Tijdschrift voor Entomologie 157 (2014) 163–171 brill.com/tve

Ocyochterus irmae, a beautiful new species of Ochteridae () from the Northern Andes Dan A. Polhemus & John T. Polhemus

The new species Ocyochterus irmae is described from the Andes Mountains of Colombia and adjacent Ecuador. This species is only the second known in the , and the first in which male specimens have been available for dissection of the genitalic structures. A new record is also provided for the previously described O. victor (Bolivar), constituting only the fourth known specimen of this taxon since the original description in 1879, and the first new collection in over 90 years. Color photographs of the dorsal habitus and anterior head are presented for both O. irmae and O. victor, as well as photographic figures of the male parameres, aedeagus, proctiger and pygophore for O. irmae, representing the first illustrations of the male genitalic structures in this genus. A distribution map for O. irmae and O. victor is also provided. Keywords: Heteroptera, Ochteridae, Ocyochterus, new species. Dan A. Polhemus*, Department of Natural Sciences, Bishop Museum, 1525 Bernice Street, Honolulu, HI 96817, USA. [email protected] John T. Polhemus1, Colorado Entomological Institute, 3115 S. York Street, Englewood, CO 80112, USA. 1Deceased.

Introduction lation to the left, and a phallotheca with a cruci- The genus Ocyochterus as currently interpreted is form basal apparatus and an elongate distal flagel- confined to the Neotropical region and has until lum. In contrast to species of occurring in the present time has held a single species, O. victor the Neotropical region, Ocyochterus species are much (Bolivar), described from Ecuador (Bolivar 1879). larger in size (body length equal to or exceeding We now describe a second species, O. irmae,based 6.5 mm), have a smooth frons with at most incipient on material taken in the Cordillera Occidental of transverse striations and no channels parallel to the Colombia and adjacent northern Ecuador. We also inner margins of the eyes, and possess 10–12 closed provide the first illustrations of the male genitalic cells in the forewing membrane. structures in this genus. Up to the present time, specimens of Ocyochterus As with other genera of Ochteridae, Ocyochterus have been very rare in world collections. The original species possess large eyes, ocelli, slender four-seg- species in the genus, O. victor, has until now been mented antennae with the basal two segments short known from only three specimens, all of them col- and stout and the distal two segments longer and lected in the nineteenth century; we now report one more slender, a long rostrum with segment III much more specimen herein, the first collected in over 90 longer than the others, slender legs with the fore years. By contrast, our new species O. irmae is based femora enlarged, a tarsal formula of 2-2-3, male gen- on 60 specimens taken at multiple sites in Colombia italia with the right paramere greatly enlarged in re- and Ecuador. This appears to be largely the result of

Tijdschrift voor Entomologie 157: 163–171, Figs 1–11. [ISSN 0040-7496]. brill.com/tve © Nederlandse Entomologische Vereniging. Published by Koninklijke Brill NV, Leiden. Published 20 November 2014. DOI 10.1163/22119434-00002045

