (USNM). Chiapas: 3 ♂, 7 mi. S of Pichucalco, than ♀ (4.2 versus 4.6 mm body length). small quiet stream with vertical banks [Rio Ixtacomitan, Distribution. Known only from the type series taken 17°25'21"N, 93°06'13"W, elev. 180 m], 5 May 1964, CL in eastern Panama from the Serrania de Mahé 1103, J. T. and M. S. Polhemus (USNM ex JTPC); 1 ♀, (Fig. 110). Chiapa del Corzo, 6 July 1963, D. R. Whitehead (USNM). Ecological notes: This species occurs on wet rocks Guanajuato: 1 ♂, 10 mi. W of Guanajuato, small stream along upland streams. with dam, 16 June 1975, CL 757, J. T. Polhemus (USNM ex JTPC). Guerrero: 1 ♀, Acapulco, 23 July 1950, Drake Ochterus perbosci (Guérin-Menéville) and Hottes (USNM). Jalisco: 1 ♀, small stream in fields nr. Figs 6, 13, 111–117 Tonila, 11 June 1975, CL 739, J. T. Polhemus (USNM ex JTPC). Mexico: 1 ♂, Tejupilco, Temescaltepec, 21 June Pelogonus perbosci Guérin-Menéville, 1843: 113. 1933, H. E. Hinton and R. L. Usinger (USNM). Morelos: Pelogonus marginatus: Uhler 1893: 706. Misidentification. 5 ♂, 1 ♀, sulphur springs 2 mi. S of Cuautla [vic. Ochterus perbosci: Kirkaldy & Torre-Bueno, 1909: 179. 18°48'56"N, 98°56'48"W, elev. ~1315 m], 25 April 1964, Downloaded from Brill.com10/08/2021 06:09:55PM via free access
CL 1036, J. T. and M. S. Polhemus (USNM ex JTPC). Nayarit: 3 ♂, 1 ♀, 10 mi. E of San Blas, small rocky stream, 28 November 1968, CL 1232, J. T. Polhemus (USNM ex JTPC). Oaxaca: 1 ♂, 8 mi. NE of Oaxaca, small rocky mountain stream [vic. 17°06'21"N, 96°38'32"W, elev. ~1705 m], 29 April 1964, CL 1060, J. T. and M. S. Polhemus (USNM ex JTPC). Puebla: 1 ♀, 20 mi. SE of Acatlan, seeps and small stream in ravine [vic. 17°59'54"N, 97°53'28"W, elev. ~1600 m], 28 April 1964, CL 1053, J. T. and M. S. Polhemus (USNM ex JTPC); 1 ♂, 1 ♀, 1 mi. S of Tehuitzingo, 22 July 1963, Ackermann & Whitehead (USNM). Sinaloa: 8 ♂, 1 ♀, Los Mochis, muddy banks of irrigation canal, 1 October 1959, W. W. Gibson (USNM); 4 ♂, 2 ♀, Rosario, Rio Baluarte, 200 ft. [65 m], 31 December 1971, ME-45, R. R. & M. E. Murray (TAMU). Tamaulipas: 2 ♀, 4 mi. S of Ciudad Victoria, 1200 ft. [365 m], 5 August 1963, Duckworth and Davis (USNM); 1 ♀, Tampico, 24 August 1974, W. E. Clark (TAMU). Veracruz: 2 ♀, Veracruz, 29 July 1950, Drake and Hottes (USNM). Nicaragua: 3 ♂, 1 ♀, Esteli, 9 July 1957, D. R. Lauck (USNM). Panama, Panama: 1 ♂, 1 ♀, 3 km S of Ipeti, 30 m, 3 May 1992, A. R. Gillogly (USNM ex JTPC). Venezuela, Aragua: 1 ♀, 2 km N of Ocumare de la Costa, 31 March–2 April 1981, A. S. Menke and L. Hollenberg (USNM). United States, Texas: 2 ♀, Cameron Co., Brownsville, 29 June 1938, R. H. Beamer (USNM); 2 ♂, 2 ♀, Hidalgo Co., pond at entrance to Bentsen-Rio Grande State Park, 11 June 1975, C. L. Smith (USNM ex JTPC). Utah: 1 , Salt Lake Co., Jordan River, 27 November ♀ Fig. 111. Ochterus perbosci (Guérin-Menéville), dorsal 1959, G. R. Hansen (USNM). habitus. Specimen from Belize, Sittee Point. Diagnosis. Body form elongate ovate; male body of 2 small patches on basal section of clavus, and length 4.90–5.50, maximum width (across hemely- 6–7 irregular patches scattered on inner corium, tra) 2.75–2.95; female body length 5.80–6.30, max- membrane dark blackish brown, irregularly maculate imum width (across hemelytra) 3.00–3.50. with pale bluish pruinose patches (Fig. 111). Frons General coloration dark blackish grey with scattered with fine transverse striations, central sections of lateral yellowish white markings, hemelytra flecked with pale margins with coarser striations adjacent to inner mar- bluish pruinose patches (Fig. 111); frons metallic gins of eyes (Fig. 112). Basal head, pronotum, scutel- green with reddish overtones centrally and laterally lum and hemelytra set with scattered short, stout, pale (Fig. 112), eyes dark blackish grey, remainder of head setae (Fig. 111); pronotum, clavus and corium lacking black except extreme posterior margin pruinose grey; obvious punctations (Fig. 111), anterolateral angles of pronotum black, lateral flanges pale yellow, posterior pronotum produced to form small hooks (Fig. 115). margin narrow brown medially, pronotal disk with Male genitalia with right paramere U-shaped, dis- scattered pale bluish pruinose patches, 4 transverse tal arm long, slender, bearing prominent fin on outer patches centrally near transverse midline, a small patch margin (Fig. 113), apex weakly crescentic (Fig. 114); inside set well inside each posterolateral angle, and a left paramere small, finger-like (Fig. 115); pygophore small patch posteromedially; scutellum black, with a with apex roughly triangular, apex acute; aedeagus pair (1+1) of pale bluish patches at anterior margin to with distal flagellum moderately elongate, over 2.5× either side of longitudinal midline, another similar the length of phallobase, distal portion recurved and pair of patches immediately posterior to the preceding, enclosed within prominent phallotheca (Fig. 13); and a final pair of small patches to either side of proctiger broad, lateral margins nearly parallel, pos- posterior apex; hemelytra dark blackish grey, each terior margin rounded, apex slightly truncate (Fig. 6). hemelytron marked with a pair of narrow yellowish Comparative notes. Ochterus perbosci was described patches at base of costal margin, followed by 3 larger based on specimens collected by naval surgeon M. and regularly spaced whitish yellow patches on poste- Perbosc at the Bay of Campeche, Mexico. The small, rior portion of costal margin, inner corium and clavus hook-like points lying at the anterolateral angles of with scattered pale blue pruinose patches consisting the pronotum (Fig. 116) are a diagnostic character Downloaded from Brill.com10/08/2021 06:09:55PM via free access
Fig. 112. Ochterus perbosci (Guérin-Menéville), head, anterior view. Specimen from Belize, Sittee Point. for this widespread species, as is the lack of obvious punctations on the pronotum, scutellum or hemely- tra (Fig. 111). The male right paramere has a promi- nent flange along the outer margin (Figs 113, 114), a character state similar to that seen in O. banksi (Figs 43, 44); this appears to be a synapomorphy linking these two species. A somewhat similar but less well-developed expression of this character state is also seen in O. americanus (Figs 33, 34). The distal flagellum of the male aedeagus is moderately elongate in this species, with the distal point of articulation weakly developed, and the terminal phallotheca broadly expanded (Fig. 13). The posterior tip of the pygophore is produced and triangular, with the apex acute, very similar to the character state seen in O. banksi and O. americanus (Figs 36, 45). Distribution. This species is widespread in the Western Hemisphere, occurring at elevations from sea level to at least 1600 m, and from Paraguay, Brazil, Peru, Ecuador and Suriname (Cordiero et al. 2014) northward through the West Indies and Mexico to the southwestern United States, with pre- vious confirmed records for the latter area only from Arizona and Texas (Polhemus & Polhemus 1988). Only a representative set of Mesoamerican records are provided above; these do not encompass the full hemispheric range of this widespread species as noted above, but do include a new United States Figs 113–116 Ochterus perbosci (Guérin-Menéville), record for Utah, and more comprehensive distribu- morphological details – 113, male right paramere; 114, tional data for Mexico (Fig. 117). detail of terminal section of male right paramere; 115, Champion (1901) examined the holotype from male left paramere; 116, pronotal margin, left side, Campeche, Mexico, and listed additional Mexican showing location of small hooked process at anterolat- specimens from Mazatlan and Veracruz, as well as eral angle. from the Caribbean Islands of Cuba, Grenada and St. Vincent. The records from Grenada and Downloaded from Brill.com10/08/2021 06:09:55PM via free access
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Fig. 117. Distribution of Ochterus perbosci (Guérin-Menéville) in Mesoamerica.
