Ecologica Montenegrina 20: 222-256 (2019) This journal is available online at: www.biotaxa.org/em

https://zoobank.org/urn:lsid:zoobank.org:pub:5BDFDF5E-EBC5-4547-8488-C7563F1C21DF

Three new genera and four new species of katydids (: ) from and Hispaniola, Greater Antilles

SHEYLA YONG

Grupo de Sistemática y Ecología de Artrópodos Caribeños Calle 200 # 3759, e/ 37 y 45, Reparto Versalles; La Lisa 13500; La Habana. Cuba E-mail: [email protected]

Received 21 February 2019 │ Accepted by V. Pešić: 25 March 2019 │ Published online 31 March 2019.

Abstract Three new genera and four new species of katydids belonging to the subfamilies () and Pseudophyllinae (Cocconotini) are described from Cuba and Hispaniola. Fully illustrated descriptions and detailed comparisons are provided, as well as precise distribution maps and color photographs of habitus, main diagnostic structures and habitat.

Key words: Conocephalinae, Copiphorini, Pseudophyllinae, Cocconotini, , new taxa, new records, Neotropics.

Introduction

The katydid fauna of the Greater Antilles is currently composed by 68 nominal species, 67 of them living taxa plus a single amber fossil (Perez-Gelabert, 2008; Yong & Perez-Gelabert, 2014; Cigliano et al., 2018). Recent field studies conducted by the author and her collaborators in Cuba and Hispaniola have yielded many new species of Orthoptera. As another part of the ongoing revision of Caribbean orthopteroids by the author, here are described some of them, i.e., three new genera and four new species. The presented descriptions are thorough and fully complemented by complete collecting data and high-resolution photographs, plus detailed taxonomic comparisons for every taxon.

Materials and methods

The katydids were detected mainly on the vegetation by direct nocturnal visual search, with the aid of standard white LED headlamps. Once spotted, high resolution, full-color digital photographs were taken (habitat included) with a Nikon Coolpix S8100 digital camera. Then, all specimens were sacrificed by either exposition to 70% ethanol vapors or immersion in the same liquid; final preservation was made either in 80%

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YONG ethanol or dry-pinned. Collecting and identification labels were laser-printed in Spanish, but transcribed into English here. The specimens were studied and measured under an AmScope SM-1T-PL LED trinocular stereomicroscope, equipped with a 20x calibrated line scale ocular micrometer. All microscopic photographs were taken with a Nikon Coolpix S8100 digital camera attached to the upper ocular tube of the microscope. Photographs of habitus were taken with a Nikon Coolpix B500. Images were processed with Adobe Photoshop CS5, only for contrast and brightness optimization, background cleanup and plate composition. As for the measurements, the interocular distance corresponds to the shortest straight line between the facing eye margins. Fastigium width was taken at its base. Distribution maps were constructed in Mapinfo Professional 10.0, using precise coordinates either taken in situ with a portable GPS device (Datum WGS84) or extracted from 1: 25,000 military reference maps. All along the text, counts are given as fractions for left/right sides. Some data were verified using the Orthoptera Species File Online (OSF), version 5.0/5.0 (Cigliano et al., 2019). Repositories are abbreviated using the following acronyms: IES: Instituto de Ecología y Sistemática, Havana, Cuba. MNHNCu: Museo Nacional de Historia Natural de Cuba, Havana, Cuba. FZ: Fernando de Zayas private collection (formerly at Havana, Cuba, current whereabouts unknown). SY: Personal collection of Sheyla Yong, Havana, Cuba.

Systematics Family Tettigoniidae Krauss, 1902 Subfamily Conocephalinae Burmeister, 1838 Tribe Copiphorini Karny, 1912 Erioloides Hebard, 1927

Erioloides santiago new species Figures 1–3, 15, 19, 22. Table I "Genus unknown 6": Yong & Perez-Gelabert, 2014: 427.

Types. CUBA: Province: Santiago de Cuba Municipality: Baconao: Playa Verraco (19°53'34"N - 75°34'32"W, 10 m a.s.l.); 18/April/2017; at night, on bushes; S. Yong, R. Teruel, E. Fonseca, J. L. Reyes; molted to adult on 20/May/2017; 1♀ holotype (SY, in ethanol 80%). Same data as holotype, but molted to adult on 5/June/2017; 1♀ paratype (SY, in ethanol 80%). Siboney (19°57'56"N - 75°42'38"W, 125 m a.s.l.); diurnal net-sweeping of vegetation; 25/October/2017; B. Lauranzón, J. L. Reyes; 1♀ juvenile paratype (SY, in ethanol 80%). Suburbs of Santiago de Cuba City; no more data; 1♀ paratype (SY, dry pinned). Palma Soriano Municipality: Presa Charco Mono (20°04'54"N -76°00'31"W, 300 m a.s.l.); 8/December/ 2017; S. Yong, R. Teruel; 1 ♀ juvenile paratype (SY, in ethanol 80%).

Additional material examined. CUBA: Camagüey Province: Santa Cruz del Sur Municipality: Archipiélago de los : Cayo Anclitas (20°46'54"N - 78°54'42"W, 2 m a.s.l.); 2/May/1971; L. F. de Armas; 1♂ (IES, dry pinned). Las Tunas: Manatí Municipality: Puerto Manatí (21°21'41"N - 76°49'34"W, 2 m a.s.l.); December/1988; L. R. Hernández; 1♂ (MNHNCu, dry pinned). Province Santiago de Cuba Province: Santiago de Cuba Municipality: Santiago de Cuba City; February/1930; P. Alayo; 1♂ (FZ, dry pinned). Cuabitas (20°03'59"N - 75°48'20"W, 120 m a.s.l.); January/1952; 1♀ (FZ, dry pinned). Versalles (20°03'59"N - 75°48'20"W, 100 m a.s.l.); June/ 1971; I. García; 1♀ (IES, dry pinned). Note. See additional discussion on these specimens below, in Remarks section.

Diagnosis (Based mostly on females). Body slender and medium-sized for the genus (18–19 mm). Both sexes macropterous, tegmina lanceolate, over 4.0 times the length of abdomen. Head with tegument coarsely and densely punctate, lacking any keels. Fastigium moderately short (slightly shorter than scapus length), shaped as an equilateral triangle in lateral view and with tip acute. Eyes small and spherical. Pronotum almost as long as wide, coarsely punctate; longitudinal sulcus inconspicuous. Legs moderately slender,

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THREE NEW GENERA AND FOUR NEW SPECIES OF KATYDIDS FROM CUBA AND HISPANIOLA ventrally armed with minute spines. Supra-anal plate basically paraboloid, much wider than long; posterior margin trilobed. Female cerci unmodified, conical. Subgenital plate wider than long; posterior margin apically with two lateral digitiform projections and a median notch. Ovipositor broad and slightly curved upwards, slightly longer than abdomen; dorsal and ventral edges entirely smooth; tip blunt and truncate. Upper valves with a file of stretches along its dorsal edge.

Figure 1. Adult female holotype of Erioloides santiago n. sp., full-body views: a) dorsal; b) lateral; c) ventral. Scale bar in millimeters.

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Figure 2. Adult female holotype of Erioloides santiago n. sp., close-up views: a) head and pronotum, dorsal; b) head and pronotum, lateral; c) head, frontal; d) apex of abdomen, dorsal; e) apex of abdomen, ventral; f) Sternites; g) apex of abdomen and ovipositor, lateral.

Etymology. The selected epithet is a noun in apposition, taken from the toponym of both the province and municipality where the type-locality of this species is enclaved.

Description of adult female holotype. Size medium for the tribe (total length 18.5 mm). General coloration pale green except as follows: eyes bluish gray, central ocellus white; antennal sockets, border of fastigium and face punctations yellowish. Head dorsally with a thin whitish yellow line extending from behind the eyes through the lateral borders of pronotum; clypeus yellowish to whitish, with a black U-shaped mark; labrum whitish, progressively darkened to brownish orange apically; mandibles and maxillary palps brownish orange, labial palps whitish. Tegmina with veins pale yellow, forming a reticulate pattern. Legs green, except for brownish tympani and tarsi; metafemur and metatibia dorsally with a yellowish sheen. Apex of ovipositor dark reddish brown. See figures 1, 15 and table I.

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Table I. Measurements of type specimens of Erioloides santiago n. sp from Santiago de Cuba, Cuba. Abbreviations: length (L), width (W), depth (H), not applicable (NA).