Downloaded from Brill.com10/06/2021 06:02:24AM * Corresponding author via free access 164 Tijdschrift voor Entomologie, volume 157, 2014 more effective modern collection techniques, such as of forewing with 10–12 closed cells in api- localized pyrethrin fogging, applied to the hygropet- cal three-fourths; large species, body length ric habitats preferred by Ocyochterus species. 6.75–8.20 mm ...... Ocyochterus – Eyes separated from vertex by channels par- allel to their inner margins; membrane of Materials and methods forewing with 4–5 cells in apical three- All measurements in the descriptions below are fourths; smaller species than above, body given in millimeters, and were made using a Wild length 3.30–6.30 mm ...... Ochterus M3Z dissecting microscope equipped with an ocu- lar micrometer. Descriptions of color and setifera- Genus Ocyochterus Drake & Gomez-Menor, tion were taken from dry-pinned specimens. Color 1954 photographs of the dorsal color pattern and anterior head were taken using an AutoMontage digital imag- Key to the species of Ocyochterus Drake & ing system linked to a Leica M165-C dissecting stere- Gomez-Menor omicroscope, with post-processing using Photoshop 1. Dorsal coloration dark reddish brown, with software. a narrow, creamy white fascia running trans- Synonymies under taxon headings are nomenclat- versely across each hemelytron from the pos- ural only. The CL number in the Material Examined terior margin of the embolium to the apex section refers to a set of collection locality coding of the clavus (Fig. 9); pronotum, clavus and numbers used by the Polhemus Collection to cross- inner corium lacking pruinose markings; lat- reference field notes, photographs, and other meta- eral pronotal margins broadly marked with data. Because the specimens dealt with in this pa- pale yellow along their entire lengths (Fig. 9); per were collected prior to the advent of modern bodyformbroadlyovate(Fig.9)...... global positioning systems, certain collecting locali- ...... O. victor (Bolivar) ties have been retroactively geo-referenced by using – Dorsal coloration black with metallic red data from Web-based geospatial browsers. This lat- overtones, hemelytra lacking a transverse pale itude and longitude data, as well as other notations fascia (Fig. 1); pronotum and hemelytra in- that add clarity to the locations of particular local- cluding clavus and inner corium bearing scat- ities in a modern context where the original label tered large bluish pruinose markings (Fig. 1); data is limited, have been added in brackets. The pre- lateral pronotal margins bearing limited dark fix ‘vic.’ preceding geographic coordinates in brackets yellow on anterior halves only; body form indicates that the specimens were taken in the vicin- elongateovate(Fig.1)...... O. irmae n. sp. ity of the coordinates provided, but not necessarily at that point location. Localities in close proximity Ocyochterus irmae sp. n. are represented by single symbols on the distribution Figs 1–8 maps. The following abbreviations are used for specimen Type material. Holotype, male: Colombia, Valle de depositories: Cauca, Farallones de Cali National Park, seeping bedrock faces along upper Rio Ponce, nr El Topacio, BPBM Bernice P. Bishop Museum, Honolulu, Hawaii, USA. 1550 m, [vic. 3°19 47 N, 76°38 13 W], 27 July BMNH The Natural History Museum, London, 1989, CL 2423, D.A. & J.T. Polhemus (USNM). United Kingdom. Paratypes: Colombia, Valle de Cauca: 18 males, 20 MNHN Muséum National d’Histoire Naturelle, females, same data as holotype (USNM, BPBM); 1 Paris, France. male, road from Saladito to Anchicaya, 500–1000 m, USNM United States National Museum, Smith- 19 August 1967, P. and B. Wygodzinsky (AMNH). sonian Institution, Washington, DC, Ecuador, Manabi: 2 males, 1 female, 38 km E of USA. Porto Viejo, in splash zone at waterfall, 11 May 1975, Ecuador-Peace Corps-Smithsonian Institution Aquatic Survey (USNM). Cotopaxi: 1 female, small waterfall 79 km E of Quevado, 1225 m, 12 Family OCHTERIDAE Kirkaldy, 1906 May 1975, P. Spangler, A. Langley and G. Cohen (USNM); 6 males, 10 females, 66 km E of Quevado, Key to the genera of Ochteridae occurring in the 1325 m, 15 January 1978, P.J. Spangler and J. Western Hemisphere Andersen (USNM). 1. Vertex of head lacking channels parallel to in- ner margins of eyes (Figs 2, 10); membrane

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and bordering inner margins of eyes dull dark brown with small dark yellowish patches at extreme pos- terolateral angles, vertex and upper frons dark metal- lic green (Fig. 2), lower edge of frons with trans- verse dark yellow fascia, this coloration extending centrally to base of clypeus, flanked to either side by a pair (1 + 1) of transverse blackish patches on lateral portions of ventral margin, clypeus, labrum and rostrum golden brown; pronotum matte black, lateral flanges dark yellow on anterior halves, black posteriorly and narrowly along extreme lateral mar- gins, lateral portions of pronotum immediately in- ward of lateral flanges bearing longitudinally elon- gate pale blue pruinose patches, remainder of prono- tum bearing small, widely and irregularly scattered pale bluish pruinose flecks, posterior pronotal mar- gin bearing 3 small dark yellow patches, lying imme- diately inward of each posterolateral angle and me- dially; scutellum matte black with metallic red over- tones, bearing small, widely and irregularly scattered pale bluish pruinose flecks, posterior apex bearing a longitudinally elongate dark yellow mark; heme- lytra matte black with metallic red overtones, bear- ing small dark yellow patches at each basal embo- lar angle and on each inner corium behind and lat- eral to claval apex, each hemelytron bearing mul- tiple scattered large pale bluish pruinose patches, consisting of a large arcuate patch on basal half of clavus, 4 irregular but roughly evenly spaced patches on outer corium adjacent to costal margin, and on inner corium adjacent to claval apex, wing mem- brane matte black, irregularly mottled with multiply branching pale blue pruinose marks (Fig. 1); thoracic venter dull black, bearing large dark yellow patches on anterior halves of lateral pronotal flanges and at basal angles of embolium; abdominal venter includ- ing genital segments dark reddish-brown; legs shin- ing chestnut brown, distal apices of femora, all tibiae and terminal tarsal segments narrowly black, claws golden brown. Structural characters. Head. Length (along mid- line as measured from directly above)/width (across eyes) = 0.90/2.20 mm, angled downward at greater than 45° when viewed laterally; frons and clypeus Figs 1, 2. Ocyochterus irmae n. sp., photographs of dorsal color patterning. – 1, dorsal habitus; 2, anterior smooth, with barely suggested transverse striations, view of head, showing eyes, frons and clypeus. lacking longitudinal medial carina or channels along inner margins of eyes (Fig. 2); eyes large, protrud- ing, not projecting posterolaterally beyond posterior Description margin of vertex, upper margins not rising above Male. General form broadly ovate (Fig. 1), body level of vertex when viewed anteriorly; antennae with length 6.75–7.30 mm, maximum width (across bases segments I and II short, globose, segments III and of hemelytra) 4.00–4.10 mm. IV slender, filiform, lengths of segments I–IV = Colour. General coloration matte black with metal- 0.15, 0.10, 0.40, 0.40 mm; rostrum long, length = lic red overtones, sparingly marked with patches of 3.80 mm, exceeding hind coxae and extending onto dark yellow and pale pruinose blue; head with eyes base of second visible abdominal sternite. reddish brown, portion of head posterior to ocelli