St. Vincent have been reconfirmed, but no speci- JTPC); 1 ♀, Cola de Caballo [vic. 25°21'45"N, mens from Cuba were present in any of the major 100°09'49"W], 3000 ft. [915 m], 6 July 1959, R. B. collections so far examined in the course of this Selander and J. C. Schaffner (USNM ex JTPC). San Luis study. A recent review of the aquatic Heteroptera Potosi: 2 ♂, 6 ♀, 32 miles E of San Luis Potosi, 26 July of Cuba (Naranjo et al. 2010) also concluded on 1970, Murray, Phelps, Hart & Schaffner (TAMU). the basis of literature review and local museum col- Tamaulipas: 2 ♂, 4 ♀, same data as holotype (USNM, lections that this species does not occur on Cuba. BPBM). Veracruz: 1 ♀, 11 mi. S of Misantla, 21 March Similarly, extensive collections by the authors on 1975, Clark & Schaffner (TAMU). Hispaniola in 1998 and 1999, and limited sam- pling in suitable habitats on Puerto Rico by the Description first author in 2015 have not revealed the presence Male. General form elongate ovate, body length 5.30– of this species. Based on the preceding, it appears 5.70, maximum width (across hemelytra) 3.00–3.20. that O. perbosci may not occur on any of the islands Color. General coloration dull black flecked with in the Greater Antilles. bluish and purple pruinose patches, lateral flanges of Ecological notes. Ochterus perbosci occurs on damp, pronotum orange-brown (Figs 118, 119); head with horizontal, littoral substrates, and has a notable vertex dull black on anterior half, dark bluish pru- degree of salt tolerance, being found along seacoasts inose on posterior half, eyes shining black, frons and the margins of saline lakes. brilliant metallic green with overtones of yellow, clypeus yellowish orange, labrum black (Fig. 120), Ochterus pseudorotundus n. sp. rostrum pale yellowish brown, antennae with seg- Figs 7, 118–125 ments I and II pale yellowish brown, segments III and IV medium brown. Pronotum dull black, lateral Type material. Holotype, male: Mexico, Tamaulipas, flanges orange-brown narrowly margined with black, Bocatoma, 7 km SSE of Gomez Farias [vic. 23°01'04"N, paired small, pale, bluish pruinose patches presentat 99°07'28"W, elev. 135 m], 19–23 May 1979, E. G. Riley inner angles of lateral flanges adjoining lateral tuber- (USNM ex JTPC). Paratypes: Mexico, Nuevo Leon: 1 ♂, cles, and as a pair (1+1) centrally to either side of 1 ♀, Santiago, Potrero Redondo [vic. 25°14'49"N, longitudinal midline. Scutellum including mesoscu- 100°09'43"W], 9 June 1985, A. Contreras R. (USNM ex tum dull black, with basolateral angles, small mark Downloaded from Brill.com10/08/2021 06:09:55PM via free access
Fig. 119. Ochterus pseudorotundus n. sp., dorsal habi- tus. Female specimen from Mexico, Nuevo Leon, Potre- ro Redondo.
= 0.40/1.40, angled downward at greater than 45° when viewed laterally; frons with coarse striations centrally, inner margins of eyes bordered by narrow channels, these channels bearing numerous trans- verse striations, upper half of frons with low carina along longitudinal midline, ventral half of frons above clypeus slightly raised to form a triangular elevation with apex directed upward (Fig. 120); eyes large, protruding, not projecting posterolaterally beyond vertex, upper margins equal with vertex when viewed anteriorly; antennae with segments I and II short, globose, segment III slender, parallel sided, segment IV fusiform, lengths of segments I–IV = 0.10, 0.15, 0.30, 0.25; rostrum moderately long, length = 2.10, extending to hind coxae. Pronotum — Pronotal length (midline)/width = 0.90/2.50, anterior lobe bearing lateral tubercles, entire surface set with scattered very short, obscure, recumbent, gold setae, rugulose and bearing coarse, shallow punctations on posterior lobe; anterior collar weakly demarcated; anterolateral margins explanate, narrow, well demarcated from disk; calli weakly Fig. 118. Ochterus pseudorotundus n. sp., dorsal habitus. tumescent; posterior margin multisinuate, posteri- Female specimen from Mexico, Tamaulipas, Bocatoma. orly concave centrally above base of scutellum. Downloaded from Brill.com10/08/2021 06:09:55PM via free access
Fig. 120. Ochterus pseudorotundus n. sp., head, ante- rior view. Female specimen from Mexico, Nuevo Leon, Potrero Redondo.
Scutellum — Large, triangular, length/width = 0.90/1.30, bearing scattered very short, obscure, recumbent, gold setae and shallow coarse punctations, mesoscutum not raised; basal portion of posterior portion of scutellum behind mesoscutum weakly transversely raised; scutellar apex acute. Hemelytra — Long, attaining tip of abdomen, with corium, clavus and embolium well defined, surfaces set with scattered very short, obscure, recumbent, gold setae; clavus with coarse shallow punctations on inner half bordering scutellum, corium and embo- lium without evident punctations; anterolateral embo- lar margin only very narrowly explanate; length of clavus along outside margin 1.80; membrane vena- tion defined by small gold setae, with 8 evident closed cells basally. Ventral surface — Venter of thorax smooth and dull, bearing scattered extremely short, pale, recum- bent setae; scattered tiny, dark punctations present on pro-, meso-, and metepisterna, these punctations absent on ventral surfaces of pronotal flanges; abdomi- nal venter covered with very short, fine, appressed pale setae, this setal covering interrupted by ovate glabrous patches surrounding spiracles laterally on paratergites. Figs 121–124. Ochterus pseudorotundus n. sp., mor- Legs — All leg segments covered by short, fine, phological details – 121, right paramere; 122, detail recumbent pale setae; fore and middle trochanters and of terminal section of right paramere; 123, left para- all femora with moderately long, semi-erect, slender, mere; 124, apex of pygophore. Specimen from Mexico, pale setae ventrally; hind femur not bowed when viewed Tamaulipas, Bocatoma. laterally; all tibiae bearing short, bristly golden setae, these setae interspersed with scattered short, stout, golden spines; hind tibia bearing fringe of very long, segments as follows: fore femur/tibia/tarsal 1/tarsal 2 = slender, golden setae along posterior margin on basal 1.40/1.25/0.05/0.25; middle femur/tibia/tarsal 1/tarsal half; claws golden, gently curving, arolia short, slender, 2 = 1.50/1.50/0.05/0.35; hind femur/tibia/tarsal length distinctly less than length of claws. Lengths of leg 1/tarsal 2/tarsal 3 = 1.70/2.50/0.05/0.20/ 0.40. Downloaded from Brill.com10/08/2021 06:09:55PM via free access
115 110 105 100 95 90 85 80 75 70 65 60
35 35
30 30
25 25
20 20
15 15
Ochterus pseudorotundus 10 10
Ochterus shepardi 5 5 SCALE 0 100 200 300 400 500 600 MILES 0 200 400 600 800 KILOMETERS CONIC PROJECTION
110 105 100 95 90 85 80 75 70 65 WEST LONGITUDE
Fig. 125. Distribution of Ochterus species in Mesoamerica. ● O. pseudorotundus n. sp; ■︎ O. shepardi n. sp.