Dimensions (mm) Female holotype Female paratype Head L / W / H 3.25 / 5.40 / 6.15 3.50 / 5.75 / 6.25 Interocular distance L 2.25 2.50 Eye diameter L 1.10 1.15 Scapus L / W 1.60 / 0.90 1.90 / 0.80 Pedicel L / W 0.65 / 0.45 0.70 / 0.40 Fastigium L / W / H 1.25 / 1.10 / 1.25 1.70 / 1.15 / 1.25 Pronotum L / W 6.65 / 6.50 6.85 / 6.35 Tegmen L / W 32.30 / 6.00 35.00 / 5.85 Abdomen L 8.60 8.50 Cerci L / W 2.10 / 0.40 2.10 / 0.45 Tergite X L / W 1.50 / 2.55 0.80 / 2.40 Supra-anal plate L / W 0.55 / 2.50 0.30 / 1.50 Subgenital plate L / W 1.40 / 3.05 1.65 / 2.55 Styli L / W NA NA Ovipositor L / W 11.25 / 2.35 13.00 / 2.60 Profemur L / W 5.50 / 1.50 5.50 / 1.60 Protibia L 6.25 6.25 Mesofemur L / W 5.40 / 1.45 5.50 / 1.55 Mesotibia L 11.35 6.30 Metafemur L / W 13.00 / 2.30 13.05 / 2.50 Metatibia L 13.15 13.75 Body (total) L 18.50 18.85

Head (fig. 2a–c). Large, wider than long (ratio = 1.6). Tegument coarsely and densely punctate. Vertex flat in lateral view. Fastigium moderately short, pentagonal in dorsal view, triangular in lateral view and hexagonal in frontal view; tip slightly shorter than scapus and acute but not sharp. Eyes small and spherical. Median ocellus small, oval and translucent, located between the antennal sockets. Frons shallowly convex, coarsely punctate and lacking any keels. Genae convex in frontal view. Antennal sockets situated right between the eyes. Antennae standard for Conocephalinae, slightly surpassing the length of the tegmina and covered with minute setae; scapus subcylindrical, longer than wide (ratio = 1.8) and slightly longer than fastigium, oval in cross-section and covered with minute setae; pedicel cylindrical, shorter than scapus (ratio = 0.4), and with the same ornamentation. Thorax. Tegument granulose and coarsely punctate, with scattered minute setae. Pronotum slightly longer than wide, trapezoidal in dorsal view; anterior margin almost straight, longitudinal sulcus inconspicuous; posterior margin very shallowly convex; lateral lobes slightly sinuose and angled at 90°; anterior sulcus deep and narrow, median and typical sulci inconspicuous. Tegmina and wings fully developed, surpassing ovipositor tip. Tegmina narrow, much longer than abdomen (ratio = 3.7). Wings not surpassing the tips of tegmina. All thoracic sterna unarmed (fig. 2f). Legs (fig. 1). Covered with minute setae all over. Legs slender, genicular lobes apically armed with a pair of small, triangular spines. Procoxa dorsally with a large spine pointing forwards. Profemur moderately slender, subrectangular and moderately compressed laterally; ventral pair of carinae slightly prominent and straight, intercarinal space shallowly concave; dorsal surface unarmed, ventral surface with 0:5 / 6:0 spines. Protibia longer than profemur (ratio = 1.1), slender and rectangular in cross-section, with 7:8 / 8:7 ventral spines; tympanal area moderately swollen, tympanic slits equal, small, narrow and located dorsally at the base of protibia. Mesofemur very similar to the profemur, with 6:0 / 0:5 ventral spines. Mesotibia with 8:7 / 7:8 ventral spines. Metafemur slender, almost as long as metatibia, oval in cross-section, with 13:4 / 4:10 ventral spines; left metatibia with 9:12 and 9:7 subapical spines on dorsal and ventral surfaces, respectively; right metatibia with 10:11 and 9:9, respectively. Metatibia with two pairs of apical spurs slightly curved

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YONG inwards: one lateroventral and another ventral (smaller). Tarsi three-segmented: I and II of the same length and short, III apically bilobed; claws short. Abdomen. Moderately robust, subcylindrical; tegument glossy. Tergite X (penultimate) wider than long (ratio = 1.7), with posterior margin sloping downwards and widely bilobed by a deep V-shaped median notch. Supra-anal plate (fig. 2d) basically paraboloid, much wider than long (ratio = 4.5); posterior margin unequally trilobed, i.e., with two large lateral projections (digitiform) and a small median projection (sharp); tegument glossy, with minute setae scattered all over. Subgenital plate (fig. 2e) wider than long (ratio = 2.2); tegument glossy; anterior margin shallowly convex, posterior margin apically complex, with two lateral digitiform projections and a deep median notch. Cerci short, conical and densely covered with short, rigid setae. Ovipositor (fig. 2g) slightly longer than abdomen (ratio = 1.3), broad and slightly curved upwards; upper edge shallowly flanged medially, lower edge evenly and shallowly convex, both edges entirely smooth; ovipositor tip blunt and truncate. Upper valves with a file of parallel stretches along its longitudinal axis, near the dorsal edge.

Figure 3. Additional adults of Erioloides santiago n. sp., from other collections: a) IES male from Cayo Anclitas; b) MNHNCu male from Puerto Manatí; c) FZ male from Santiago de Cuba; d) FZ female from Cuabitas.

Adult male. In September 2013, the author examined and photographed three adults of this sex from three Cuban collections: IES, MNHNCu and FZ (fig. 3), but all these specimens are currently unavailable (see below, in Remarks section).

Variation. The second female paratype from Santiago de Cuba City is actually slightly larger than the holotype (see table I), but it looks smaller because it was preserved dry-pinned and the abdomen shrank back by desiccation.

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Comparison. On morphological grounds, the closest relative to E. santiago n. sp. is also its nearest neighbor, but this has gone unnoticed so far. The detailed color photographs of the types of Eriolus jamaicensis Brunner, 1915 (available on OSF), clearly show that this species endemic to Jamaica does possess all characters currently diagnostic for Erioloides (see Naskrecki, 2000: 79), with the single exception of a longer fastigium (almost twice the diameter of the eye), which alone does not justify its generic segregation. Thus, it is formally transferred here as Erioloides jamaicensis (Brunner, 1915), n. comb. It differs from the new species as follows (female only). According to the measurements given by Bruner (1915: 388), E. jamaicensis n. comb. is remarkably larger, e.g., in the single female paralectotype size is 32 mm and lengths of pronotum, tegmina and metafemur are 7.25, 37.0 and 12.5 mm, respectively. Moreover, according to the photographs available on OSF, its fastigium is much longer (more than twice as long as scapus) and sharper, spiniform in lateral view and lanceolate in frontal view.

Distribution (fig. 22). From the scant available data, this species seems endemic to Cuba but widespread across the eastern half of the archipelago; see below, in Remarks section.

Ecological notes. The two females from Playa Verraco were found in close proximity around midnight, on low roadside bushes (less than 1.5 m above the ground), about 10 m away from the seashore. The vegetation is coastal xerophytic scrub on limestone karstic terrain (fig. 19d); this is one of the most arid areas of the Cuban archipelago. Both specimens were collected as mid-instar juveniles (15c–d) and kept alive in captivity, where each individual molted twice to reach adulthood. The juvenile from Charco Mono was also collected at night (around 21:30 hrs. EST), on the outer foliage of the tree Senna atomaria (L.) H. S. Irwin & Barneby (Fabaceae), approximately 4.0 m above the ground. The vegetation is a bare secondary grassland (pasture) with scattered trees, but it was originally a mesic semicaducifolious forest on volcanic sandy terrain (fig. 19b), on the northern slopes of the Sierra Maestra. According to the field notes of the collectors (Beatriz Lauranzón and Jorge L. Reyes, pers. comm.), the juvenile from Siboney was captured in the backyard of an abandoned house, while sweeping with an entomological net on the low Verbenaceae bushes Stachytarpheta jamaicensis (L.) Vahl, 1804. The primary vegetation around the site is microphyllous semicaducifolious forest and subcoastal xerophytic scrub, both on limestone karstic terrain (fig. 19c), about 1 km away from the seashore; the aridity of the site is quite similar to Verraco. No ecological data are available from the labels of the dry-pinned specimens. According to the collector (Rolando Teruel, pers. comm.), the vegetation and landscape of Cayo Anclitas and Puerto Manatí is very similar: a mosaic of mesic coastal semicaducifolious forest and mangrove on karstic terrain, with open grassland along sandy beaches and in anthropic clearings (fig. 19a).