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Figs 3–7. Ocychoterus irmae n. sp., male genitalic structures. – 3, right paramere; 4, left paramere; 5, aedeagus; 6, posteror margin of pygophore; 7, proctiger.

Pronotum. Length (midline)/width = 1.25/3.70 Legs – with all coxae, trochanters, and bases mm, anterior lobe lacking lateral tubercles, surface of femora bearing short, fine, recumbent pale se- lacking evident setae, bearing scattered small, shal- tae interspersed ventrally with scattered longer, low punctations; anterior collar narrow, not strongly semi-erect, slender, pale setae, remainder femora demarcated; anterolateral margins explanate, weakly and all tibiae shining and without short setae; demarcated from disk; calli only weakly tumescent; middle and hind femora slightly bowed down- posterior margin multisinuate, posteriorly concave ward when viewed laterally; all tibiae and tarsal centrally above base of scutellum. Scutellum triangu- segments bearing scattered moderately long black lar, length/width = 1.25/1.90 mm, surface lacking spines intermixed with more numerous very short evident setae, bearing scattered small, shallow punc- black spines; claws large, golden-brown, gently tations; mesoscutum hidden beneath posterior mar- curving, arolia moderately long, hair-like, not ex- gin of pronotum. Hemelytra long, attaining tip of ceeding length of claws. Lengths of leg segments abdomen, with corium, clavus and embolium well as follows: fore femur/tibia/tarsal 1/tarsal 2 = defined, surface lacking evident setae, bearing scat- 1.90/1.70/0.15/0.35 mm; middle femur/tibia/tarsal tered small, shallow punctations; anterolateral em- 1/tarsal 2 = 1.80/1.90/0.20/0.40 mm; hind fe- bolar margin well produced and explanate; length of mur/tibia/tarsal 1/tarsal 2/tarsal 3 = 2.10/3.40/0.15/ clavus along outside margin 3.00 mm; membrane ve- 0.40/0.50 mm. nation well defined, with 10 evident closed cells. Genital segment well retracted into abdomen. Ventral surface of thorax – smooth and dull, bear- Pygophore with caudal tip blunt, broadly rounded, ing scattered very short silvery setae on pleurae; scat- lateral lobes weakly developed (Fig. 6). Proctiger tered tiny, deep, dark punctations present on proepis- roughly triangular, apex forming a blunt angle terna; abdominal venter densely covered with very (Fig. 7). Right paramere slender and very elongate, short, fine, appressed silvery setae, this setal cover- bearing a U-shaped bend basally followed by a long ing interrupted by ovate glabrous patches surround- distal arm, this arm bearing a transverse ridge near ing spiracles laterally on paratergites. midpoint (Fig. 3). Left paramere small, hatchet-

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Fig. 8. Distribution of Ocychoterus irmae n. sp. shaped, basal arm slender, distal head broadly ex- nate lateral projections; distal portion of aedeagus panded (Fig. 4). Aedeagus with shape as in Fig. 5. forming elongate flagellum, this flagellum articulated consisting of cruciform phallobase bearing acumi- at basal point of connection to phallobase, and again