Male genitalia — Right paramere U-shaped, distal broadly and consistently embrowned, and the large arm relatively long, slender basally, apical section pruinose patch behind the claval apex is slightly moderately expanded, paddle-like (Figs 121, 122); smaller, although also paler and thus more promi- left paramere small, angled distally, apex slightly nent. The male right paramere of O. pseudorotundus bifurcate (Fig. 123); pygophore with medial process lacks the flanges on the inner margin seen in O. moderately produced, apex rounded, flanked by rotundus (compare Figs 121, 122 to 128–130), and concave subapical notch (Fig. 124); aedeagus with the shapes of the male proctigers are very distinct distal flagellum moderately short, approximately (compare Figs 7 and 9). Despite these differences, the 4× length of phallobase, terminal portion contained taxa are clearly closely related, and seem to be the within expanded phallotheca; proctiger elongate, eastern and western segr egates of a common stock conical, basal lateral lobes weakly developed (Fig. 7). that have speciated allopatrically in the mountain Female. Similar to male in general structure and col- chains of northern Mexico. oration but slightly larger in overall size, body length Distribution. This species occurs in the Sierra Madre 5.60–5.80, maximum width (across hemelytra) Occidental of eastern Mexico (Nuevo Leon, San Luis 3.10–3.30. Potosi, Tamaulipas, Veracruz) (Fig. 125). Etymology. The name ‘pseudorotundus’ refers to the Ecological notes. Nothing is recorded regarding the superficial resemblance of this species toO. rotundus. habits of this species, but given its similarities to O. Comparative notes. Ochterus pseudorotundus is a large rotundus it is likely to be found on vertical rock seeps. species occurring in the Sierra Madre Occidental of northeastern Mexico. It has been confused with Ochterus rotundus J. Polhemus & M. Polhemus O. rotundus by past workers, and intermixed in Figs 126–133 museum holdings. The body form is somewhat vari- able in terms of its breadth (Figs 118, 119), but has Ochterus viridifrons: Schell, 1943: 37 (in part). the costal margins nearly parallel basally and as Ochterus rotundus J. Polhemus & M. Polhemus, 1976: 223. such is never as broad and evenly rounded as in O. rotundus (Fig. 126). In addition, the lateral Material examined. Mexico, Durango: 4 ♂, 5 ♀, seeping margins of the hemelytra in O. rotundus are more roadside rock faces 7 mi. W of Los Bancos [vic. 23°38'54"N, Downloaded from Brill.com10/08/2021 06:09:55PM via free access
105°47'56"W, elev. ~2320 m], 20 April 1964, CL 1017, J. T. & M. S. Polhemus (holotype and paratypes, USNM ex JTPC); 1 ♂, seep at Km 183 W of Durango on Hwy. 40, 2 Janaury 1979, CLH 244, C. L. Hogue (USNM ex JTPC). Jalisco: 1 ♀, seeping roadside rock face S of Mismaloya [vic. 20°27'06"N, 105°17'37"W], 1500–2000 ft. [460–610 m], 9 June 1975, CL 734, J. T. Polhemus (USNM ex JTPC). Michoacan: 1 ♂, 1 ♀, waterfall at El Salto, ~17 mi. E of Morelia [vic. 19°39'13"N, 100°56'59"W, elev. ~2250 m], 15 June 1975, CL 751, J. T. Polhemus (paratypes, USNM ex JTPC); 9 ♂, 8 ♀, waterfall, cascades and seeps in gorge 6 mi. W of Uruapan [vic. 19°24'09"N, 102°07'07"W], ~5000 ft. [1525 m], 14 June 1975, CL 747, J. T. Polhemus (paratypes, USNM ex JTPC). Nayarit: 1 ♀, wet rock faces along swift stream with waterfall in ravine at Km 50 on highway west of Tepic, W of Compostela, 2600 ft [792 m], 8 June 1975, CL 730, J.. T Polhemus (USNM ex JTPC). Sinaloa: 1 ♂, 1 ♀, 1 immature, seeping roadside rock face 4 mi. E of La Palmita [vic. 23°35'45"N, 105°50'49"W], 6700 ft. [2040 m], 6 June 1975, CL 722, J. T. Polhemus (paratypes, USNM ex JTPC); 1 ♂, 1 ♀, E of Santa Lucia [1 mi. E of Santa Lucia, vic. 23°26'38"N, 105°50'21"W, elev. ~1230 m], 26 April 1974, M. S. Polhemus (paratypes, USNM ex JTPC). Sonora: 10 ♂, 2 ♀, Rancho de los Baños, NE of Nacozari, wet rock faces along river in canyon downstream of ranch [30°26'11"N, 109°28'56"W, elev. 990 m], 3 June 1975, CL 709, J. T. Polhemus (paratypes, USNM ex JTPC). United States, Arizona: 1 , Grand Canyon, ♀ Fig. 126. Ochterus rotundus J. Polhemus & M. Polhe- Brooklyn Museum collection, 1929 (USNM); 1 ♂, mus, dorsal habitus. Female specimen from Mexico, Coconino Co., seeps on bedrock in middle section of Buck Durango, 7 mi. W of Los Bancos, CL 1017. Farm Canyon, trib. to Colorado River in Marble Canyon, Mile 41 below Lee’s Ferry, [36°23'52"N, 111°53'80"W, pronotum narrowly margined with orange, central elev. 930 m], 28 May 1972, CL 546, J. T. and M. S. section with 4 transverse pale bluish patches cen- Polhemus (paratype, USNM ex JTPC); 1 ♀, Grand trally near transverse midline behind calli; mesoscu- Canyon, seeping rock face in upper Stone Creek Canyon, tum and scutellum black, with small, elongate pale above confluence with Colorado River, Mile 132 below bluish pruinose patches at basal angles, as a small Lee’s Ferry, [vic. 36°20'48"N, 112°27'15"W, elev. ~615 very small spot medially on anterior margin, and as a m], 31 May 1972, CL 551, J. T. & M. S. Polhemus (para- pair (1+1) of very small patches to either side of pos- type, USNM ex JTPC). terior apex; hemelytra black with dark green over- tones, costal margins diffusely orange-brown, inner Diagnosis. Body form broadly ovate and rotund; male margins of claval commissure orange, pale bluish body length 4.85–5.10, maximum width (across pruinose patches and flecks scattered on each hem- hemelytra) 2.90–3.10; female body length 5.80–6.00, elytron, consisting of a set of irregular mottling on maximum width (across hemelytra) 3.40–3.50. basal half of clavus and basal corium, a larger patch General coloration black with scattered yellow- on posterior inner corium immediately posterior to orange markings, hemelytra flecked with pale bluish claval apex, 5 relatively evenly spaced patches along to pruinose patches (Fig. 126); frons brilliant shining costal margin, forewing membrane dark brown, bluish green with metallic purple overtones centrally, irregularly maculate with pale bluish pruinosity metallic blue to aqua overtones laterally (Fig. 127), (Fig. 126). Frons with coarse transverse striations, eyes silvery, posterior portion of head behind eyes inner margins of eyes bordered by wide, shallow blackish overlain with pale bluish pruinosity, lateral channels, these channels bearing transverse striations sections adjoining inner margins of eyes black, along their entire lengths, upper portion of frons narrowly margined with orange; pronotum black, with a weakly raised striation along longitudinal anterior margin diffusely frosted with pale bluish midline (Fig. 127). Dorsal surface bearing very short, pruinosity, lateral flanges orange narrowly margined slender, recumbent gold setae, lacking prominent with black along outer edges, posterior edge of punctations (Fig. 126). Head vertex not sunken Downloaded from Brill.com10/08/2021 06:09:55PM via free access