Remarks. Though widespread in Cuba, this species seems to be quite rare and/or sporadic. Most of its collecting sites have been extensively sampled by the present author for three years now, and for decades by her collaborators and other skilled collectors (even well-reputed entomologists). Nevertheless, the listed specimens remain the only ones known so far. The four adults from IES, MNHNCu and FZ collections (three males and one female, see above in Additional Material Examined and Male sections), have all been assigned tentatively to E. santiago n. sp. Their conspecificity must be confirmed when they become available again (see below), or when additional samples from the same localities are obtained. The IES and MNHNCu single males were officially loaned to Daniel E. Perez-Gelabert in September/2013 but remain unpublished and neither have been returned yet to their legal repositories, despite the loan is already overdue. The FZ male and female pair is also missing: on October 14, 2018, the author contacted Fernando de Zayas' heir by phone, who stated that the entire FZ collection had been sold out and refused to clarify its final destination. Prior to the present contribution, Erioloides comprised 11 nominal species distributed from Mexico through Central America, down to northern South America (Cigliano et al., 2019). The description of E. santiago n. sp. and the transfer of E. jamaicensis n. comb. implicate the first official records of the genus from the Greater Antilles. It is worth noting that Naskrecki (2000) mentioned this genus to occur "throughout the Caribbean", but did not list any Greater Antillean location. Moreover, Erioloides is even more

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Rombophora new genus Figures 4–6, 16, 20c–d, 23. Table II Type species: Rombophora sierramaestrae n. sp., by present designation.

Diagnosis (Based mostly on females). Body stout and medium-sized for the subfamily (18–19 mm). Both sexes macropterous; tegmina rhomboidal, sexually dimorphic (shorter, wider and more angulose in male), surpassing tip of abdomen (male), but not reaching tip of ovipositor (female); hind wings fully developed. Head with smooth tegument, lacking any keels. Fastigium very large, sharp and slightly curved downwards, upper surface with minute denticles in basal half, base of fastigium with a very large tooth, vertically separated by a deep notch from a large frontal tooth. Eyes small and spherical. Pronotum almost as long as wide; dorsal disk strongly projected, paraboloid; longitudinal sulcus inconspicuous. Legs slender and ventrally armed with minute spines; genicular spines very long and sharp. Female cerci unmodified, broadly conical. Ovipositor moderately short (but longer than abdomen), slender, straight and entirely smooth.

Etymology. The generic epithet is an arbitrary combination of letters, which mixes the Spanish word ''rombo'' (meaning "rhomb" and alluding to tegmina shape) with the final syllables of Serville, 1831 (its putative closest relative). It is declared here feminine in gender.

Comparisons. Rombophora n. gen. most closely resembles Copiphora, Acantheremus Karny, 1907 (both Neotropical) and Stål, 1873 (native to Asia and America). From both Copiphora and Acantheremus, the new genus can be separated as follows: 1) Size smaller. 2) Prosternum with a pair of conical spines. 3) Base of fastigium narrower than scapus. 4) Genae lacking lateral carinae. 5) Mesotibia dorsally unarmed. 6) Ovipositor moderately short with smooth apex. In the other two genera, size is larger, prosternal spines are absent, fastigium base is wider than scapus, genae have moderate to strong lateral carinae, mesotibia is dorsally armed with spines (always in Copiphora, usually in Acantheremus), and ovipositor is remarkably different (much to exaggeratedly longer in Copiphora, longer and with apex minutely serrate in Acantheremus). Last, Rombophora n. gen. can be separated from Pyrgocorypha by: 1) Size smaller. 2) Tegument mainly smooth and glossy, on pronotum feebly punctate. 3) Base of fastigium narrower than scapus. 4) Fastigium dorsally with minute denticles in basal half and apically not hooked. 5) Meso- and metasternum lacking lateral basisternal lobes. 6) Ovipositor shorter and slightly curved upwards. In Pyrgocorypha size is larger, the tegument is moderately granulose in a reticulate pattern, fastigium base is wider than scapus, fastigium shape is different (dorsally lacking denticles and apically hooked), meso- and metasternum possess lateral basisternal lobes and ovipositor is longer and straight.

Distribution (fig. 23). Monotypic genus, known only from the high peaks (1,000–1,752 m. a.s.l) of the Sierra Maestra. From the scarce data available, the single species is endemic to these mountains, but widespread across its two main sections: the western Turquino Range and the eastern Gran Piedra Range.

Rombophora sierramaestrae new species Figures 4–6, 16, 20c–d, 23. Table II Pyrgocoripha? [sic] Zayas, 1976: 63, 64; fig. 53. Misidentification and Incorrect Subsequent Spelling (ISS).

Types: CUBA: Santiago de Cuba Province: Santiago de Cuba Municipality: Cordillera de La Gran Piedra: La Gran Piedra (20°00'42"N - 75°37'37"W, 1,000–1,230 m a.s.l.); on the vegetation; 14/April/2017; S. Yong, R. Teruel, E. Fonseca; 1♀ adult holotype (SY, in ethanol 80%). Same locality; 4–6/October/2015; S. Yong, R. Teruel; 1♀ juvenile paratype (SY, in ethanol 80%). Same locality; 23/May/2016; S. Yong, R. Teruel; 1♀ adult (dry pinned), 1♀ juvenile (ethanol 80%), both paratypes (SY). Same locality; 5/July/2016; Y. Ricardo; 1♀ adult paratype (SY, dry pinned). Same locality; 23/July/2017; S. Yong, R. Teruel; 1♂ juvenile, 1♀ juvenile paratypes (SY, in ethanol 80%). Same locality; 11–16/July/2018; S. Yong, R. Teruel; 1♀ adult paratype (SY, in ethanol 80%). Guamá Municipality: Cordillera del Turquino: Pico La Bayamesa (20°02'46"N - 76°35'19"W, 1,752 m a.s.l.); August/2017; J. R. Fuentes; 1♂ juvenile (SY, in ethanol 80%).

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Figure 4. Adult female holotype of Rombophora sierramaestrae n. gen., n. sp., full-body views: a) dorsal; b) lateral; c) ventral. Scale bar in millimeters.

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Figure 5. Adult female holotype of Rombophora sierramaestrae n. gen., n. sp., close-up views: a) head and pronotum, dorsal; b) head and pronotum, lateral; c) head, frontal; d) apex of abdomen, dorsal; e) apex of abdomen, ventral; f) Sternites; g) apex of abdomen and ovipositor, lateral.

Aguada de Joaquín (20°00'47"N - 76°50'02"W, 1,663 m a.s.l.); 27/January–2/February/2018; A. A. Méndez, A. Rapado; 1♂ juvenile, 1♀ juvenile, both paratypes (SY, in ethanol 80%).

Additional material examined. CUBA: Santiago de Cuba Province: Santiago de Cuba Municipality: Cordillera de La Gran Piedra: La Gran Piedra (20°00'42"N - 75°37'37"W, 1,230 m a.s.l.); June/1964; F. de Zayas; 1♀ adult (FZ, dry pinned). Guamá Municipality: Cordillera del Turquino: Alto del Cardero (19°58'31"N - 76°51'15"W, 1,238 m a.s.l.); F. de Zayas; 2♀♀ adults (FZ, dry pinned).

Record from high-quality photograph. CUBA: Santiago de Cuba Province: Guamá Municipality: Sierra Maestra: Cordillera del Turquino: Aguada de Joaquín (20°00'47"N - 76°50'02"W, 1,663 m a.s.l.); 1♂ (see Navarro, 2010: 304).

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Diagnosis. See genus diagnosis above.

Etymology. The specific epithet is a Latinized eponym, derived from the name of the mountain range that contains all localities known for the species (the Sierra Maestra).

