TRANSACTIONS RESEARCH ARTICLE TAES 140: 163-184 AMERICAN ISSO 0002-8320 ENTOMOLOGICAL www.aes.entomology.org SOCIETY

Two new species of Neoxabea, three new species of Oecanthus, and documentation of two other species in Nicaragua (: : Oecanthinae)

Nancy Collins, Eric van den Berghe and Lisa Carson

[NC] [email protected] [EVDB] [email protected] (University of the Americas, Managua, Nicaragua) [LMC] [email protected] (Dartmouth College, Hanover, NH, USA)

ABSTRACT

Two new species of Neoxabea and three new species of Oecanthus are described from Nicaragua: N. cerrojesusensis and N. ottei, O. belti, and O. symesi occur at higher elevations in the northern half of Nicaragua. A large population of O. bakeri was found in dry tropical forest west of Lake Nicaragua. The presence of O. allardi in Nicaragua is documented as the first record outside of the West Indies, and waveforms from the first known recordings of a singing maleO. leptogrammus are provided. Keys are provided for Neoxabea and Oecanthus in Central America.

INTRODUCTIONS of forest along waterways, transitioning to cloud Oecanthinae have been described from the forest around 1,100 m. Domitila Reserva Silvestre Americas, Europe, Africa, Asia, India, Australia Privada (DRSP) is located 46 km southeast of San and islands in the Caribbean and Pacific. Eadeset Marcos, Nicaragua. The property extends to the al. 2013 lists 11 species in the genus Neoxabea and western shore of Lake Nicaragua (Lago Cocibolca). 65 species in the genus Oecanthus. All 11 of the Other sites where Oecanthines were Neoxabea species (Table 1) and 23 of the Oecanthus encountered or collected include: Cerro Tisey, species (Table 2) occur in the Western Hemisphere Datanli Diablo, Jinotega, Kahka Creek, Kilambé, south of the United States. Only five species have Montelimar, San Marcos and Tepesomoto. been previously documented in Central America: N. bipunctata, O. comma, O. leptogrammus, O. MATERIALS AND METHODS prolatus and O. varicornis (Walker 1967). Collections by EV and NC from ten sites in Genus verification.—There are nine genera Nicaragua (Fig. 1) yielded two unknown species of of Oecanthinae worldwide, but only two occur in Neoxabea and three unknown species of Oecanthus. the New World – Oecanthus and Neoxabea. Tree The principal collection sites are Cerro Jesus (cloud crickets encountered in Nicaragua were identified forest) and Domitila Reserva Silvestre Privada to genus with a key from Walker (1967). Two of (tropical dry forest). Cerro Jesus (peak elevation the new species were identified as members of the 1,792 m), is northwest of Jalapa, Nicaragua, and genus Neoxabea due to the lack of spines on the straddles the northern border with Honduras. The hind tibiae, and the presence of a tubercle on the top 600 m is covered in dense virgin cloud forest distal border of the scape (Figs. 4 & 12). Three of on the Nicaraguan side, but is clear-cut on the the new species were identified as members of the Honduran side. The lower slopes on the Nicaraguan genus Oecanthus due to the presence of spines on side are cultivated with shade coffee and corridors the hind tibiae (Figs. 20, 26, 34 & 58).

Publication date:15 May 2014 urn:lsid:zoobank.org:pub:5534B48B-DDD3-40E4-AFD3-859F0CD4F0FF 164 NEOXABEA AND OECANTHUS OF NICARAGUA