Fig. 127. Ochterus rotundus J. Polhemus & M. Polhe- mus, head, anterior view. Female specimen from Mexi- co, Durango, 183 km W of Durango. below level of eyes when viewed anteriorly; prono- tum bearing small lateral tubercles arising from from lateral margins of anterior lobe (Fig. 126). Male genitalia with right paramere U-shaped, dis- tal arm moderately long (Fig. 128), distal section slightly expanded, bearing an obtusely angled flange on inner margin, tip rounded (Fig. 130); left para mere small, weakly crescentic (Fig. 131); pygophore with caudal tip well produced, apex blunt, forming aright angled notch on right side (Fig. 132); aedeagus relatively short, distal section not recurved, tip acu- minate, lying within an inflated phallotheca. Comparative notes. Ochterus rotundus is easily recog- nized within the Mesoamerican ochterid biota by Figs 128–132. Ochterus rotundus J. Polhemus & M. Pol- the character combination of a rotund and broadly hemus, male genitalic structures – 128, right paramere; ovate body form (Fig. 126); a brilliant green frons 129, right paramere, inner view of apex; 130, detail of with purple overtones (Fig. 127); broadly orange- terminal section of right paramere; 131, left paramere; brown hemelytral margins that each bear 6 promi- 132, apex of pygophore. Specimen from Mexico, nent transverse lavender flecks along their length Durango, 7 mi. W of Los Bancos. (Fig. 126); the structure of the male genital capsule, with the posterior apex of of the pygophore truncate & M. S. Polhemus (on yellow paper)/C. J. Drake (on (Fig. 132); and the distal section of the right param- white paper)”. The specimen is in excellent condition ere expanded, flattened, and paddle-like, with several and retains all its appendages; it has not been dissected. small lateral flanges (Figs 128–130). The male para- The locality label on the holotype contains an error, mere is less tightly curved basally than that in many since the seep locality west of Los Bancos lies in the other Mesoamerican species, and the broad flange on Mexican state of Durango (as was correctly indicated in the inner margin serves to separate it from the simi- the original description by Polhemus & Polhemus lar O. pseudorotundus (compare Figs 128–130 to 1976), rather than in Sonora. This locality was origi- 121, 122) (see additional discussion above). nally visited by J. T. Polhemus and M. S. Polhemus in We have examined the holotype of this species, a April 1964, with M. S. Polhemus subsequently return- brachypterous male held in the Smithsonian Institution. ing there on his own in April 1974. The collection log The specimen bears the labels: “MEX Sonora, 7 mi. W books held in J. T. Polhemus collection make it clear of Los Bancos, IV-26, 1974, M. S. Polhemus (on white that on this second trip M. S. Polhemus was on the paper)/Holotype, Ochterus rotundus, J. T. Polhemus road from Mazatlan to Durango on the date indicated, Downloaded from Brill.com10/08/2021 06:09:55PM via free access
115 110 105 100 95 90 85 80 75 70 65 60
35 35
30 30
25 25
20 20
Ochterus rotundus 15 15
Ochterus smaragdinus 10 10
5 5 SCALE 0 100 200 300 400 500 600 MILES 0 200 400 600 800 KILOMETERS CONIC PROJECTION
110 105 100 95 90 85 80 75 70 65 WEST LONGITUDE
Fig. 133. Distribution of Ochterus species in Mesoamerica. ● O. rotundus J. Polhemus & M. Polhemus; ■︎ O. smaragdinus n. sp. and therefore could not have been in Sonora. The type bedrock faces. It has not been found on horizontal locality is therefore amended, and a note has been alluvial substrates along the margins of ponds or added to the tray holding the type specimen in the streams. Although O. rotundus occurs in the United Smithsonian Institution collections. States only in the bottom of the Grand Canyon, in The Arizona record for O. viridifrons listed by Mexico it is by contrast a species of higher elevations, Schell (1943) in fact refers to this species, which was indicating that it may have a preference for a particu- undescribed at the time of her monograph. lar microclimatic regime that it tracks along an eleva- Distribution. This species occurs primarily in Mexico, tional gradient from north to south. with collections at hand from the states of Durango, Jalisco, Michoacan, Nayarit, Sinaloa, and Sonora Ochterus schellae Drake (Fig. 133). The range of O. rotundus extends barely Figs 110, 134–139 into the United States along the Colorado River in Arizona, with records from this river and its tributar- Ochterus schellae Drake 1952: 73. ies in the bottom of the Grand Canyon (Polhemus & Polhemus 1988). Material examined. Guatemala, Sacatepequéz: 1 ♂, 1 ♀, In the original description of this species, S of Sololá, small waterfalls and wet rock faces [Panajachel J. Polhemus & M. Polhemus (1976) listed two series cascades, 3 mi. S of Solola, 14°45'25"N, 91°09'43"W, consisting only of female specimens that were not elev. 1770 m], 13 January 1970, CL 1324, J. T. Polhemus designated as paratypes, the first from near (USNM ex JTPC). Mexico, Mexico: 3 ♂, 1 ♀, Tejupilco Mismaloya, Jalisco, and the second from west of Dist., Temescaltepec, 1340 m, June–July 1933, H. E. Compostela, Nayarit. A re-examination of these Hinton (holotype and paratypes, USNM); 1 ♂, Temescal specimens indicates that they probably represent tepec, 11 January 1933, H. E. Hinton and R. L. Usinger females of O. schellae, and they are provisionally (USNM ex JTPC). listed under that species in the present work. In addition to the above, the following series that con- Ecological notes. This species is apparently an obligate tain females only also appear to belong to this species, and inhabitant of seeping rheocrenes flowing over vertical are provisionally assigned here: Downloaded from Brill.com10/08/2021 06:09:55PM via free access
El Salvador, Chalatenango: 1 ♀, La Majadita [N of Metapan nr. Cerro Montechristo, vic. 14°23'17"N, 89°23'33"W, elev. ~ 1455 m], 21 December 1969, CL 1257, J. T. Polhemus (USNM ex JTPC). Guatemala, Baja Verapaz: 1 ♀, Dept. Baja V. P. [Baja Verapaz], Los Encuentros, 25 June 1966, Flint and Ortiz (USNM). Mexico, Jalisco: 12 ♀, Atentique, CL 740, 12 June 1975, J. T. Polhemus (USNM ex JTPC); 1 ♀, 10 mi. N of Barra de Navidad, 12 April 1977, Davis and Hanson (USNM ex JTPC). Sinaloa: 1 ♀, rocky stream crossing Hwy. 40, 1 mi. E of Santa Lucia [vic. 23°26'38"N, 105°50'21"W, elev. ~1230 m], 20 April 1964, CL 1019, J. T. & M. S. Polhemus (USNM ex JTPC); 4 ♀, stream flowing over granite at Santa Lucia, seeps and pools, 4000 ft. [1220 m], 6 June 1975, CL 720, D. A. & J. T. Polhemus (USNM ex JTPC).