Description of adult female holotype. Size medium for the tribe (total length 19.35 mm). General coloration pale green, except as follows: eyes black, central ocelli white; antennae pale reddish brown; clypeus, labrum and mandibles vivid reddish purple; labial and maxillary palps white, with orange tips. Tegmina scattered with small red spots. Metafemur spines white, with black tips. Subapical spines and claws of metatibiae reddish brown. Abdomen very pale green, almost white. Ovipositor reddish brown. See figures 4, 16 and table II. Head (fig. 5a–c). Large, wider than long (ratio = 1.9). Tegument smooth and glossy, with minute and thin setae scattered all over. Vertex shallowly convex in lateral view. Fastigium very large, sharp and slightly curved downwards; in lateral view with upper surface shallowly convex and lower surface concave; in dorsal view subtriangular; basal half dorsally with minute denticles, base of fastigium ventrally with a very large tooth, vertically separated by a deep notch from a large apical tooth in frons. Eyes small and spherical. Median ocellus small, round and translucent, located between antennal sockets. Frons shallowly convex and glossy, lacking lateral edges. Genae convex in frontal view, lacking lateral carinae. Antennal sockets situated right between the eyes. Antennae standard for Conocephalinae, remarkably longer than body; scapus subcylindrical, longer than wide (ratio = 2.0) and much shorter than fastigium, oval in cross- section and covered with minute setae; pedicel cylindrical, almost half the length of scapus. Thorax. Tegument glossy and feebly punctate, with minute setae scattered all along the margins. Pronotum almost as long as wide, subrectangular in dorsal view, with dorsal disk paraboloid and strongly projected posteriorly; anterior margin very shallowly notched medially, posterior margin strongly convex and projected backwards, lateral lobes subquadrate; longitudinal sulcus inconspicuous, anterior and typical sulci shallow and narrow. Tegmina and wings fully developed; tegmina narrow, angulose and much longer than abdomen (ratio = 2.2), but not reaching ovipositor tip. Wings not surpassing the tips of tegmina. Prosternum armed with a pair of relatively short and thin, conical spines. Basisternum of meso- and metasternum lacking lateral lobes (fig. 5f). Legs (fig. 4). Slender, covered with minute setae all over. Genicular lobes armed with a pair of apical spines, which are triangular, moderately long, sharp and pointing backwards. Procoxa dorsally with a large spine pointing forwards. Profemur subquadrate in cross-section, slightly compressed; ventral pair of carinae weak to moderate and straight, with 3:3 / 4:2 spines. Protibia longer than profemur (ratio = 1.2), rectangular in cross-section; ventral pair of carinae slightly sinuose, with 6:6 / 6:6 spines; tympanal area moderately swollen, slits equal, large, oval, and located dorsally at the base of protibia. Mesofemur very similar to profemur, with 4:4 / 4:4 ventral spines. Mesotibia with 6:7 / 5:7 ventral spines. Metafemur robust, shorter than metatibia (ratio = 0.9), oval in cross-section and with 11:4 / 4:10 ventral spines. Left metatibia with 9:9 and 10:7 subapical spines in its dorsal and ventral surfaces, respectively; right metatibia with 9:8 and 7:10, respectively; three pairs of apical spurs curved inwards: one dorsal, one lateroventral and one ventral (smallest). Tarsi four-segmented: I and II short and same-length, III much wider; claws longer than tarsi I + II, curved. Abdomen. Short and stout, subcylindrical, tegument glossy, with minute setae scattered all over. Tergite X (penultimate) much wider than long (ratio = 3.4), weakly furrowed longitudinally, with posterior margin widely and deeply notched (M-shaped). Supra-anal plate (fig. 5d) acutely paraboloid, wider than long (ratio = 2.9), with longitudinal furrow large, deep and distally wider, tegument coriaceous, with scattered minute setae. Subgenital plate (fig. 5e) hexagonal, wider than long (ratio = 2.0), and slightly tectiform; tegument glossy to coriaceous, with minute setae scattered all over; anterior margin shallowly convex, lateral margins angulose, posterior margin shallowly convex. Cerci very large and thick (compared to supra-anal plate length, ratio = 3.8), conical, densely covered with long and thin setae. Ovipositor (fig. 5g) rather short (but longer than abdomen, ratio = 1.1), slender, straight and with entirely smooth edges (i.e., lacking any teeth, serrations or crenulations); tegument glossy and vestigially punctate; upper and lower margins basically straight and parallel, upper margin with tip shallowly curved downwards, lower margin with tip shallowly curved upwards.

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Table II. Measurements of type specimens of Rombophora sierramaestrae n. gen. n. sp. from La Gran Piedra, Santiago de Cuba. Abbreviations: length (L), width (W), depth (H), not applicable (NA).

Dimensions (mm) Female holotype Female paratype Head L / W / H 3.25 / 6.10 / 0.75 3.00 / 6.30 / 6.90 Interocular distance L 2.65 2.85 Eye diameter L 1.10 1.20 Scapus L / W 1.85 / 0.90 2.00 / 0.85 Pedicel L / W 0.80 / 0.50 0.65 / 0.50 Fastigium L / W / H 3.70 / 1.55 / 1.80 3.55 / 1.55 Pronotum L / W 7.25 / 7.40 7.70 / 7.60 Tegmen L / W 19.50 / 5.25 20.65 / 5.75 Abdomen L 8.85 8.25 Cerci L / W 2.30 / 0.65 2.30 / 0.65 Tergite X L / W 0.80 / 2.70 NM Supra-anal plate L / W 0.60 / 1.75 NM Subgenital plate L / W 1.40 / 2.80 1.05 / 2.25 Styli L / W NA NA Ovipositor L / W 10.00 / 1.20 12.50 / 1.15 Profemur L / W 5.65 / 1.55 6.00 / 1.65 Protibia L 7.00 7.75 Mesofemur L / W 5.75 / 1.55 6.15 / 1.70 Mesotibia L 7.00 7.07 Metafemur L / W 13.80 / 3.00 14.15 / 2.90 Metatibia L 15.35 15.15 Body (total) L 19.35 18.95

Male. Only three juveniles available for this study, plus a high resolution, full-color photograph of an adult specimen from Aguada de Joaquín (Cordillera del Turquino, Sierra Maestra; see fig. 16a). The latter is very similar to the described females, but shows the following sexually dimorphic characters: 1) habitus stouter; 2) clypeus, labrum and mandibles yellowish orange with black edges; 3) pronotum with dorsal disk remarkably more projected posteriorly, covering stridulatory apparatus; 4) tegmina wider and dorsally more angulose, strongly rhomboidal.

Variation. One female paratype from La Gran Piedra is slightly smaller than the holotype (see table II) and the other adult paratypes.

Distribution (fig. 23). As for the genus (see above).

Ecological notes. The specimens collected by the present author at La Gran Piedra were all found before midnight, on the vegetation but in variable conditions. The holotype and one adult paratype were both found walking on roadside bushes, about 1 m above the ground. Another adult paratype was walking on a fern frond 0.2 m above the ground, in the bottom of a rainforest creek. The fourth adult paratype was standing on a tree twig 2.0 m above the ground, also in the rainforest. In contrast, the juveniles (including additional individuals that were observed but not collected), were all feeding on the fresh seeds of grass spikes, less than 1 m above the ground, at the clear sides of the road, trails and sidewalks. La Gran Piedra is the highest peak of the eastern sector of the Sierra Maestra. The vegetation is mostly composed of montane rainforest and pine forest (fig. 20d), but human settlements in the area for more than 150 years have introduced coffee plantations and cleared areas to construct buildings and gardens. The site is wet and cold, especially at night when temperature usually drops below 15°C (even in summer) and the air humidity reaches saturation point. Daytime temperature seldom raises above 25°C and the mountaintop is frequently misty or enveloped in low clouds (fig. 21c).

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According to the field data of the collectors (Abelardo A. Méndez and José R. Fuentes, pers. comm.), the specimens from Aguada de Joaquín and Pico La Bayamesa (Turquino Range) were all captured under exactly the same conditions (fig. 20c), but at higher altitude and in less anthropized areas.

Remarks. Despite having been collected repeatedly and all through the year, this species is remarkably rare and scarce everywhere it occurs. Only one or a few specimens were found in each sampling event, each usually about one week long (see Material Examined section above).

Figure 6. Adult females of Rombophora sierramaestrae n. gen., n. sp., from FZ collection.

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Genus Unicorniella new genus Figures 7–10, 17, 21, 24. Table III Type species: Unicorniella hatueyi n. sp., by present designation.

Diagnosis. Body slender and size large for the subfamily (26–29 mm). Both sexes macropterous; tegmina lanceolate, sexually not dimorphic (size and shape similar in both sexes), clearly surpassing tip of abdomen (male) and ovipositor (female); hind wings fully developed. Head with tegument smooth, lacking any keels. Fastigium very large, very sharp and very slightly curved downwards, base of fastigium with a very large tooth, vertically separated by a deep notch from a large frontal tooth. Eyes small and spherical. Pronotum longer than wide; dorsal disk strongly projected, truncate; longitudinal sulcus inconspicuous. Legs very long, slender and ventrally armed with minute spines; genicular spines very long and sharp. Male cerci curved inwards and armed apically with two sclerotized and strongly incurved spines; female cerci unmodified, narrowly conical. Ovipositor long, slender, straight and entirely smooth.

Etymology. The generic epithet is an arbitrary combination of letters, which mixes the Spanish noun "Unicornio" with the Latin diminutive ending "-ella". It is declared feminine in gender. Unicorns are fantastic creatures believed to live deep inside the remotest forests, thus, only seldom seen; the epithet alludes to both the habitat of this katydid and its sporadic findings.

Comparisons. Unicorniella n. gen. is very closely related to two other Copiphorini genera: Karny, 1907 and Pyrgocorypha. The new genus can be separated from Neoconocephalus as follows: 1) Habitus slenderer. 2) Fastigium very long and sharp. 3) Genicular spines conspicuously larger. In Neoconocephalus, the habitus is generally stouter, the fastigium is short and blunt (the single exception being N. aduncus (Scudder, 1878), which has it C-shaped), and the genicular spines are much shorter. Last, Unicorniella n. gen. can be separated from Pyrgocorypha by: 1) Habitus slenderer. 2) Fastigium apically not hooked. 3) Ovipositor longer than abdomen. 4) Mirror of right tegmen longer than wide. In Pyrgocorypha, the habitus is generally stouter, the fastigium is apically hooked, the ovipositor is as long as abdomen, and the mirror of right tegmen is essentially circular.