Collection methods, sites and dates.—Areas including antennae, tegmina, limbs or cerci. The inhabited by Oecanthines at DRSP and Montelimar tegminal width (at the widest portion) and the were determined by locating actively singing males; length to width ratio (l/w) were measured with the tree crickets were collected by hand or with sweep tegmina resting together atop the abdomen. Femur nets. Oecanthines at other collection sites were length was measured from the proximal to distal drawn to sheets set up in conjunction with 120-175 ends of the femur. watt mercury vapor bulbs, and BioQuip model Number of teeth on the stridulatory file.—The 2807A 15 watt blacklight tubes. The collection dates right tegmen was mounted on a slide and viewed for all species ranged from April 2006 to June 2013. using a First Lab microscope with a 10X digital Dates for Cerro Jesus were 28-29 April 2006, 1-6 eyepiece through a 10X power objective lens in June 2011, 17-20 May 2012, and 6-11 June 2013. order to count the stridulatory file teeth. Other collection dates are: Cerro Tisey ca 25 April Metanotal glands.—The metanotal glands of 2011; Datanli Diablo ca 17-21 May 2011; DRSP 2-7 male specimens can be an important taxonomic January 2010 and 20-25 January 2011; Jinotega July character for tree crickets (Walker TJ and Gurney 2013; Kahka Creek 5-11 May 2011; Kilambé 22-25 AB 1967). Since not all species have been May 2011; Montelimar 2-7 January 2010 and 20-25 catalogued by metanotal gland appearance, they January 2011; San Marcos 15 March 2011 and 6-11 are not yet a reliable diagnostic tool. This paper June 2013; and Tepesomoto April 2011. provides photographs for documentation and Specimens relevant to this study were comparisons (Figs. 48-56). Photographs were taken examined in the entomological collections at the using a Canon Powershot S5IS though a First Lab Museo Entomologico in Leon, Nicaragua, and at microscope with a 10X digital eyepiece through a Ave Maria University, San Marcos, Nicaragua. 4X power objective lens. None were identified beyond genus. Tibial armature.— Subapical, dorsal, hind Documentation and preservation of tibial spines on Oecanthus specimens were specimen.— Males and females were collected examined using a First Lab microscope with a 10X and photographed; singing males were video digital eyepiece through a 4X power objective lens. recorded with sound. Measurements and visual Song and temperature recordings.—Songs were observations for morphological study were made recorded using a Canon PowerShot S5 IS digital after the specimens were euthanized using acetone. camera set in video mode at 30 frames/sec, with Specimens were preserved in alcohol or dried using the camera held as close to the singing anhydrous calcium sulfate crystals. Some specimens as possible. An avi-to-wav converter marketed had their forewings removed in order to photograph online by 008soft.com was used to extract the the metanotal gland and to count the teeth on the sound track from the video file. AVS4YOU Audio stridulatory file. Editor Software versions 4.2 and 6.1 were used to Depositories.—Academy of Natural Sciences analyze the sound tracks and to make images of of Philadelphia (ANSP); Bernice Pauahi Bishop their waveforms. The temperatures within 15 cm Museum (BPBM); California Academy of Sciences of actively singing males were recorded using a (CAS); Florida State Collection of hand-held digital thermometer attached to a 3-meter (FSCA); Museo Entomologico, Leon (MEL); and wire cord manufactured by LaCrosse Technology. University of Wisconsin-Madison (UW-Mad). Accuracy of the digital thermometer was confirmed Photographs, audio files and videos of all seven by comparing, in a water bath, its readings with a species discussed in this paper will be made precision glass laboratory thermometer. available as supplementary material for viewing The pulse rate is documented as pulses per at www.oecanthinae.com and http://orthoptera. second (p/s), and is calculated during periods of speciesfile.org. continuous pulse production. It should be noted that Morphological measurements.—The total the tree crickets presented in this paper sang only at length refers to the midline length from the night. Recordings were made during limited visits fastigium to the apex of the subgenital plate--not to Nicaragua since import permits of live specimens NANCY COLLINS ET. AL. 165 into the U.S. for longer observations could not be areas (Figs. 3 & 18). Specimen deposited at ANSP. obtained. Due to consistent night temperatures Song diagnosis - The calling song of males while in locations visited in Nicaragua, the consists of intermittent bursts of rapid trilling. recordings of singing males were made within an This species starts out with staccato-like clicking extremely narrow range of temperatures; therefore, sound for 5-25 seconds, then has 0.9-1.3 seconds no graphs of pulses per second over ranges of of a continuous trill with ascending volume. No temperatures are possible. set patterns of silence, staccato or trilling were Molecular Analysis.—Specimens were identified. Pulse production rate 90 pulses per sequenced at the 12S locus using primers and second (Figs. 7, 8 & 9) and a dominant frequency of conditions from Robillard and Desutter-Grandcolas 2.7 kHz at 26.1 C. (2006). Molecular analysis - The aligned and trimmed 12S of sequences of N. cerrojesusensis [Genbank RESULTS Accession Number (GAN): KJ024364] were 278 base pairs. Neoxabea cerrojesusensis n. sp. Paratypes: one male and one female (Cerro Collins & van den Berghe Jesus, Jun 2013) to: MEL and UW-Mad; one male urn:lsid:zoobank.org:act:A9A7771B-EB7B-4557-931B- and one female (Cerro Jesus, Apr 2006) to: BPBM, D2C273671C04 CAS and FSCA.