Diagnosis. Body form broadly ovate, robust; male body length 5.20–5.50, maximum width (across hemelytra) 2.85–2.90; female body length body length 6.00–6.70, maximum width (across hemely- tra) 3.30–3.70. General coloration dark brown with scattered yellowish and orange markings, hemelytra flecked with bluish to lavender pruinose patches (Fig. 134); frons shining black overlain with metallic green, trending to metallic blue on vertex, clypeus and labrum yellowish brown (Fig. 135), eyes silvery, remainder of head black except for narrowly orange adjacent to inner margins of eyes; pronotum with Fig. 134. Ochterus schellae Drake, dorsal habitus. anterior section dull black, lateral flanges pale yel- Female specimen from Guatemala, Sacatepequéz, near low with their outer margins broadly blackish Sololá, CL 1824. brown, posterior section dull blackish brown with extreme posterior margin narrowly orange, 4 small transverse bluish pruinose patches present along channels, these channels bearing transverse striations transverse midline behind calli; mesoscutum and along their lengths, upper portion of frons with a scutellum black, with small bluish pruinose patches slightly raised carina along longitudinal midline; at basal angles, as a small roughly circular patch vertex not sunken below eyes when viewed anteri- medially on anterior margin, and another similar orly (Fig. 135); pronotum bearing lateral tubercles; patch immediately basal to posterior apex, extreme posterolateral pronotal angles acutely rounded, not posterior apical angle sordid white; hemelytra dark pointed; dorsal surface bearing numerous short, blackish brown, grading to orange brown on the stout, pale setae (Fig. 134); posterior pronotum, cla- costal margins, claval commissure narrowly orange, vus and corium bearing numerous small puncta- another small orange marking present on costal tions (Fig. 134). margin at base of wing membrane, pale bluish to Male genitalia with right paramere U-shaped, dis- lavender pruinose patches present on each hemely- tal arm relatively long, slender basally (Fig. 136), api- tron consisting of a large bluish pruinose patch at cal section broadly expanded, paddle-shaped, with a base of clavus followed by a another patch on inner low, broadly rounded flange on outer margin, apex central clavus, 3–4 small pale bluish pruinose broadly rounded (Figs 137, 138); left paramere small, patches scattered on the basal corium, diffuse pale angled, apex blunt (Fig. 139); pygophore with caudal blue pruinosity to either side of basal half of claval tip moderately produced, apex broadly rounded; suture, a large bluish pruinose patch on inner proctiger elongate, tapering, roughly triangular; corium adjacent to claval apex, 4–5 lavender pru- aedeagus relatively short, distal section not recurved, inose patches somewhat evenly spaced along costal tip acuminate, lying within an inflated phallotheca. margin, forewing membrane black to dark brown, Comparative notes. This species is not common in mottled with pale bluish pruinosity, veins rusty collections, but is readily recognized by its relatively brown (Fig. 134). Frons with coarse transverse stria- large size within the Mesoamerican ochterid assem- tions, inner eye margins bordered by broad, shallow blage and its sexual dimorphism in body length, with Downloaded from Brill.com10/08/2021 06:09:55PM via free access
Fig. 135. Ochterus schellae Drake, head, anterior view. Female specimen from Guatemala, Sacatepequéz, near Sololá, CL 1824. males always exceeding 5.0 mm, and female lengths 6.0 mm or greater. The robust body form, the somber, dusky coloration without bright pruinose dorsal markings (Fig. 134), and the large, twisted, paddle-shaped male right paramere (Figs 136–138) are also diagnostic. Drake (1952) described this species based on 6 specimens taken at Tejupilco, Mexico. We have examined the holotype, a brachypterous male held in the Smithsonian Institution. The specimen bears the labels: “MEXICO, June-July 1933, H. E. Hinton (on white paper)/Tejupilco Dist., of Temescaltepec, Alt. 1340 m. (on white paper)/Holotype, Ochterus schellae (on red paper)”. The specimen is in moder- Figs 136–139. Ochterus schellae Drake, male genitalic ately good condition, with all appendages present structures – 136, right paramere; 137, right paramere, but the left forewing somewhat detached and askew; inner view of apex; 138, detail of terminal section of the genitalia have been dissected and are in a vial right paramere; 139, left paramere. Specimen from below the specimen on the same pin. Our illustra- Mexico, Temescaltepec. tion of the male genitalia is taken from a topotypic paratype specimen. as currently interpreted, although confirmation of Schell (1943) misinterpreted this species as some series will require subsequent collection of O. acutangulus Champion, and illustrated the male males from the localities in question. genitalia as belonging to that species. Drake (1952) Distribution. Of sporadic but widespread occurrence noted this error, but did not provide an alternative in Mexico, with its range extending south into illustration of the male genitalia for O. acutangulus. Guatemala and El Salvador (Fig. 110). We now provide proper illustrations for this charac- Ecological notes. Ochterus schellae is a hygropetric ter system in both species (compare Figs 136–139 species that occurs on vertical seeping rock faces. and Figs 18–21). Because many of the series in hand consist of Ochterus shepardi n. sp. female specimens only, it is possible that more than Figs 140–146 one species is represented in the material listed above and shown on the map in Fig. 110. Based on general Material examined. Holotype, male: Costa Rica, Alajuela, size and coloration, however, all of this material La Fortuna, Querado Burio, 15 January 2000, WDS-A- appears to fall within the species concept of O. schellae 1274, W. D. Shepard (USNM ex JTPC). Paratypes: Costa Downloaded from Brill.com10/08/2021 06:09:55PM via free access
Fig. 141. Ochterus shepardi n. sp., head, anterior view. Male specimen from Costa Rica, Alajuela, La Fortuna. pruinose patches present at inner angles of lateral flanges adjoining lateral tubercles, and as a pair (1+1) centrally to either side of longitudinal midline. Scutellum including mesoscutum dull blackish brown, with basolateral angles, small mark centrally at middle of anterior margin, and subapical angular projections on either side of longitudinal midline Fig. 140. Ochterus shepardi n. sp., dorsal habitus. Male before posterior apex pale bluish pruinose. Hemelytra specimen from Costa Rica, Alajuela, La Fortuna. dull medium brown; embolium bearing 3 evenly spaced very small lavender pruinose patches along its length, with middle patch slightly larger than others; Rica, Alajuela: 7 ♂, 5 ♀, same data as holotype (USNM ex clavus bearing very small lavender pruinose patches JTPC, BPBM, EMEC); 1 ♂, Rio Chachagua at Chachagua, at base and centrally along inner margin adjoining 16 January 2000, WDS-A-1276 W. D. Shepard (EMEC). scutellum; basal corium with 3 scattered small laven- Guanacaste: 2 ♀, Rio Lajas, 6 km S of San Miguel, der pruinose patches, central corium with a larger 23 January 2000, WDS-A-1290, W. D. Shepard lavender pruinose patch adjacent to claval apex, dis- (EMEC); 2 ♂, 3 ♀, unnamed stream at Rincon de la Vieja tal corium with a single small lavender pruinose Lodge, 19 January 2000, WDS-A-1286, W. D. Shepard patch at base of wing membrane; wing membrane (EMEC); 2 ♂, 3 ♀, Liberia, 27 March 1987, J. M. Hill dark blackish brown with medium brown veins, (USNM). bearing scattered small lavender pruinose patches across closed cells, mottled with lavender pruinose Description posterior to the closed cells (Fig. 140); thoracic Male. General form elongate ovate, body length 4.80– venter dull black extensively overlain by blue- 5.00 maximum width (across hemelytra) 2.40–2.80. grey pruinosity, posterior margins of acetabulae Color. General coloration brown sparingly flecked pale yellowish, abdominal venter blackish brown; with small pale lavender pruinose patches, lateral legs yellowish brown, ventral surfaces of femora flanges of pronotum dark yellow (Fig. 140); head yellowish. with vertex dull dark blackish brown with very small Structural characters. Head — Length (along midline bluish pruinose patches at each posterolateral angle, as measured from directly above)/width (across eyes) eyes silvery, frons shining metallic green below, shin- = 0.40/1.25, angled downward at greater than 45° ing black above, clypeus orange-yellow, labrum when viewed laterally; frons with coarse transverse golden (Fig. 141), rostrum orange-brown, antennae striations centrally, inner margins of eyes bordered with all segments medium brown. Pronotum dull by narrow channels, these channels bearing numer- blackish brown on anterior half, dull medium brown ous transverse striations (Fig. 141); eyes large, pro- on posterior half, lateral flanges orange-brown nar- truding, not projecting posterolaterally beyond rowly margined with dark brown, small, pale, bluish vertex, upper margins equal with vertex when viewed Downloaded from Brill.com10/08/2021 06:09:55PM via free access