Distribution. Monotypic genus, known only from the mountains of the neighboring Greater Antillean islands of Cuba and Hispaniola. In the former, it is scattered all across the main island, but in the latter, it is known from a single locality in southern Dominican Republic (fig. 24).

Unicorniella hatueyi new genus, new species Figures 7–10, 17, 21, 24. Table III "Especie indeterminada, presumiblemente de..." [Neoconocephalus]: Zayas, 1976: 64. Misidentification "Genus unknown 5": Yong & Perez-Gelabert, 2014: 427.

Types. CUBA: Santiago de Cuba Province: Santiago de Cuba Municipality: Cordillera de La Gran Piedra: La Gran Piedra (20º00'35''N - 75º37'42''W, 1,200 m a.s.l.); on the vegetation; 11–16/July/2018; S. Yong, R. Teruel; 1♂ adult holotype (SY, green phase, in ethanol 80%), 1♂ adult (brown phase), 4♂♂ juveniles (green phase), 4♀♀ juveniles (green phase), all paratypes (SY, in ethanol 80%). Same locality; 4–6/October/2015; S. Yong, R. Teruel; 1♂ juvenile, 1♀ juvenile, both paratypes (SY, green phase, in ethanol 80%). Same locality; 11/March/2016; R. Teruel, J. L. Reyes; 1♀ adult paratype (SY, green phase, in ethanol 80%). Same locality; 23/July/2017; S. Yong, R. Teruel; 1♂ juvenile paratype (SY, green phase, in ethanol 80%).

Additional material examined. CUBA: Artemisa Province: San Cristóbal Municipality: Sierra de Rangel (22°44'11"N - 83°11'29"W, 600 m a.s.l.); 12–15/January/1934; J. Acuña, A. R. Otero; 1♀ adult (brown phase, FZ, dry pinned). Province: Cumanayagua Municipality: Guamuhaya Massif: Pico San Juan (21°59'23"N - 80°08'51"W, 1,140 m a.s.l.); 7–14/August/2015; S. Yong, E. Fonseca, T. M. Rodríguez- Cabrera; 3♂♂, 9♀♀ adults (both green and brown phases, now lost). Guantánamo Province: Baracoa Municipality: Nibujón (20°30'26"N - 74°39'13"W, 10 m a.s.l.); 6/March/1979; L. B. Zayas; 1♂ adult (green phase, IES, dry pinned). DOMINICAN REPUBLIC: Santo Domingo Province: San Felipe de Villa Mella

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Figure 7. Adult male holotype of Unicorniella hatueyi n. gen., n. sp., full-body views: a) dorsal; b) lateral; c) ventral. Scale bar in millimeters.

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Figure 8. Adult male holotype (a–i) and adult female paratopotype (j) of Unicorniella hatueyi n. gen., n. sp., close-up views: a) head and pronotum, dorsal; b) head and pronotum, lateral; c) head, frontal; d) stridulatory area, dorsal; e) stridulatory areas, left and right sides, dorsal; f) sternites; g) apex of abdomen, dorsal; h) apex of abdomen, ventral; i) apex of abdomen, posterior; j) apex of abdomen and ovipositor, lateral.

Municipality: Sierra Prieta (18°38'56"N - 69°58'22"W, 150 m a.s.l.); 12/September/2016; R. Teruel; 1 ♂ juvenile (green phase, SY, in ethanol 80%). Note. The complete sample from Pico San Juan was destroyed

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THREE NEW GENERA AND FOUR NEW SPECIES OF KATYDIDS FROM CUBA AND HISPANIOLA shortly after being collected, by an ant attack in the field. Fortunately, the specimens were photographed in situ when found, so the evidence remains (see figs. 17b, d herein).

Diagnosis. See genus diagnosis above.

Etymology. This species is named after Hatuey (birth date unknown, murdered by fire at stake in 1512), the well-known Taino cacique. He was the first rebel native to America and coincidently, he lived in the same two islands where this species occurs: Cuba and Hispaniola.

Table III. Measurements of type specimens of Unicorniella hatueyi n. gen., n. sp. from La Gran Piedra. Abbreviations: length (L), width (W), depth (H), not applicable (NA), not measured (NM).

Dimensions (mm) Male holotype Female paratype Head L / W / H 4.25 / 5.60 / 7.25 4.50 / 7.10 / 8.00 Interocular distance L 1.80 2.30 Eye diameter L 1.30 1.50 Scapus L / W 1.25 / 0.60 1.40 / 0.85 Pedicel L / W 0.90 / 0.43 0.80 / 0.55 Fastigium L / W / H 4.00 / 1.40 / 1.75 5.00 / 1.75 / 2.25 Pronotum L / W 7.45 / 6.10 7.65 / 7.00 Tegmen L / W 40.60 / 5.65 44.75 / 5.60 Abdomen L 15.25 17.40 Cerci L / W 1.75 / 1.05 NM Tergite X L / W 2.00 / 3.30 4.15 / 2.20 Supra-anal plate L / W 1.50 / 1.50 1.10 / 3.00 Subgenital plate L / W 2.25 / 2.65 2.25 / 2.40 Styli L / W 0.90 / 0.22 NA Ovipositor L / W NA 23.20 / 1.90 Profemur L / W 7.30 / 1.10 7.75 / 1.25 Protibia L 8.65 9.60 Mesofemur L / W 9.00 / 1.05 9.50 / 1.10 Mesotibia L 10.00 10.90 Metafemur L / W 22.90 / 2.50 24.75 / 2.55 Metatibia L 23.50 26.20 Body (total) L 26.95 29.55

Description of adult male holotype. Size large for the tribe (total length 26.95 mm). General coloration dark green, except as follows: a bright white stripe all along the sides of the body (from the base of fastigium through the lateral margins of pronotum and the costal margin of the tegmina), eyes white with a dark shadow, median ocellus white; antennae reddish brown, with black socket and scapus. Head with fastigium laterally and ventrally black, labrum and mandibles reddish purple, palps greenish white. Tegmina with the reticulate venation sharply contrasting in immaculate whitish. Legs pinkish to reddish brown, metafemur externally with small whitish to yellowish spots. See figures 7, 17a and table III. Head (fig. 8a–c). Large, wider than long (ratio = 1.3). Tegument smooth. Vertex flat in lateral view. Fastigium very large, very sharp and only slightly curved downwards; in lateral view with upper and lower surfaces basically straight; in dorsal view subtriangular; basal half smooth, base of fastigium ventrally with a very large tooth, vertically separated by a deep notch from a large apical tooth in frons. Eyes small and spherical. Median ocellus medium-sized, oval and translucent, located between antennal sockets. Frons very shallowly convex and glossy, lacking lateral edges. Genae convex in frontal view, lacking lateral carinae. Antennal sockets situated right between the eyes. Antennae standard for Conocephalinae, slightly surpassing the length of the tegmina; scapus subcylindrical, longer than wide (ratio = 2.0) and much shorter than

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Figure 9. Juvenile males of Unicorniella n. gen., full-body lateral view: a) paratopotype; b) specimen from Sierra Prieta (Dominican Republic). Scale bar in millimeters.