Neoxabea cerrojesusensis (Figs. 2 & 3) is Neoxabea ottei n. sp. currently known only from northern Nicaragua. Collins & van den Berghe urn:lsid:zoobank.org:act:C8CCE77F-C462-45B8-B0AB- Etymology.— Named for the mountain where BD7F4DC72720 this new species was discovered. Habitat.—Drawn to lights; host plants/trees Neoxabea ottei (Figs. 10 & 11) is currently unknown. known only from northern Nicaragua. Holotype.—Male. Cerro Jesus, in northern Etymology.—Named in honor of Dr. Daniel Nicaragua. GPS – 13° 59’ N; 86° 11’ W, elevation Otte who has described over 60 species of tree 1,100m. 6 Jun 2013. E. van den Berghe, collr; crickets, and is founder of the Orthoptera Species deposited at ANSP. Body length 15.5 mm; femur File (an online orthoptera database). length 7.5 mm; pronotal length 3.0 mm; tegminal Habitat .—Drawn to lights; host plants/trees length 11.0 mm; tegminal width 4.0 mm. Antennal unknown. tubercles as in Fig. 4. Head color brown; pronotum Holotype.—Male. Cerro Jesus, in northern tan; tegmina translucent tan; limbs translucent Nicaragua. GPS 13° 59’ N; 86° 11’ W, elevation yellow. Cerci curve inward proximally then outward 1,100m. 6 Jun 2013. N. Collins, collr; deposited and end with a pointed tip (Fig. 5). Eight pale at ANSP. Body length 19.5 mm; femur length 8.5 ovoid knobs on the dorsal abdomen - one knob per mm; pronotal length 3.0 mm; tegminal length 11.5 tergum (Fig. 6). Tegmina of the males have a faint mm; tegminal width 5.0 mm. Antennal tubercles X pattern when at rest on the abdomen (Figs. 2 & as in figure 12. Head color brown; pronotum tan; 18). Stridulatory teeth total 23. Metanotal gland at tegmina translucent beige; limbs translucent yellow. figure 48. Cerci curve inward proximally then outward and Female allotype description.—Collected at end with a pointed tip (Fig. 13). Eight pale ovoid Cerro Jesus. Body length 15.0 mm; pronotal length knobs on the dorsal abdomen - one knob per tergum 2.9 mm; ovipositor length 4.8 mm; femur length which are noticeable through the wings at rest (Figs. 7.0 mm. Antennal tubercles same as in male. 14 & 18). Tegmina of the males have dark markings Head color mahogany; pronotum light brown, which form an X pattern of varying clarity when lighter abdominal region; limbs translucent yellow. resting on the abdomen. Stridulatory teeth total 20. Forewings display two dark maculae with whitish Metanotal gland photo at figure 49. 166 NEOXABEA AND OECANTHUS OF NICARAGUA

Female allotype description – Also located at mm with wings at rest atop abdomen. Tegmina Cerro Jesus. Length 18.0 mm; ovipositor 5.0 mm. have scattered darkened areas along veins (Fig. Antennal tubercles same as in male. Medium brown 19). Stridulatory teeth total 25. Dark brown ventral but lighter on the lateral abdominal regions; head markings (Fig. 21) in whitish fields on the straw- and pronotum mahogany. Forewings display four colored scape and pedicel. Color uniformly pale dark maculae – located dorsally (Fig. 11). Specimen green, lighter in the abdominal region. Pale orange- deposited at ANSP. tan head. Limbs translucent milky green color. Song diagnosis - The calling song of males Femoral-tibial joints dark orange with a prominent consists of intermittent bursts of rapid trilling. This black half-circle. Three rows of light black slash species starts out with a staccato-like clicking sound markings down the length of the femur. Front for up to 25 seconds, then has 0.9-1.3 seconds of tibial joints deep golden yellow. Cerci straight and a continuous trill with ascending volume. Pulse slender. Metanotal gland as at figure 52. production rate 94 pulses per second (Figs. 15, 16 & Female allotype description.— Collected at 17) and a dominant frequency of 2.8 kHz at 26.2 C. DRSP. Overall yellowish-green color. Head has Molecular analysis - The aligned and trimmed same orange-tan head and black slash markings on 12S of sequences of N. ottei [GAN: KJ024365] the hind femurs as on the male (Fig. 20). Antennal were 278 base pairs. markings same as on male. Deposited at ANSP. Paratypes: one male and one female (Cerro Immature description.—Fifth stage male Jesus, Jun 2013) to MEL and UW-Mad; one male same in color as adults with same black slashes on and one female (Cerro Jesus, Apr 2006) to BPBM, hind femurs. Developing wings have black slash CAS and FSCA. markings. Younger instars: dorsal abdomen light green with two longitudinal whitish lines bordering Oecanthus bakeri n. sp. a central strip of reddish-brown, with rows of paired Collins & van den Berghe dark grey circles urn:lsid:zoobank.org:act:37032B94-CDA7-444D-B444- Song diagnosis.— The calling song of DC21535C5549 males consists of a rapid trilling - which occurs continuously or as intermittent bursts of trilling Oecanthus bakeri (Figs. 19 & 20) is currently of various lengths (Figs. 23 & 24). Recordings known only from Nicaragua. were made of DRSP individuals in the field and Etymology.—Named in honor of Charles as captives in containers made of netting, with Fuller Baker for his contribution to the study of temperatures ranging from 25.6 C to 27.1 C. Oecanthinae in Nicaragua. Baker described O. Pulse rate at 25.7 C is 80 pulses per second with a rileyi in 1905. He also described O. marcosensis dominant frequency of 3.0 kHz. which was collected near San Marcos, Nicaragua. Tibial armature .—Four large spines in pairs on Further analysis led to listing it as a synonym of O. the distal hind tibiae, with two to four small spines varicornis. between the large spines – all aligned in two rows Habitat.— Individuals were readily located at (lateral and medial edges). Small spines continue DRSP in the month of January on dense patches proximally but do not extend into the femur. Spines of Acanthaceae and Verbena sp. next to a rarely pale with black tips. traveled dirt road between wire-fenced pastureland Paratypes.—one male each (DRSP, Jan 2011) and dry tropical forest. One specimen also collected to MEL, UW-Mad, and FSCA. at lights on Cerro Tisey in 2012. Holotype.—Male. Nicaragua, Domitila Reserva Silvestre Privada - dry tropical forest in southwestern Nicaragua. GPS – 11° 48’ N 86° 31’ W, elevation 9m. 22 Jan 2011. N. J. Collins, collector. Deposited at ANSP. Body length 16.8 mm; tegminal length 12.0 mm; tegminal width 6.0 NANCY COLLINS ET. AL. 167