Ochterus smaragdinus n. sp. Figs 133, 147–153
Type material. Holotype, macropterous male: Mexico, Chiapas: 1 ♂, 3 ♀, swift mountain river, 7 mi. N of Santa Fe [vic. 17°25'56"N, 93°02'05"W, elev. ~150 m], 5 May 1964, CL 1102, J. T. & M. S. Polhemus (USNM ex JTPC). Paratypes: Mexico, Chiapas: 3 ♀, same data as holotype (USNM ex JTPC, BPBM); 1 ♀, 3 mi. S of Solusuchiapa, 15 June 1965, Burke, Meyer and Schaffner (TAM). Veracruz: 4 ♂, 6 ♀, 5 km. from Catemaco, 9 May Fig. 147. Ochterus smaragdinus n. sp., dorsal habitus. 1981, J. Bueno (UNAM); 5 ♂, 2 ♀, hydroelectric dam nr. Female specimen from Mexico, Chiapas, 7 mi. N of Catemaco, 9 May 1981, G. Ortega León (UNAM). Santa Fe, CL 1102. Downloaded from Brill.com10/08/2021 06:09:55PM via free access
rising above level of vertex when viewed anteriorly; antennae with segments I and II short, globose, seg- ments III and IV slender, filiform, lengths of seg- ments I–IV = 0.10, 0.10, 0.25, 0.15; rostrum long, length = 2.20, exceeding hind coxae and extending onto base of abdominal venter. Pronotum — Pronotal length (midline)/width = 0.90/2.60, anterior lobe with small, angular lateral tuber- cles, entire surface set with scattered short, recumbent, golden setae, bearing numerous shallow, glabrous foveae; anterior collar not strongly demarcated; anterolateral margins explanate, well demarcated from disk; calli weakly tumescent; posterior margin multisinuate, poste- riorly concave centrally above base of scutellum. Scutellum — Triangular, length/width = 0.90/1.40, bearing scattered short, recumbent, golden setae, mesoscutum slightly transversely raised along poste- rior margin, posterior portion of scutellum bearing weakly suggested sulcus along longitudinal midline. Fig. 148. Ochterus smaragdinus n. sp., head, anterior Hemelytra — Long, attaining tip of abdomen, view. Female specimen from Mexico, Chiapas, 7 mi. N with corium, clavus and embolium well defined, sur- of Santa Fe, CL 1102. faces set with scattered short, recumbent, golden setae; clavus, corium and embolar margin lacking posterior apex; hemelytra dark brown overlain with evident punctations; anterolateral embolar margin emerald green overtones on clavus, outer corium, narrowly produced and explanate; length of clavus and basal membrane, costal margin, inner claval along outside margin 1.90; membrane venation well margin adjacent to scutellum, and claval commissure defined, with 7 evident closed cells. orange-brown; pale bluish to lavender pruinose Ventral surface — Venter of thorax smooth and patches present on each hemelytron consisting of dull, lacking evident setae; scattered tiny, deep, dark extensive pale bluish pruinose maculations on basal punctations present on pro-, meso-, and metepi- half of clavus followed by a another small pale bluish sterna; abdominal venter densely covered with short, patch near claval apex, scattered small pale bluish fine, appressed pale setae, this setal covering inter- pruinose patches flecked on the basal corium, a large rupted by ovate glabrous patches surrounding spira- bluish pruinose patch on inner corium adjacent to cles laterally on paratergites. claval apex, 3 roughly circular lavender pruinose Legs — All leg segments covered by short, fine, patches somewhat evenly spaced on embolium along recumbent pale setae; all coxae, trochanters and basal costal margin, 2 large lavender patches on distal portions of femora ventrally bearing long, semi- corium adjacent to costal margin; forewing mem- erect, slender, pale setae, remainder of ventral femo- brane dark brown, flecked with multiple small pale ral margins bearing moderate length, slender, pale bluish patches; veins medium brown (Fig. 147); tho- semi-erect setae; hind femur slightly bowed down- racic venter dull black slightly frosted with greyish ward when viewed laterally; all tibiae bearing numer- pruinosity, acetabulae and posterior margins meta- ous short, bristly golden setae, these setae becoming pleurae yellowish brown; abdominal venter medium more numerous distally, interspersed with scattered brown, genital segments dark yellow; legs with coxae, short, stout, golden spines, posterior margin of hind trochanters, and basal portions of femora straw yel- tibia with thick fringe of long, golden hairs, lengths low, distal apices of femora, all tibiae and basal tarsal of these hairs over 3× the thickness of the tibia; claws segments pale brown, terminal tarsal segments and golden, gently curving, arolia moderately long, hair- claws dark brown. like, not exceeding length of claws. Lengths of leg Structural characters. Head — Length (along midline segments as follows: fore femur/tibia/tarsal 1/tarsal 2 as measured from directly above)/width (across eyes) = 1.20/1.15/0.05/0.25; middle femur/tibia/tarsal 1/ = 0.40/1.50, angled downward at greater than 45° tarsal 2 = 1.40/1.20/0.07/0.25; hind femur/tibia/ when viewed laterally; frons with fine transverse stri- tarsal 1/tarsal 2/tarsal 3 = 1.70/2.10/0.10/0.20/0.20. ations, becoming more obscure on upper half, inner Male genitalia — Right paramere U-shaped, distal margins of eyes bordered by broad channels cen- arm relatively short, stout (Fig. 149), apical section trally, these channels bearing transverse striations expanded, with a projecting angular spur on inner (Fig. 148); eyes large, protruding, not projecting margin, apex acutely rounded (Fig. 150); left para posterolaterally beyond vertex, upper margins not mere small, angled basally, apex slightly bifurcate Downloaded from Brill.com10/08/2021 06:09:55PM via free access
(Fig. 151); pygophore with medial process moder- ately produced, apex truncate, with angled shoulder laterally (Fig. 153); aedeagus with distal flagellum long, tip slender and acuminate, contained within a moderately large phallotheca (Fig. 152). Female. Similar to male in general structure, color- ation, and body proportions; body length 5.20–5.40, maximum width (across hemelytra) 2.90–3.00. Etymology. The name ‘smaragdinus’ is derived from the Latin ‘smaragdus’, or emerald-colored, and refers to the greenish dorsal coloration of this species. Comparative notes. Ochterus smaragdinus is an uncommonly encountered species found in south- eastern Mexico. Similar to O. panamensis n. sp., to which it appears to be related, it exhibits distinct green overtones over the dark ground color on the pronotum and hemelytra (Fig. 147), these being the only two Ochterus species in Mesoamerica to possess such coloration. It may be separated from O. pana- mensis by the structure of the male right paramere with its expanded distal section bearing an angled spur on the inner margin (Figs 149, 150), and by the blunt rather than angled shape of male pygophore (Fig. 153). Distribution. Known only from southern Mexico (Chiapas and Veracruz) (Fig. 133). Ecological notes. This species occurs along swift, rocky upland streams.