Thorax. Tegument glossy and densely rugose, with scattered minute setae. Pronotum longer than wide (ratio = 1.2), subrectangular in dorsal view, with dorsal disk truncate and strongly projected posteriorly; anterior margin shallowly concave, posterior margin shallowly convex and projected backwards, lateral lobes angled at 90°; longitudinal sulcus inconspicuous, anterior sulcus narrow but conspicuous, median and typical sulci inconspicuous. Tegmina and wings fully developed; tegmina slender, lanceolate and much longer than abdomen (ratio = 2.7); mirror large and subrectangular, longer than wide (fig. 8e). Wings not surpassing tegmina tip. Prosternum armed with a pair of relatively short and thin, conical spines. Basisternum of meso- and metasternum with lateral lobes (large, sub-trapezoidal and medially in contact, see fig. 8f). Legs (fig. 7). Very long and slender, covered with minute setae all over. Genicular lobes armed with a pair of apical spines, which are triangular, moderately long, sharp and pointing backwards. Procoxa dorsally with a large spine pointing forwards. Profemur subcylindrical, compressed; ventral pair of carinae

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THREE NEW GENERA AND FOUR NEW SPECIES OF KATYDIDS FROM CUBA AND HISPANIOLA strong and straight, separated by a deep sulcus, with 0:1 / 1:0 spines. Protibia slightly longer than profemur (ratio = 1.2), rectangular in cross-section; ventral pair of carinae slightly sinuose, with 5:6 / 6:5 spines; tympanal area slightly swollen, slits equal, relatively short, oval, and located dorsally at the base of protibia, below each slit there is a short but narrow pit. Mesofemur very similar to profemur, with 0:0 / 1:0 ventral spines. Mesotibia with 7:7 / 6:5 ventral spines. Metafemur slender, as long as metatibia (ratio = 1.0), oval in cross-section and with 5:8 / 8:7 ventral spines. Left metatibia with 18:21 and 9:8 subapical spines in its dorsal and ventral surfaces, respectively; right metatibia with 15:19 and 9:8, respectively; three pairs of apical spurs curved inwards: one dorsal, one lateroventral and one ventral (the smallest). Tarsi four- segmented, I and II long and of same length, III wider; claws longer than tarsi I + II, curved. Abdomen. Short and slender, subconical, tegument glossy, with minute setae scattered all over. Tergites without any modification. Tergite X (penultimate) wider than long (ratio = 1.6), with posterior margin widely V-shaped and sloping down. Supra-anal plate (fig. 8g, i) partially concealed under tergite X, rhomboidal, as long as wide (ratio = 1.0), and with a wide and deep median longitudinal furrow; tegument glossy to coriaceous, with minute setae scattered all over. Subgenital plate (fig. 8h) symmetrical, slightly wider than long (ratio = 1.2); tegument smooth and glossy, median keel only slightly perceptible in distal half; anterior margin shallowly convex with a shallow median concavity, lateral margins convex, posterior margin deeply V-notched, all margins covered with minute setae; styli relatively short (compared to subgenital plate) and densely covered with minute setae all over. Cerci large and thick (compared to supra- anal plate length, ratio = 1.6), densely covered with short and rigid setae, cylindrical and apically bifurcate, with each lobe ending in a strongly sclerotized, very sharp tooth (figs. 8 g–i).

Female (paratopotype). Very similar to male, differing as follows. Body larger (29.5 mm) and slightly more robust. Tegmina clearly surpassing ovipositor tip. Head with width/length ratio slightly greater (= 1.6). Pronotum as long as wide (ratio = 1.0). Profemur with 0:1 / 2:0 ventral spines. Protibia with 6:5 / 6:6 ventral spines. Mesofemur with 3:0 / 0:1 ventral spines. Mesotibia with 7:6 / 6:7 ventral spines. Metafemur with 7:8 / 5:8 ventral spines. Left metatibia with 13:19 and 8:6 subapical spines in its dorsal and ventral surfaces, respectively; right metatibia with 18:21 and 7:9, respectively. Tergite X (penultimate) longer than wide (ratio = 1.9). Supra-anal plate much wider than long (ratio = 2.7). Subgenital plate as long as wide (ratio = 1.0), convex and almost tectiform; anterior margin deeply V- notched, lateral margins shallowly concave, posterior margin paraboloid. Cerci short, conical and densely covered with short and rigid setae. Ovipositor (fig. 8j) longer than abdomen (ratio = 1.3), slender, straight and with edges entirely smooth (i.e., lacking any teeth, serrations or crenulations); tegument minutely and densely punctate; upper and lower margins basically straight and parallel, upper margin with tip shallowly curved downwards, lower margin with tip shallowly curved upwards.

Variation. As the majority of Copiphorini, this species also has a brown phase (figs. 17d, f). The pattern concurs with that described for the green phase, differing only in that the base color is a straw brown, the legs lack the pinkish to reddish shades and the tegmina are conspicuously spotted with dark to blackish brown.

Distribution (fig. 24). As for the genus (see above); see also Remarks section below.

Ecological notes. According to the available data, this species is restricted to very humid and mostly cold montane sites (fig. 21). The specimens collected personally by the present author were all found feeding on the fresh seeds of grass spikes, less than 1 m above the ground, mostly in open places such as forest clearings and sides of the roads, trails and sidewalks. According to the field notes of the collector (Rolando Teruel, pers. comm.), the adult female paratype from La Gran Piedra was captured in early evening with already complete darkness (19:10 hrs. EST), about 2 m up a white outer wall of a house, where it landed attracted by the white light coming through a glass window. The same collector also documented that the juvenile from Sierra Prieta (Dominican Republic) was found at dawn, clinging to a tall grass spike about 1.5 m above the ground, in an open roadside environment surrounded by pine forest.

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Figure 10. Adults of Unicorniella hatueyi n. gen., n. sp., from other collections: a) IES male from Nibujón; b) FZ female from Rangel.

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Figure 11. Adult male holotype of Anacaona bellatrix n. gen., n. sp., full-body views: a) dorsal; b) lateral; c) ventral. Scale bar in millimeters.

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Figure 12. Adult female paratype of Anacaona bellatrix n. gen., n. sp., full-body views: a) dorsal; b) lateral; c) ventral. Scale bar in millimeters.

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Figure 13. Adult male holotype (a–c, g–h) and adult female paratype (d–f) of Anacaona bellatrix n. gen., n. sp., close- up views: a) head and pronotum, dorsal; b) head and pronotum, lateral; c) head, frontal; d) head and pronotum, dorsal; e) head and pronotum, lateral; f) head, frontal; g) stridulatory area, dorsal; h) stridulatory areas, left and right sides, dorsal.

For a description of the ecological conditions at the type-locality, see above under Rombophora sierramaestrae n. sp. The other four localities known for this species have very similar conditions (fig. 21), despite their important differences in altitude.

Remarks. Unicorniella hatueyi n. sp. is distributed across the entire main island of Cuba, but it is extremely localized and scarce. A single, small outbreak of adults was observed in August/2015 at Pico San Juan, as well as an unusually greater abundance of late-instar juveniles and subadults in July/2018 at the type-locality (a possible prelude of another adult outbreak). But apart from this, only one or a few specimens have been obtained in all other positive collecting events (usually about a week long each, see Material Examined section above); even a previous one-week search by the present author on February/2015 at Pico San Juan, did not yield any specimens.

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The scant material examined from Cuba and Hispaniola appears to correspond with a single species occurring on both islands. Nevertheless, all populations known so far are highly isolated from each other (even three of them imply mountaintops) and the possibility that it could actually involve a complex of cryptic species cannot be discarded. This will be clarified only when additional adult specimens from Cuba and especially Hispaniola can be compared.

Figure 14. Adult male holotype (a–c) and adult female paratype (d–f) of Anacaona bellatrix n. gen., n. sp., respectively, close-up views: a) apex of abdomen, dorsal; b) apex of abdomen, ventral; c) apex of abdomen, posterior; d) apex of abdomen, dorsal; e) apex of abdomen, ventral; f) ovipositor, lateral.

Subfamily Pseudophyllinae Burmeister, 1838 Tribe Cocconotini Brunner von Wattenwyl, 1895 Genus Anacaona new genus Figures 11–14, 18, 20a, b, 25. Table IV Type species: Anacaona bellatrix n. sp., by present designation.

Diagnosis. Body medium-sized (26–27 mm) and relatively slender for the tribe. Both sexes macropterous; tegmina lanceolate, sexually dimorphic (shorter in male), surpassing tip of abdomen (male), but not reaching tip of ovipositor (female); hind wings fully developed. Head with tegument coarsely and densely punctate, lacking any keels. Fastigium short (slightly shorter than scapus) and blunt-conical. Eyes moderately small and spherical. Pronotum slightly longer than wide; dorsal disk moderately projected, truncate; tegument dorsally smooth, laterally feebly punctate; longitudinal, anterior and typical transverse sulci shallow and

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THREE NEW GENERA AND FOUR NEW SPECIES OF KATYDIDS FROM CUBA AND HISPANIOLA narrow. Prosternum armed with two conical spines. Legs moderately slender for the tribe. Male penultimate abdominal tergite (10th) not especially modified. Supra-anal plate in male widely triangular and with a deep longitudinal sulcus, in female rhomboidal and not sulcate. Subgenital plate very wide, symmetrical and with posterior margin deeply U-notched; in male much larger and with a pair of lateral sulci converging backwards. Male cerci curved inwards and armed apically with two sclerotized and strongly incurved spines; female cerci unmodified, broadly conical. Ovipositor very long, slender, with parallel edges, distally curved upwards and mostly smooth (except for a dorsobasal tooth).

Etymology. This genus is named after Anacaona (born in 1474, murdered by hanging in 1503), the well- known Taino woman that ruled the Jaragua chiefdom in Hispaniola and was also said to be the greatest Caribbean beauty. The generic epithet is declared here feminine in gender.