Oecanthus belti n. sp. Oecanthus symesi n. sp. Collins & van den Berghe Collins & van den Berghe urn:lsid:zoobank.org:act:76B568CB-5D91-4D52-9BF6- urn:lsid:zoobank.org:act:89B0CCE8-C2E5-40AD-AF15- C331F9303792 1796F0C322A9

Oecanthus belti (Figs. 25 & 26) is currently Oecanthus symesi (Figs. 30 & 31) is currently known only from northern Nicaragua. known only from Cerro Jesus, Nicaragua. Etymology.—Named after Thomas Belt, Etymology.—Named in honor of Dr. an amateur naturalist who traveled throughout Laurel Symes. Dr. Symes earned her PhD using Nicaragua while doing engineering work 1868- Oecanthinae as her research study group, and 1872. Belt’s book, The Naturalist in Nicaragua, verified distinct species status for the two newly includes many passages about Nicaraguan . described Nicaraguan Neoxabea tree crickets using Habitat .— Drawn to lights. One female DNA sequencing with LC. found on Asteraceae: Dahlia dallium (Fig. 26) in a Habitat .—One female individual was collected landscaped garden at a coffee farm on Cerro Jesus via net approximately 15 feet high from a Guava (elevation 1,100m). tree aka “Guava roja” (Psidium guajava). Holotype.—Male. Collected at Cerro Jesus, Holotype.— Male. Collected at Cerro Jesus, in northern Nicaragua. GPS 13° 59’ N; 86° 11’ W, in northern Nicaragua. GPS 13° 59’ N; 86° 11’ W, elevation 1,100m. 6 Jun 2013. N. Collins, collr; elevation 1,100m. 6 Jun 2013. E. Van den Berghe, deposited at ANSP. Body length 15.5 mm; femur collr; deposited at ANSP. Body length 16.5 mm; length 9.0 mm; pronotal length 2.5 mm; tegminal femur length 10.0 mm; pronotal length 2.5 mm; length 12.0 mm; tegminal width 5.0 mm. Antennal tegminal length 11.5 mm; tegminal width 3.5 markings as in figure 27. Head color pale green mm. Single deep brown vertical lines on first two and cream; antennae black; scape red and pedicel antennal segments as in figure 32. Color uniformly blackish red; pronotum pale green bordered with rich golden yellow with pale yellow limbs and deep white; tegmina translucent green; limbs translucent orange head and orange pronotum. Cerci slender pale green with orange femoral-tibial joints. and pale with dark pointed distal tips (Fig. 33). Stridulatory teeth total 32. Metanotal gland at figure Mouth palps orange with white distal tips. Front 53. tibial and femoral-tibial joints brownish-orange. Female allotype description.—Collected at Stridulatory teeth total 18. Metanotal gland at figure Cerro Jesus 6 Jun 2013. Length 16.0 mm; ovipositor 54. 5.0 mm. Pale green body, limbs and pronotum. Rich Female allotype description.—Collected at red on head. Antennae black. Appearance similar to Cerro Jesus 6 Jun 2013. Body length 15.5 mm. that of female O. varicornis. Specimen deposited at Ovipositor length 4.5 mm. Antennal markings and ANSP. overall color same as in male. Specimen deposited Song diagnosis .—The calling song of males at ANSP. consists of a rapid trilling (Figs. 28 & 29). Pulse Song diagnosis.—The song pattern matches rate at 26.7 C. is 81 pulses per second with a those of U.S. species in the niveus group - dominant frequency of 4.2 kHz. intermittent bursts of trilling (Figs. 35 & 36). Pulse rate at 22.7 C is 58 pulses per second with a Tibial armature.—Four large spines in pairs on dominant frequency of 2.0 kHz. It is noteworthy the distal hind tibiae, with two to four small spines that while four different species were singing between the large spines – all aligned in two rows from inside separate net containers, including N. (lateral and medial edges). Small spines continue cerrojesusensis, N. ottei, O. belti and O. symesi, proximally but do not extend into the femur. Spines the male O. symesi was noticeably lower in volume pale with black tips. to the human ear than the other three species, Paratype.—Females to: MEL, UW-Mad, CAS presumably due its narrow tegmina. Given the and FSCA. large number of singing insects on Cerro Jesus, and 168 NEOXABEA AND OECANTHUS OF NICARAGUA