Ochterus viridifrons (Champion) Figs 1, 15, 154–159
Pelogonus viridifrons Champion 1901: 346. Ochterus viridifrons: Schell 1943: 37.
Material examined. Costa Rica, San José: 1 ♂, San Jose, June–July 1931, H. Schmidt (USNM ex JTPC); 2 ♂, 1 ♀, San Jose, 30 June 1957, F. Truxal and A. S. Menke (USNM). Guatemala, Baja Verapaz: 1 ♂, San Geronimo [San Jerónimo], G. C. Champion (BMNH). San Marco: 1 ♂, Rio Naranjo, 450 ft., G. C. Champion (syntype, BMNH). Honduras, Choluteca: 2 ♀, 10 mi. S of Choluteca, stream with waterfall and cascades [13°11'22"N, 87°08'46"W, elev. 35 m], 22 December 1969, CL 1260, J. T. Polhemus (USNM ex JTPC). Mexico, Chiapas: 2 ♀, 19 mi. N of Arriaga, small rocky stream with sandy bed [vic. 16°26'59"N, 93°49'27"W, elev. ~640 m], 1 May 1964, CL 1076, J. T. & M. S. Polhemus (USNM ex JTPC). Guerrero: Figs 149–153. Ochterus smaragdinus n. sp., male geni- 14 ♀, Puente Terreros at Km 21 on Mexico highway 134, talic structures – 149, right paramere; 150, detail of NE of Ixtapa, vic. 17°47'44"N, 101°27'20"W, 395 m terminal section of right paramere; 151, left paramere; [1290 ft.], 29 January 1985, CL 1896, J. T. Polhemus 152, aedeagus; 153, apex of pygophore. Specimen from (USNM). Oaxaca: 8 ♂, 1 ♀, rocky stream at Hwy. 190, Mexico, Chiapas, 7 mi. N of Santa Fe, CL 1102. 29 mi. W of Tequisistlan [vic. 16°30'24"N, 94°54'27"W, elev. ~825 m], 30 April 1964, CL 1064, J. T. & M. S. Polhemus (USNM ex JTPC). Panama, Canal Zone: 4 ♂, 3 ♀, Canal Zone, 6–8 February 1939, C. J. Drake (USNM). Downloaded from Brill.com10/08/2021 06:09:55PM via free access
Fig. 155. Ochterus viridifrons (Champion), head, ante- rior view. Female specimen from Costa Rica, San Jose.
present on costal margin at base of wing membrane, pale bluish to lavender pruinose patches present on each hemelytron consisting of a moderately large bluish pruinose patch at base of clavus followed by another patch on inner central clavus and an elon- gate patch on outside basal margin along claval Fig. 154. Ochterus viridifrons (Champion), dorsal hab- suture, 3–4 small pale bluish pruinose patches scat- itus. Female specimen from Costa Rica, San Jose. tered on the basal corium, a large bluish pruinose patch on inner corium adjacent to claval apex, 4–5 lavender pruinose patches somewhat evenly spaced Diagnosis. Body form elongate ovate; male body along costal margin, forewing membrane black, length 4.80–5.00, maximum width (across hemely- mottled with pale bluish pruinosity, veins brown tra) 2.80–2.90; female body length body length (Figs 1, 154). Frons with coarse transverse striations, 5.20–5.50, maximum width (across hemelytra) inner eye margins bordered by moderately broad, 2.90–3.00. shallow channels, these channels bearing transverse General coloration dark greyish black with scat- striations along their lengths, upper portion of frons tered orange-brown markings, hemelytra flecked with a slightly raised carina along longitudinal mid- with bluish pruinose patches (Figs 1, 154); frons line (Fig. 155); vertex slightly sunken below eyes shining black overlain with brilliant metallic green, when viewed anteriorly; pronotum bearing lateral clypeus and labrum yellowish brown (Fig. 155), eyes tubercles; posterolateral pronotal angles slightly gold, remainder of head black except for narrowly pointed (Figs 1, 154); dorsal surface bearing scat- orange-brown adjacent to inner margins of eyes, pos- tered very short, stout, pale setae (Figs 1, 154); pos- terolateral angles with small bluish pruinose patches; terior pronotum, basal clavus, inner corium bearing pronotum dull black, lateral flanges orange-brown small punctations (Figs 1, 154). with their outer margins broadly blackish brown, Male genitalia with right paramere U-shaped, extreme posterior margin narrowly orange-brown, 4 distal arm relatively short, stout (Fig. 156), apical small transverse bluish pruinose patches present section highly expanded, with a broadly rounded along transverse midline behind calli; mesoscutum flange on inner margin, apex acutely rounded and scutellum black, with small bluish pruinose (Fig. 157); left paramere small, angled basally, apex patches at basal angles, as a small patch medially on narrowed (Fig. 158); pygophore with apex moder- anterior margin, and at extreme posterior apex; hem- ately produced, broadly rounded; aedeagus with elytra dark greyish black, grading to orange-brown distal flagellum relatively short, less than 4× length of on the costal margins, claval commissure narrowly phallobase, tip slender and bifurcate, contained orange-brown, another small orange-brown marking within an elongate, expanded phallotheca (Fig. 15).
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do not approach the prominently angled condition seen in O. acutangulus (Fig. 16) or O. shepardi (Fig. 140), with which it might potentially be con- fused on the basis of external characters and overlap- ping geographic range. The second author examined a syntype male of this species, held in the Natural History Museum, London. The specimen is in good condition, with rather pale coloration, indicating that it may be ten- eral. It bears the following labels: “Type (round label)/ Rio Naranjo, 450 ft., Champion/B. C. A. Rhyn. II, Pelogonus viridifrons, male, Ch./Sp. Figured.” Distribution. Originally described from two speci- mens taken in Guatemala, a male from Rio Naranjo and a female from San Gerónimo, with subsequent records by Schell (1943) from Panama and Costa Rica, and Cianferoni (2012) from Ecuador. We now provide further records from Honduras and Mexico (Chiapas, Guerrero, Oaxaca) (Fig. 159). A specimen from the Grand Canyon, in the U. S. state of Arizona, noted by Schell (1943) as possibly belong- ing to this species, is in fact referable to the subse- quently described O. rotundus J. Polhemus & M. Polhemus (1976). Ecological notes. This is a species that occurs on hori- zontal littoral substrates. Drake (1952) noted that O. viridifrons may sometimes be found inhabiting the margins of brackish lagoons and marshes along sea- coasts in Panama and Mexico, indicating a potential degree of salt tolerance.