Downloaded from Brill.com10/06/2021 06:02:24AM via free access 168 Tijdschrift voor Entomologie, volume 157, 2014 at a transverse joint near midpoint, extreme distal section very slender and whip-like. Female. Similar to male in general structure, col- oration, and body proportions, but slightly larger in overall size; body length 8.10–8.20 mm, maximum width (across bases of hemelytra) 4.45–4.50 mm. Subgenital plate roughly pentagonal, recessed basally into abdominal ventrite VI, posterorolateral margins converging at approximately 90° angle with apex di- rected posteriorly, extreme apex with small V-shaped notch. Etymology. The name irmae honors Irma Polhemus, the mother of the first author and the wife of the second author. She accompanied both authors on many collecting expeditions to remote parts of the world, and it is a pleasure to honor her spirit of adventure with this patronym. Comparative notes. Ocyochterus irmae is easily distin- guished from the previously described O. victor by its large size (body length exceeding 6.5 mm in males and 8 mm in females); the dorsal color pattern, con- sisting of a matte black ground color with metallic red overtones, marked with numerous pale blue pru- inose patches (Fig. 1); the smooth, metallic green frons, which has only faintly suggested transverse striations, and bears a dark yellow patch along the ventral margin, with this dark yellowish coloration not extending upward adjacent to the inner margins of the eyes (Fig. 2). It is likely that the structures of the male genitalia (Figs 3–7) will also prove diagnos- tic, although these are currently unknown for O. vic- tor (see subsequent discussion). This is the first species of Ocyochterus for which the male genitalia have been dissected and illustrated, and in general they conform to the typical aspects of this family. The pygophore is cup-like and ven- trally keeled, with its posterior apex produced and setiferous (Fig. 6). The proctiger is roughly triangu- lar in shape (Fig. 7), and forms a dorsal lid above the pygophore. The aedeagus has a cruciform basal apparatus, to which is connected a very long dis- tal flagellum; this flagellum has two points of artic- ulation, one basally where it connects to the phal- lobase, and another near the halfway point dis- Figs 9, 10. Ocyochterus victor (Bolivar), photographs of tally, where the flagellum makes a transition from dorsal color patterning. – 9, dorsal habitus; 10, anterior a broadly curving tube of relatively even width to a view of head, showing eyes, frons and clypeus. more complex, coiled distal section contained within a partially membranous phallotheca (Fig. 5). The male parameres differ greatly in size, with the left distal arm on the right paramere and the distal flagel- paramere reduced, possessing a slender basal arm lum of the aedeagus are character states that represent and a transversely expanded distal head (Fig. 4), potential apomorphies for Ocyochterus. whereas the right paramere is extremely elongate, Ecological notes. The Rio Ponce at the El Topacio with a U-shaped bend basally, followed by a slender type locality was a cool, rocky mountain river flow- and weakly sinuate distal arm that tapers gradually ing down from the high peaks of the Farallones de to a rounded apex (Fig. 3). In comparison to other Cali. The river channel above the parking area at Neotropical Ochteridae, the elongation of both the the end of the road was bordered by bedrock seeps

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Fig. 11. Distribution of Ocyochterus victor (Bolivar). overlain by roots and tangles of vegetation that could pyrethrin fog to these wet bedrock faces, causing the be pulled back to reveal the wet rock beneath. The to emerge from hiding places amid the vege- type series of O. irmae was taken by applying a light tative mass. Other taxa also taken in company with