the collective volume of their songs, it would be DISCUSSION difficult to locateO. symesi calling high in trees. Tibial armature .—Total of five to six large New species status was determined by spines on the distal end of the hind tibia, not comparing morphological characteristics, locations situated as pairs, with zero to three small spines and song patterns to those of species documented in between – all aligned in two rows (lateral and Central America. Special attention was also given medial). Small spines continue proximally but do to species with similar stridulatory teeth count not extend into the femur. Large spines pale yellow totals, regardless of location, as presented in Walker large spines transitioning to orange and then black TJ (1962) and Walker TJ (1967). Metanotal gland tips; small spines orange with black tips (Fig. 34). features were compared to those in Walker TJ, Molecular analysis.—The aligned and Gurney AB (1967) and Walker TJ (1967). Walker’s trimmed 12S sequence of Oecanthus symesi [GAN: papers can be accessed at: http://entomology.ifas. KJ024362] was 433 base pairs. ufl.edu/walker/buzz/s576a.htm. Paratypes.—One male (Cerro Jesus, Jun 2013) Molecular analysis.—DNA analysis was to CAS; one female (Cerro Jesus, Apr 2006) to used to confirm divergence between the new MEL and one female UWM (Cerro Jesus, Jun Neoxabea species. The 5.0% divergence between 2013). N. cerrojesusensis and N. ottei is comparable to the divergence between each of these species and Documentation of other Oecanthus species in the North American species, N. bipunctata [GAN: Nicaragua KJ024363]. There is 5.8% divergence between N. bipunctata and N. cerrojesusensis; and 4.3% Oecanthus allardi Walker TJ & Gurney AB divergence between N. bipunctata and N. ottei. 1960. Identification based on color, size, antennal DNA analysis of O. symesi revealed 8.7% markings and sound recordings compared to those divergence and two deletions relative to the most posted online at Macaulay Library. Photographs similar North American species O. pini [GAN: and song waveforms (Figs. 37-41) are offered KJ024361]. However, while it is clear that the for documentation of this species outside of sequence from O. symesi is distinct from that of O. the West Indies. Metanotal gland at figure 55. pini, caution should be exercised before inferring Dominant frequency 3.0 kHz at 27.2 C. Collected at that O. symesi is much more closely related to O. Montelimar, DRSP, Kilambé and Cerro Jesus. Male pini than to other North American species in the and female specimens deposited at UW-Madison. nigricornis species group, given the high levels of shared polymorphism and low sequence divergence Oecanthus leptogrammus Walker TJ 1962. observed among O. pini and several other members Identification based on color, size, long slender of the nigricornis species group. hind femurs, and antennal markings. Photographs Neoxabea cerrojesusensis and N. ottei - and song waveforms (Figs. 42-47) are offered Although males of the two new Neoxabea species for documentation of this species. Metanotal are very similar in appearance, there are slight gland at figure 56. Dominant frequency 2.0 differences in the tegminal width. N. cerrojesusensis kHz at 27.3 C. Sound recording can be heard at males have slightly narrower wings with distal ends supporting material. Collected at Cerro Jesus. that appear to blend in with the hindwings; while N. This species was also found at San Marcos, and a ottei have slightly wider wings with broader distal female was collected at DRSP. The singing male ends (Fig. 18). Visual differences are more evident in the recording and one female from Cerro Jesus in the females of both species. N. cerrojesusensis deposited at UW-Mad. has an overall darker color than N. ottei, and has two prominent dark maculae compared to four for N. ottei (Fig. 18). The two new Neoxabea species were also ruled out as being three other species with stridulatory NANCY COLLINS ET. AL. 169 teeth counts closest down the list on Table 1 by 30km NE of DRSP. using characters presented in Walker TJ (1967). The Baker’s description (Baker 1905) included: distal tip of the cerci of N. astales and N. quadrula Length 18mm. Color a faded brownish, darker are club-shaped; and N. femorata has no knobs on on the posterior legs. The antennae are stout, the tergites. the two basal joints brownish, and each with a Oecanthus bakeri, O. belti and O. symesi slender, longitudinal, dark brown dash beneath; - Out of an abundance of caution, comparisons the flagellum is straw colored and with short white were made to species documented in Mexico: O. pubescence. The tegmina are 14mm long by 5.5mm argentinus, O. celerinictus, O. quadripunctatus, O. wide at the widest place; on the reflexed portion are exclamationis and O. alexanderi. These five species fourteen oblique veins. The wings reach to the tips are green with tegminal widths greater than 4.0 mm of the tegmina. The cerci are rather stout, extending and thus do not match O. symesi. None match O. to the wing tips, and covered with rather long white bakeri or O. belti as all five lack red on the pedicel hairs of varying lengths. The hind femora have or scape. Other non-matching factors are that O. three longitudinal series of short, oblique, straight exclamationis has distinctly different antennal brown dashes on the outer side, while both femurs markings (upside down exclamation mark) and O. and tibiae are sprinkled with minute brown dots; alexanderi chirps rather than trills. the tibiae bear five pairs of large spines besides the numerous small teeth; the apex of hind tibiae Oecanthus bakeri.—Although the stridulatory and the hind tarsi are thickly covered with a short teeth total for O. immaculatus is 24 and the golden pubescence. total for O. bakeri is 25, O. immaculatus has no Further analysis led to listing O. marcosensis antennal markings and is documented only in Peru. as a synonym of O. varicornis. The type specimen Overall colors and antennal markings appear quite is deposited at the National Museum of Natural different between O. bakeri and O. varicornis. History (NMNH), but is in a damaged state and is While O. bakeri has dark brown ventral markings missing its head and thorax. Walker TJ indicated in whitish fields on a straw-colored scape and in his 1962 paper that the possibility existed for O. pedicel, O. varicornis has black markings on a marcosensis to be a species “similar to but distinct red scape and pedicel., The stridulatory teeth total from O. varicornis.” and song pulse rate are similar to O. varicornis, Since the type specimen cannot be utilized and DNA sequences of O. bakeri showed minimal for an accurate comparison to the specimens divergence from O. varicornis; therefore, questions found in Nicaragua for this paper, it seemed more initially existed whether this was a pallid form reasonable to name a new species after Baker than of O. varicornis. Determination of new species to try to prove they are what Baker described as O. status, however, was made due to differences in marcosensis. the appearance of the tegmina (Fig. 22) and of the DNA was not analyzed for O. belti; therefore metanotal gland. The tegmen of O. varicornis has other characteristics were compared. an overall rounder shape than that of O. bakeri. The l/w ratio of the tegmina at rest: O. varicornis Oecanthus belti.—The stridulatory teeth total from Texas = 1.8 and O. bakeri = 2. Differences in of O. belti is close to that of O. varicornis, O. the metanotal glands can be viewed at figures 51 pallidus and O. major. With the tegmina closed and & 52. Arrows point to rounded lateral borders of at rest, the wings of O. belti present as oval-shaped; the scutum for O. varicornis, and scalloped lateral while the wings of O. varicornis are pear-shaped. borders for O. bakeri. These wing differences can be viewed at figures 25 It is interesting to note that in 1905, CF Baker and 57. described O. marcosensis, a tree cricket with The red coloring on the head and black similarities to O. bakeri found at DRSP. Baker antennae of O. belti is distinctly different from the described O. marcosensis from a site near San yellow on the head and pale antennae of O. pallidus. Marcos, Nicaragua. San Marcos is approximately The wings of O. pallidus are also somewhat pear- 170 NEOXABEA AND OECANTHUS OF NICARAGUA