Discussion Of the 20 species treated in this work, only two, O. americanus and O. banksi, are confined to North America above Mexico, occurring in the central and eastern United States. These species both share a common putative synapomorphy, in the form of an arcuate flange along the outer margin of the male right paramere, a character state otherwise seen only in the widespread O. perbosci, which also occurs in some of the southern and western United States, as well as widely through Mesoamerica, into South America, and on certain Caribbean islands. In the Figs 156–158. Ochterus viridifrons (Champion), male west, another predominantly North American spe- genitalic structures – 156, right paramere; 157, detail of cies, O. barberi, occurs in the deserts of the south- terminal section of right paramere; 158, left paramere. western United States, extending into Mexico only Specimen from Mexico, Oaxaca, 29 mi. W of Tequisist- in the northern desert state of Sonora. The final spe- lan, CL 1064. cies occurring in the United States north of Mexico is O. rotundus, which is primarily a species of the Comparative notes. Ochterus viridifrons is a widely Sierra Madre Occidental of western Mexico, extend- distributed but only intermittently collected species. ing northward into the United States only in the It may be recognized by the brilliant green coloration thermal refugium of the Grand Canyon of the of the frons (Fig. 155), and the distinctive shape of Colorado River in Arizona. the male right paramere (Figs 156, 157). The lateral Mesoamerica, by contrast, contains a more diverse pronotal angles are slightly pointed (Figs 1, 154), but and predominantly endemic ochterid biota. Three
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115 110 105 100 95 90 85 80 75 70 65 60
35 35
30 30
25 25
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Ochterus viridifrons 5 5 SCALE 0 100 200 300 400 500 600 MILES 0 200 400 600 800 KILOMETERS CONIC PROJECTION
110 105 100 95 90 85 80 75 70 65 WEST LONGITUDE
Fig. 159. Distribution of Ochterus viridifrons (Champion) in Mesoamerica. species occurring in this region, O. aeneifrons, Acknowledgements O. manni, and O. viridifrons, have ranges that The first author dedicates this paper to his late father include various portions of northern and western and posthumous second author, Dr John T. South America, whereas 12 species are confined Polhemus, and also to the late Martin S. Polhemus, entirely to the area between the Mexican border and the brother of the deceased second author and the Panama. There is clear geographic segregation of uncle of the first author. The collaborative field work these Mesomarican endemics, with O. explanatus, O. of the two Polhemus brothers in Mexico during the mexicanus, O. pseudorotundus and O. schellae occur- 1960s led to the discovery of many of the new ring in the sierras of northern and central Mexico, in ochterid species described in this paper, some of some cases as far south as Guatemala; O. acutangulus, which have not been collected since. Although it has O. chiapensis, O. ovatus, and O. smaragdinus occur- taken half a century to finally bring the results of ring predominantly in the highlands of Chiapas, their pioneering work on the aquatic Heteroptera of Guatemala and Belize at the base of the Yucatan Mesoamerica to publication, the first author believes Peninsula; and O. costaricensis, O. obscurus, O. pana- they both would have been pleased with the result. mensis and O. shepardi known only from El Salvador, This monograph is based in large part on speci- Costa Rica and Panama. It is possible that one or mens formerly held in the J. T. Polhemus collection more of the latter species may eventually be shown in Englewood, Colorado, which now resides at the to occur in Colombia as well. Two of the putative United States National Museum of Natural History Mesoamerian endemics, O. explanatus and O. obscu- of the Smithsonian Institution in Washington, rus, are taxa living in wet forest leaf litter, and appear DC (USNM ex JTPC). Thanks are given the follow- to be allied on the basis of their explanate pronotal ing curators who allowed examination of further structure to the two eastern United States endemics, specimens held under their care: Thomas J. Henry O. americanus and O. banksi. The remaining for access to the USNM line collection (USNM), Mesoamerican endemic species show more complex and the C. J. Drake Collection, which also resides at affinities, including apparent sister group relation- the USNM; Dr William Shepard of Sacramento, ships with South American taxa. Deciphering the California, whose specimens are deposited in the Essig origins and affinities of this biota, however, must Museum of Entomology, University of California, await a future phylogenetic analysis. Berkeley (EMEC); DrDownloaded Robert fromBrooks Brill.com10/08/2021 of the Snow 06:09:55PM via free access
Entomological Museum, University of Kansas, Kormilev, N. A. 1973. Ochteridae from Western and Lawrence (SEMK); Dr Richard Shaffner of Texas A Southern Africa (Hemiptera: Heteroptera). – Occasional & M University, College Station (TAMU); and Dr Papers of the California Academy of Sciences 106: Harry Brailovsky, Universidad Nacional Autónoma 1–9. de México, Mexico City (UNAM). Holotypes of the Menke, A. S. 1979. Family Ochteridae/Velvety Shore new species described are deposited in the USNM, Bugs. – In: A. S. Menke (Ed.), The Aquatic and Semi- with paratypes of some species also deposited in the Aquatic Hemiptera of California (Heteroptera: Bishop Museum, Honolulu, Hawaii (BPBM) and at Hemiptera). Bulletin of the California Insect Survey, EMEC. Vol. 21, pp. 124–125. University of California Press, Berkeley, CA, USA. Finally, the first author also thanks the Bishop Moreira, F. F. F., J. F. Barosa, J. R. I. Ribiero and V. P. Alecrim. Museum, Honolulu, Hawaii, which provided 2011. Checklist and distribution of semiaquatic and facilities support, digital imaging technology, and aquatic Heteroptera (Gerromorpha and Nepomorpha) access to collection resources that allowed comple- occurring in Brazil. – Zootaxa 2958: 1–74. tion of this manuscript. That institution's con- Naranjo, C., S. M. Riviaux, F. F. F. Moreira and R. C. tinuing support of research into the taxonomy and Court. 2010. Taxonomy and distribution of aquatic biogeography of aquatic Heteroptera is deeply and semiaquatic Heteroptera (Insecta) from Cuba. – appreciated. Revista de Biologia Tropical (International Journal of Tropical Biology) 58: 897–907. Nieser, N. 1975. The water bugs (Heteroptera: Nepomorpha) of the Guyana region. – Studies of the Fauna of Suriname References and other Guyanas 16: 1–308. Baehr, M. 1990a. Ochterus noualhieri sp. n., a new ochterid Nieser, N. and P.-P. Chen. 1992. 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A new Ochterus (Hemiptera: Heteroptera: Ochteridae and Gelastocoridae. – The Raffles Bulletin Ochteridae) from northeastern Brazil, with a key to the of Zoology, 60: 343–359. speecies recorded from the country. – Zootaxa 3860 Polhemus, J. T. and M. S. Polhemus. 1976. Aquatic and (5): 493–497. semiaquatic Heteroptera of the Grand Canyon (Insecta: Drake, C. J. 1952. Concerning American Ochteridae Hemiptera). – Great Basin Naturalist, 36: 221–226. (Hemiptera). – Florida Entomologist, 35: 72–75. Schell, D. V. 1943. The Ochteridae (Hemiptera) of the Gapud, V. P. & H. O. San Valentin, 1977. The Ochteridae Western Hemisphere. – Journal of the Kansas (Hemiptera) of the Philippines. – Kalikasan, Philippine Entomological Society, 16: 29–47. Journal of Biology, 6: 269–300. Uhler, P.R. 1884. Order VI.–Hemiptera. – In: J. S. Herrera Madrigal, F. 2014. First record of the genus Kingsley (Ed.), The Standard Natural History, Volume Ocyocherus from Colombia and new records of II, pp. 204–296. S. E. Cassino & Co., Boston, MA, Ochteridae from Costa Rica (Hemiptera: Heteroptera). – USA. Acta Biológica Colombia, 20 (2): 255–257. Kormilev, N. A. 1971. Ochteridae from the Oriental and Received: November 27, 2015 Australian Regions. – Pacific Insects 13: 429–444. Accepted: January 7, 2016
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