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O. irmae included Paravelia flavomarginata (Hunger- new species described above (compare Figs 1, 2 and ford), as well as other species in the genera Hebrus, Figs 9, 10). In addition, the body form of O. victor Mesovelia, Saldula and Nertha. is more broadly ovate and robust, in comparison to the slightly more elongate and posteriorly tapering Ocyochterus victor (Bolivar) body form exhibited by O. irmae.Wehavenothad Figs 9–11 an opportunity to dissect the male genitalia of O. vic- tor, because the only specimen we have in hand is a Pelogonus victor Bolivar, 1879: 144. female. Pelogonus splendidus Montandon, 1898: 73. Syn. by The species Ochterus splendidulus, originally de- Drake & Gomez-Menor, 1954: 158. scribed by Montandon (1898) in the genus Pelo- Ochterus splendidulus: Schell, 1943: 33. gonus, was synonymized under O. victor by Drake Ochterus victor: Schell, 1943: 46. & Gomez-Menor (1954). Montandon distinguished Ocyochterus victor: Drake & Gomez-Menor, 1954: his O. splendidus on the basis of minor differences 158. in coloration that he felt deviated from Bolivar’s very Material examined. Ecuador, Pichincha: 1 female, brief original description (Bolivar 1879). The male 55 km E of Santo Domingo de los Colorados on holotype of O. victor is in the Instituto Español de Chiriboga road, 1400 m, collection #8, 7 Septem- Entomologia is Madrid. We have not seen this speci- ber 1990, P.J. Spangler (USNM). 1 male, Nanegal, men, but the second author was able to examine the Équateur [vic. 0°07 53 N, 78°40 35 W], coll. V. male holotype of O. splendidulus in the Muséum Na- Ortenada (holotype of O. splendidulus, MNHN); 1 tional d’Histoire Naturelle, Paris, and a female spec- female, same data as preceding (BMNH). imen of O. splendidulus, apparently collected with Diagnosis. Female. General form broadly ovate the holotype (and incorrectly labeled as the type) (Fig. 9), body length 7.20 mm, maximum width held in The Natural History Museum, London. The (across bases of hemelytra) 4.45 mm. Dorsal ground color pattern of these latter specimens conforms to color dark, velvety reddish-black, with broad dark the illustration of the holotype provided by Drake & yellow markings on posterior head, lateral prono- Gomez-Menor (1954), and we agree with their deci- tal margins, and basal half of the costal margin on sion in regard to the synonymy of O. splendidulus.In the hemelytra; posterior section of hemelytra bear- addition to these three specimens, all collected in the ing a narrow, transverse, creamy white fascia begin- nineteenth century, we can now add one additional ning on each hemelytral margin near posterior ter- example listed herein, which represents the first re- minus of embolium and angling slightly forward collection of O. victor in over 90 years. to posterior claval apex (Fig. 9). Frons with upper Distribution. The male holotype of O. victor was and central portions dark, shining metallic blackish- taken at Pichincha, Ecuador (near the northern out- green, with the lateral and lower frons as well as the skirts of modern Quito), while both specimens of eyes yellowish-tan, this yellowish coloration extend- Montandon’s O. splendidulus came from Nanegal, ing parallel to each inner eye margin for its entire Ecuador (approximately 41 km in a straight line length (Fig. 10). Venter dull blackish-brown; legs and northwest of Quito). The specimen reported below rostrum orange-brown. comes from the same province in Ecuador as the Comparative notes. Bolivar’s original (1879) descrip- holotype, and is only the fourth known in world col- tion of this taxon is in Spanish, and consists of a sin- lections. Based on these records, and the very lim- gle paragraph dealing almost exclusively with details ited number of captures over the past 135 years since of the body coloration. Schell (1943) subsequently the original description, it appears that O. victor may provided a translation of Bolivar’s description into have a very localized distribution in the Ecuadorian English, but was unable to examine any specimens. Andes (Fig. 11). Drake & Gomez-Menor (1954) subsequently pro- vided an excellent shaded illustration of the dorsal habitus, clearly depicting the distinctive color pattern Acknowledgements of this species. The combination of a dark reddish- The following individuals kindly provided access to brown dorsal ground color cut by a striking pale fas- collections and specimens held under their care: Dr cia across the posterior hemelytra (Fig. 9), the lat- Thomas J. Henry of the USDA Systematic Ento- eral pronotal margins broadly marked with dark yel- mology Laboratory, who provided access to mate- low along their entire lengths (Fig. 9), and a metallic rial held in the Smithsonian Institution collections; green frons with a dark yellow U-shaped mark that Mme. Dominique Pluot-Sigwalt, Muséum National covers the ventral margin and extends upward on d’Histoire Naturelle; and W.R. Dolling, The Natural each side parallel to the inner eye margins (Fig. 10) History Museum, London. We also wish to thank make this species unmistakable in relation to our Maria del Rosario Manzano, of the Universidad de

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Cali, who facilitated our field work in the Cordillera Drake, C.J. & J. Gomez-Menor, 1954. A new genus of Oriental that led to the discovery of O. irmae.Fi- American Ochteridae (). – Revista Española nally, the second author thanks Dr Allen Allison and de Entomología 30: 157–159. Dr Neal Evenhuis of the Bishop Museum, Honolulu, Montandon, A.-L., 1898. Hémiptères Hétéroptères nou- Hawaii, who provided facilities support and access to veaux des Collections du Muséum de Paris. – Bulletin collection resources that allowed completion of this du Muséum National d’Histoire Naturelle, Paris 4: 72– manuscript. 75. Schell, D.V., 1943. The Ochteridae (Hemiptera) of the Western Hemisphere. – Journal of the Kansas Entomo- References logical Society 16: 29–47. Bolivar, D.I., 1879. Hemípteros Nuevos del Museo de Madrid. – Anales de la Sociedad Española de Historia Natural 8: 130–146. Received: August 15, 2014 Accepted: October 5, 2014

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