shaped (Zefa et al. 2012). O. pallidus is known only Keys to Oecanthinae in Central America from Brazil. The tegminal width for O. major is 7.2mm The following keys summarize characters (Walker TJ 1967). O. major is known only from which are useful in identifying specimens or Mexico. live tree crickets in Central America. Oecanthus specimens do not retain colors as well as Neoxabea, Oecanthus symesi.—The narrow tegmina, but the antennal markings can generally be seen overall light golden yellow color and orange on discolored specimens. These keys include five head of O. symesi, along with the low number new species from Nicaragua and other tree crickets of stridulatory file teeth, indicate it is distinctly documented in Central America, using characters different from other known Oecanthus species. from Walker TJ, Gurney AB 1960, Walker TJ 1962, Tibial armature – Tibial armature for and Walker TJ 1967. Oecanthus species is comprised of large and small spines in two rows on the dorsum of the distal Key to Genera hind tibia. One row is on the lateral edge and one row is on the medial edge. The large spines are 1 Hind tibiae smooth with no spines (have located distally, with varying numbers of small terminal spurs only); prominent tubercle on spines between them. The small spines continue the first antennal segment...... Neoxabea proximally but do not extend into the femur. 1’ Distal hind tibiae with spines; no prominent The large spines on some species are situated as tubercle on the first antennal segment. . . . . opposing pairs (one in the lateral row and one in the ...... Oecanthus medial row); while the large spines on other species are alternating in position or have no relation to Key to Neoxabea Species in Central America those on the other row. Since the numbers and positions of spines for O. symesi were different 1 Males...... 2 from O. bakeri and O. belti, spines were examined 1’ Females ...... 5 on seven additional species of Oecanthus for comparison using a First Lab microscope with a 2(1) Stridulatory teeth count 30 or less ...... 3 10X digital eyepiece through a 4X power objective 2’ Stridulatory teeth count greater than 30 . . . lens. Large spines were situated as matching pairs, ...... formosa and there were generally three or four smaller spines between the large spines for: O. bakeri, O. 3(2) Setae or brushes prominently visible within belti, O. leptogrammus, O. quadripunctatus and metanotal orifice...... 4 O. varicornis. The large spines were not situated 3’ Sparse or no visible setae within metanotal as matched pairs, and there were zero to four small orifice...... bipunctata spines between same-row pairs of large ones for: O. forbesi, O. fultoni, O. niveus, O. pini and O. symesi. 4(3) Tegmina do not extend beyond lateral sides of abdomen...... cerrojesusensis 4’ Tegmina extend slightly past sides of abdo- men...... ottei 5(1’) Dark brown with two prominent blackish maculae ...... cerrojesusensis 5’ Medium brown with four prominent blackish maculae ...... ottei NANCY COLLINS ET. AL. 171