Figure 15. Live habitus of Erioloides santiago n. sp.: a–b) adult female holotype, in captivity; c) female holotype still subadult before last ecdysis, in captivity; d) juvenile female paratopotype, in captivity

Comparisons. Anacaona n. gen. most closely resembles two other Neotropical genera: Nesonotus Beier, 1960 and Sphaeropyga Beier, 1960. The new genus can be separated from Nesonotus as follows: 1) Face coarsely punctate and lacking lateral edges. 2) Pronotum dorsally smooth and laterally feebly punctate, with anterior and typical transverse sulci narrow and shallow. 3) Tegmina apically round and narrower, with venation not conspicuously raised. 4) Male cerci with two apical teeth. 5) Female subgenital plate sub-trapezoidal. 6) Ovipositor longer, wider and with edges essentially parallel. In Nesonotus, the face is closely wrinkled and has strong lateral edges, the pronotum is coarsely granulose and has deep anterior and typical transverse sulci, the tegmina are apically truncate and wider, with venation conspicuously raised, the male cerci have a single apical tooth, the female subgenital plate is pentagonal, the ovipositor is shorter, narrower and distally tapering. Last, Anacaona n. gen. can be separated from Sphaeropyga by: 1) Size smaller. 2) Face coarsely and densely punctate, lacking lateral edges. 3) Fastigium tip surpassing the antennal sockets. 4) Pronotum dorsally smooth and laterally feebly punctate. 5) Tegmina apically narrower. 6) Male penultimate abdominal

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YONG tergite (10th) flattened and not covering the cerci or tip of abdomen. 7) Male cerci with two apical teeth but otherwise unmodified. 8) Ovipositor (in Sphaeropyga the ovipositor is robust and almost straight). In Sphaeropyga, Beier (1960) recorded sizes of 34–42 mm in males and 37–43 in females, the face is feebly punctate and has lateral edges, the tip of fastigium just reaches the antennal sockets, the pronotum is densely granulose, the tegmina are apically wider, the male penultimate abdominal tergite (10th) is greatly bi-inflate (buttock-like) and covers completely the cerci and tip of abdomen like a helmet, the male cerci lack apical teeth but are highly modified (spatulate and basally with an internal tooth and two conspicuous blunt processes).

Distribution. Monotypic genus, known only from southern Hispaniola (Dominican Republic, see fig. 25).

Figure 16. Live habitus of Rombophora sierramaestrae n. gen., n. sp., in natural habitat: a) adult male from Aguada de Joaquín; b–c) adult female paratopotypes; d) juvenile female paratopotype. Photo a credit Eladio Fernández, modified from Navarro (2010: 304).

Anacaona bellatrix new species Figures 11–14, 18, 20 a, b, 25. Table IV

Types. DOMINICAN REPUBLIC: Barahona Province: Cabral Municipality: Sierra de Bahoruco: km 7–9 of road from Cabral to Polo (18°10'55"N - 71°15'10"W, 221 m a.s.l.); on the vegetation; 23/August/1987; L. F. de Armas, E. J. Marcano, A. Abud, D. Lantigua; 1♂ holotype (SY, in ethanol 80%). Pedernales Province: Oviedo Municipality: Los Tres Charcos (17°49'10.3"N - 71°26'15.1"W, 71 m a.s.l.); on the vegetation; 14/September/2016; R. Teruel; one adult female paratype (SY, in ethanol 80%).

Diagnosis. See genus diagnosis above.

Etymology. The selected epithet is a Latin word that literally means "she-warrior". It alludes to the remarkable aggressiveness displayed by individuals of this species when handled alive.

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Figure 17. Live habitus of Unicorniella hatueyi n. gen., n. sp.: a) adult male holotype, green phase, in captivity; b) adult male from Pico San Juan, green phase, in natural habitat (low grass); c) adult female paratopotype, green phase, in natural habitat (fern frond); d) adult female from Pico San Juan, brown phase, in natural habitat (low grass), see anomalous fastigium; e) juvenile male paratopotype, green phase, in natural habitat (bush twig); f) subadult male paratopotype, brown phase, in natural habitat (thin liana). Photos b–d courtesy of Tomás M. Rodríguez-Cabrera.

Description of adult male holotype. Size medium for the tribe (total length 27.3 mm). General coloration yellowish brown, with eyes and tip of mandibles and leg spines black, tegmina diffusely spotted with dark brown. See figures 11 and table I. Head (fig. 13a–c). Large, much wider than long (ratio = 1.8). Tegument glossy, coarsely and densely punctate, with minute setae scattered all over. Vertex shallowly convex in lateral view. Fastigium short,

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(slightly shorter than scapus, ratio = 0.8), conical but with blunt tip, dorsomedially incised by a very narrow and shallow longitudinal furrow. Eyes small and spherical. Median ocellus medium-sized, prominent, spherical and translucent, located between antennal sockets. Frons very shallowly convex, glossy, coarsely and densely punctate and lacking lateral edges. Genae convex in frontal view, lacking lateral carinae. Antennal sockets situated right between the eyes. Antennae standard for Cocconotini, much longer than tegmina; scapus subcylindrical, longer than wide (ratio = 1.5), slightly longer than fastigium, oval in cross- section and essentially glabrous; pedicel cylindrical, almost half the length of scapus.

Figure 18. Live habitus of female paratype Anacaona bellatrix n. gen., n. sp., in captivity.

Figure 19. Habitat of Erioloides santiago n. sp.: a) Puerto Manatí; b) Charco Mono; c) Siboney; d) Playa Verraco. Photo a courtesy of Rolando Teruel.

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Figure 20. Habitat of Anacaona bellatrix n. gen., n. sp. (a–b) and Rombophora sierramaestrae n. gen., n. sp. (c–d): a) Los Tres Charcos; b) road from Cabral to Polo; c) Aguada de Joaquín; d) La Gran Piedra. Photos a–c courtesy of Rolando Teruel.

Thorax. Tegument glossy, dorsally smooth and laterally feebly punctate, with minute setae scattered all over. Pronotum as long as wide (ratio = 1.0), subrectangular in dorsal view, with dorsal disk truncate and moderately projected posteriorly; anterior margin convex, posterior margin almost straight and projected backwards, lateral lobes shallowly sinuose and subquadrate; longitudinal sulcus inconspicuous, anterior and median sulci narrow but conspicuous, typical sulci shallow. Tegmina and wings fully developed, greatly surpassing abdomen tip. Tegmina slender, lanceolate and longer than abdomen (ratio = 1.7); mirror of left tegmen subtriangular, of right tegmen almost circular (fig. 13g–h). Wings not surpassing the tips of tegmina. Prosternum armed with a pair of medium-sized conical spines. Basisternum of meso- and metasternum with lateral lobes (medium-sized, subtrapezoidal, laterally expanded, and almost flat). Legs (fig. 11). Moderately slender for Cocconotini, covered with minute setae all over. Procoxa dorsally with a large spine pointing forwards. Profemur subquadrate, slightly compressed; ventral pair of carinae weak to moderate and straight, with 6:0 / 5:2 spines. Protibia slightly longer than profemur (ratio = 1.1), rectangular in cross-section; dorsal pair of carinae basically straight, ventral pair of carinae prominent and slightly sinuose, with 8:7 / 8:7 spines; tympanal area swollen, slits equal, large, wide, and located dorsally at the base of protibia. Mesofemur with 5:1 / 1:5 ventral spines. Mesotibia with 9:7 / 8:9 ventral spines. Metafemur robust, shorter than metatibia (ratio = 0.9), oval in cross-section and with 6:3 / 3:7 ventral spines. Left metatibia with 11:11 and 10:7 subapical spines in its dorsal and ventral surfaces, respectively; right metatibia with 12:10 and 5:9, respectively; genicular lobes armed with a pair of apical lateral spines; three pairs of apical spurs curved inwards: one dorsal, one lateroventral and one ventral (the smallest). Tarsi four-segmented, I long, II and III shorter and basically same-length; claws long and curved.

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Figure 21. Habitat of Unicorniella hatueyi n. gen., n. sp.: a) Sierra de Rangel; b) Pico San Juan; c) La Gran Piedra; d) Sierra Prieta. Photos a, d courtesy of Rolando Teruel.

Abdomen. Long and stout, subcylindrical, tegument glossy. Tergite X (penultimate) wider than long (ratio = 1.8), deeply bilobed and sloping down. Supra-anal plate (fig. 14a, c), concealed under tergite X, widely triangular, wider than long (ratio = 2.6); tegument glossy to coriaceous, with minute setae scattered all over; median-longitudinal sulcus deep. Subgenital plate (fig. 14b) large, symmetrical and as long as wide (ratio = 1.0); tegument coriaceous to glossy, with a pair of lateral sulci slightly convergent backwards; anterior margin concave, lateral margins shallowly convex, posterior margin deeply U-notched, all margins covered with minute setae; styli medium-sized, cylindrical, densely covered with minute setae all over. Cerci relatively short (clearly shorter than styli in ventral view) and stout, densely covered with short and rigid setae, cylindrical, curved inwards and apically bifurcate, with each lobe ending in a short, sharp tooth (figs. 14a–c).