Key to Oecanthus species in Central America 7(5’) Tegminal width at rest atop abdomen 5.0mm or less; tegmen at rest have long oval appear- 1 Predominantly green in color; tegminal ance ...... belti width 4.1mm or more...... 2 7’ Tegminal width at rest atop abdomen 5.7mm 1’ Golden yellow color with dark orange head or greater; tegmen at rest have pear-shaped and pronotum; tegminal width 4.0mm or less appearance...... varicornis ...... symesi Nicaragua lacks an in depth study of 2(1) Markings on pedicel and scape not of same Orthoptera; therefore, the potential exists for configuration ...... 3 additional sites for Oecanthinae, as well as 2 Markings on pedicel and scape have same discovery of more new species. In order to further configuration; or only scape mark visible . 4 understanding of Oecanthinae in Nicaragua, investigation of host plants, and song recordings 3(2) Mark on scape round or comma-shaped, / analysis at a wider range of temperatures for mark on pedicel straight...... comma all species in Nicaragua are needed. As DNA 3’ Mark on pedicel round, mark on scape sequencing capabilities broaden, analysis of O. straight...... prolatus allardi from Nicaragua compared to O. allardi from the West Indies would be of interest. Additionally, 4(2) Markings on scape and pedicel are not round advanced DNA analysis of O. bakeri and O. dots ...... 5 varicornis would better define their placement on a 4’(2’) Round black dot in white field on both pedi- phylogeny tree. Further study of the tibial armature cel and scape; male song consists of trilled for all species of Oecanthus would be of interest. chirps...... allardi ACKNOWLEDGEMENTS 5(4). Both marks on pedicel and scape are readily visible; antennal markings are vertical lines Initial tree cricket specimens from Cerro on white fields; antennae pale; scape pale Jesus, Cerro Tisey, Datanli Diablo, Kahka Creek or straw-colored; pedicel same intensity or and Kilambé are products of LLAMA Leaf Litter lighter than scape...... 6 Survey of Mesoamerica with John 5’. Only scape mark readily visible; scape mark T. Longino and Robert S. Anderson as Project is black vertical line not on white field; an- Investigators of National Science Foundation Grant tennae black; scape dark red; pedicel darker # DEB-1157383. We are grateful to Jean Michel than scape...... 7 Maes for obtaining the necessary export permits, and the Ministerio del Ambiente y los Recursos 6(5) Medium width line on pedicel and scape; Naturales Nicaragua (MARENA) for granting the head predominantly pale orange; permits. Ricardo Antonio Jarquin Urbina, Wiston male song continuous trilling; dark slash Valle Mejia, Dan Heffern and Brian Raber assisted marks on hind femurs...... bakeri with collection of specimens; Laurel Symes 6’ Very thin line on pedicel and scape; patch of assisted with DNA sequencing; and Nydia Damaris medium orange on head; male song intermit- Molina identified the host plant for O. symesi at tent bursts of trilling; hind femurs without Cerro Jesus. We are appreciative of invaluable dark slash markings ...... leptogrammus suggestions from Thomas J. Walker and John LaPolla for improving this manuscript. Thomas J. Walker’s 1967 paper examining species found south of the United States was immensely helpful in diagnosing these new Nicaraguan species. We also acknowledge entomological equipment support from IDEAWILD to EV. 172 NEOXABEA AND OECANTHUS OF NICARAGUA

Finally, we extend thanks to Bing Kirk and the Walker TJ (1963) The and calling songs friendly helpful staff of Cerro Jesus for welcoming of United States tree crickets (Orthoptera: the advancement of science, generously leasing Gryllidae: Oecanthinae). II. The nigricornis facilities to scientists and environmentalists, and group of the genus Oecanthus. Annals of the above all having the foresight to conserve and Entomological Society of America 56: 772-789. protect one of few remaining significant examples Walker TJ (1967) Revision of the Oecanthinae of cloud forest in northern Nicaragua. (Orthoptera: Gryllidae) of America South of the United States. Annals of the Entomological REFERENCES Society of America 60: 784-796. Walker TJ (2013) Singing insects of North Baker CF (1905) Homoptera: On the Gnathodus America: crickets and katydids. [SINA] http:// Species of the Abdominalis Group. entnemdept.ifas.ufl.edu/walker/Buzz/ (accessed Invertebrata Pacifica 1:82. December 2013) Eades DC, Otte D, Cigliano MM, Braun H. Walker TJ, Gurney AB (1960) A new species of Orthoptera Species File Online. Version 2.0/4.1. Oecanthus from the West Indies (Orthoptera, Available from: http://Orthoptera.SpeciesFile. Gryllidae). The Florida Entomologist 43(1): org (accessed October 2013). 9-13. Cornell University. Macaulay Library. Available Walker TJ, Gurney AB (1967) The metanotal from: http://macaulaylibrary.org/audio/114499 gland as a taxonomic character in New World (accessed December 2013). Oecanthus (Orthoptera, Gryllidae). Proceedings Robillard T, Desutter-Grandcolas L (2006) of the Entomological Society of Washington Phylogeny of the cricket subfamily 69: 157-161. Eneopterinae (Orthoptera, Grylloidea, Zefa E, Schneid Neutzling A, Rutz Redü, D, Eneopteridae) based on four molecular loci Lobregat de Oliveira G, Martins L ( 2012) and morphology. Molecular Phylogenetics and A new species of Oecanthus and Oecanthus Evolution 40: 643-661. lineolatus Saussure, 1897 from Southern Brazil: Walker TJ (1962) The taxonomy and calling songs species description, including phallic sclerites, of United States tree crickets (Orthoptera: metanotal glands and calling song (Orthoptera: Gryllidae: Oecanthinae). I. The genus Gryllidae: Oecanthinae). Zootaxa 3360:55. Neoxabea and the niveus and varicornis groups of the genus Oecanthus. Annals of the Entomological Society of America 55: 303-322. NANCY COLLINS ET. AL. 173

Fig. 1. Collection sites in Nicaragua 174 NEOXABEA AND OECANTHUS OF NICARAGUA

Figs. 2-9. N. cerrojesusensis. 2. Male. 3. Female. 4. Antennal tubercles or knobs. 5. Cerci. 6. Knobs on dorsal abdominal tergites. 7. 1-sec waveform of staccato prelude (sound like clicks). 8. 1-sec waveform at 26.1 C; 90 p/s. 9. 14-sec waveform at 26.1 C. Staccato-like clicking sounds gradually increase in speed before onset of continuous trill which increases in volume and then ends abruptly. (One trilling period between yellow markers.) NANCY COLLINS ET. AL. 175

Figs. 10-17. N. ottei. 10. Male. 11. Female. 12. Antennal tubercles or knobs. 13. Cercus. 14. Knobs on dorsal abdominal tergites. 15. 1-sec waveform of staccato warm up transitioning to trilling. 16. 1-sec waveform of continuous trilling at 26.2 C; 94 p/s. 17. 3-sec waveform at 26.2 C. 176 NEOXABEA AND OECANTHUS OF NICARAGUA

Fig. 18. Comparisons of color and markings of N. cerrojesusensis (L) and N. ottei (R). Note the width of the tegmina of the N. cerrojesusensis males compared to those of the N. ottei. Also note the N. cerrojesusensis female has two prominent dark maculae on the forewings compared to four on N. ottei. NANCY COLLINS ET. AL. 177

Figs. 19-24. O. bakeri. 19. Male. 20. Female. 21. Tegmen of O. varicornis from Texas (L) and O. bakeri from DRSP (R). 22. Antennal markings. 23. 1-sec waveform from DRSP at 25.7 C; 80 p/s. 24. 22-sec waveform from DRSP at 25.7 C. 178 NEOXABEA AND OECANTHUS OF NICARAGUA

Figs. 25-29. O. belti. 25. Male. 26. Female. 27. Antennal markings. 28. 1-sec waveform at at 26.7 C; 81 p/s. 29. 20-sec waveform at 26.7 C. NANCY COLLINS ET. AL. 179

Figs. 30-36. O. symesi. 30. Male. 31. Female. 32. Antennal markings. 33. Cerci. 34. Tibial armature. 35. 1-sec waveform at 22.7 C; 58 p/s. 36. 10-sec waveform at 22.7 C. 180 NEOXABEA AND OECANTHUS OF NICARAGUA

Figs. 37-41. O. allardi. 37. Male. 38. Immature female. 39. Antennal markings. 40. 1-sec waveform at 27.2 C (one ‘chirp’ consisting of an opening double pulse then 11 trios). 41. 15-sec waveform at 27.2 C. NANCY COLLINS ET. AL. 181

Figs. 42-47. O. leptogrammus. 42. Male (Note: dark areas on limbs not present prior to exposure to alco- hol.) 43. Female. 44. Antennal markings. 45. 1-sec waveform at 27.3 C; 82 p/s. 46. 5-sec waveform at 27.3 C. 47. 20-sec waveform at 27.3 C. 182 NEOXABEA AND OECANTHUS OF NICARAGUA

Figs. 48-56. Metanotal glands: 48. N. cerrojesusensis 49. N. ottei 50. N. bipunctata (from US, for comparison) 51. O. varicornis (from US, for comparison with O. bakeri) 52. O. bakeri 53. O. belti 54. O. symesi 55. O. allardi 56. O. leptogrammus (Note: Some are aged specimens, thus colors are unreliable.) NANCY COLLINS ET. AL. 183

Fig. 57. Shape of tegmina of male O. varicornis from US, for comparison with tegmina shape of O. bakeri. 184 NEOXABEA AND OECANTHUS OF NICARAGUA

Fig. 58. Tibial armature of male O. varicornis from US, for comparison with O. symesi.