Female (paratype). Very similar to male, differing as follows. Body larger (26.25 mm) and slightly less robust. Coloration with only minor differences (compare figs. 12 to 18), attributable to differences in preservation time (coloration of fresh female is more reliable): tegmina with infuscate pattern darker, denser and more extended, with venation contrastingly whitish; legs femur externally spotted with pale yellowish dots; ovipositor reddish brown, basally paler and apically much darker. Head with width/length ratio slightly lower (= 1.6). Legs longer. Tegmina longer, much longer than abdomen but not reaching ovipositor tip. Profemur with 0:5 / 5:0 ventral spines. Protibia with 7:8 / 8:7 ventral spines. Mesofemur with 5:0 / 1:5 ventral spines. Mesotibia with 10:7 / 8:9 ventral spines. Metafemur with 7:2 / 2:5 ventral spines. Left metatibia with 10:11 and 11:7 subapical spines in its dorsal and ventral surfaces, respectively; right metatibia with 12:11 and 7:9, respectively

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Tergite X (penultimate) much longer than wide (ratio = 2.8). Supra-anal plate rhomboidal, slightly wider than long (ratio = 1.2). Subgenital plate (fig. 14e) subtrapezoidal, wider than long (ratio = 2.2) and slight convex; tegument coriaceous and glossy; anterior margin shallowly convex, lateral margins almost straight and convergent backwards, posterior margin narrowly and deeply U-notched. Cerci short, thickly conical and densely covered with short and rigid setae. Ovipositor (fig. 14f) very long (longer than abdomen, ratio = 1.5), slender, with parallel edges, coriaceous, distally curved upwards and largely smooth (except for a coarse, blunt dorsobasal tooth).

Table IV. Measurements of both type specimens of Anacaona bellatrix n. gen. n. sp. from Dominican Republic. Abbreviations: length (L), width (W), depth (H), not applicable (NA), not measured (NM).

Dimensions (mm) Male holotype Female paratype Head L / W / H 4.10 / 7.20 / 7.50 4.40 / 7.25 / 6.65 Interocular distance L 2.75 2.65 Eye diameter L 1.65 1.85 Scapus L / W 1.40 / 0.90 1.40 / 0.90 Pedicel L / W 0.75 / 0.55 0.85 / 0.55 Fastigium L / W / H 1.25 / 1.15 / 1.25 1.35 / 1.15 / 1.10 Pronotum L / W 8.25 / 7.75 8.35 / 8.15 Tegmen L / W 26.00 / 4.85 32.00 / 5.90 Abdomen L 15.00 13.50 Cerci L / W 1.70 / 0.95 1.35 / 0.60 Tergite X L / W 2.35 / 4.20 1.30 / 3.60 Supra-anal plate L / W 1.30 / 3.40 1.00 / 1.20 Subgenital plate L / W 3.50 / 3.80 1.55 / 3.50 Styli L / W 1.55 / 0.40 NA Ovipositor L / W NA 19.90 / 1.50 Profemur L / W 7.00 / 2.20 7.50 / 2.50 Protibia L 8.00 9.10 Mesofemur L / W 7.25 / 2.10 8.50 / 2.50 Mesotibia L 9.05 9.75 Metafemur L / W 15.75 / 4.80 19.00 / 5.40 Metatibia L 18.05 19.65 Body (total) L 27.35 26.25

Figure 22. Geographical distribution of Erioloides santiago n. sp.: type locality (red symbol), additional localities (yellow symbols). Image frame = 500 x 220 km, inset = 20 x 13 km.

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Figure 23. Geographical distribution of the genus Rombophora n. gen. and its single species Rombophora sierramaestrae n. gen, n. sp.: type locality (red symbol), additional localities (yellow symbols). Image frame = 500 x 220 km, inset = 20 x 13 km.

Distribution (fig. 25). As for the genus (see above).

Ecological notes. According to the field notes of its collector (Rolando Teruel, pers. comm.), the female paratype was found at noon, hidden deep inside a hollow horizontal branch of a tree, about 2 m above the ground. It was resting with the head pointing outside, in close proximity to several adults and juveniles of the similarly-sized scorpion Centruroides lucidus Teruel, Armas & Kovařík, 2015 (Buthidae). Another very similar case of interspecific tolerance at rest between a Pseudophyllinae tettigoniid and a different species of the same scorpion genus (Centruroides Marx, 1889) was already found in Mexico (Yong, 2018: 98).

General remarks

This contribution increases the Copiphorini genera known to occur in Cuba from four to seven, and those recorded from Hispaniola from two to four (see a habitus comparison of all these genera in fig. 26): 1. Caulopsis Redtenbacher, 1891: Cuba. 2. Erioloides Hebard, 1927: Cuba and Hispaniola (Dominican Republic). 3. Eriolus Bolívar, 1888: Cuba. 4. Neoconocephalus Karny, 1907: Cuba and Hispaniola (Haiti and Dominican Republic). 5. Pyrgocorypha Stål, 1873: Cuba and Hispaniola (Dominican Republic) 6. Rombophora n. gen.: Cuba. 7. Unicorniella n. gen.: Cuba and Hispaniola (Dominican Republic).

On the other hand, the tribe Cocconotini is here recorded for the first time to occur in Hispaniola. The description of Anacaona n. gen. raises to four the genera of Pseudophyllinae from this island, together with Caribophyllum Rehn, 1947 (Pterophyllini Karny, 1925), Polyancistrus Serville, 1831 and Spelaeala Rehn, 1943 (both Polyancistrini Brunner von Wattenwyl 1895). All four genera are endemic to this insular territory.

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Figure 24. Geographical distribution of the genus Unicorniella n. gen. and its single species Unicorniella hatueyi n. gen., n. sp.: type locality (red symbol), additional localities (yellow symbols). Image frame = 500 x 220 km, inset = 20 x 13 km.

Figure 25. Geographical distribution of the genus Anacaona n. gen. and its single species Anacaona bellatrix n. gen., n. sp.: type locality (red symbol), additional locality (yellow symbol). Image frame = 500 x 220 km, inset = 20 x 13 km.

Acknowledgements First, I am greatly indebted to Rolando Teruel (Grupo de Sistemática y Ecología de Artrópodos Caribeños, Havana, Cuba) for his invaluable field assistance and all the help given while writing this paper: preparation of images, language and style correction, expert advice on taxonomic questions and useful comments during all stages of the manuscript. This contribution is another outcome of the research project "An inventory of the Orthoptera of the Eastern Sierra Maestra Range, Cuba", funded by an OSF Grant Application from the Orthopterists' Society. Also, I am also very grateful to Idea Wild for the logistic support (especially the photographic equipment). I really appreciate the help of Elier Fonseca (Universidad de la Habana, Cuba), Yoendri Ricardo, Jorge L. Reyes, Abelardo A. Méndez and José Ramón Fuentes (all from Centro Oriental de Ecosistemas y Biodiversidad, Santiago de Cuba) and Tomás M. Rodríguez-Cabrera (Havana), for the field assistance and donating important specimens. Holger Braun (Museo de La Plata, Argentina) and Frederic Schramm (Kista, Sweden), kindly translated Beier's descriptions (originally in German) into English for me. Holger Braun, Sylvain Hugel (Strasbourg University, France) and Lianna Teruel (Riobamba, Chimborazo, Ecuador), kindly provided pertinent literature. Two anonymous reviewers gave useful suggestions to the manuscript.

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Figure 26. Genera of Copiphorini from Cuba and Hispaniola: a–b) Eriolus; c–d) Rombophora gen. n.; e–f) Erioloides; g–h) Caulopsis; i–j) Pyrgocorypha; k–l) Unicorniella gen. n.; m–n) Neoconocephalus. Photo c credit Eladio Fernández, modified from Navarro (2010: 304).

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References

Cigliano, M. M., Braun, H., Eades, D. C. & Otte, D. (2019) Orthoptera Species File Online. Version 5.0/5.0. Available at: http://orthoptera.speciesfile.org/Home-Page/Orthoptera/HomePage.aspx [accessed 22/February/2019]. Naskrecki, P. de (2000) Katydids of Costa Rica. Vol 1. Systematics and bioacoustics of the cone-headed katydids (Orthoptera: Tettigoniidae: Conocephalinae sensu lato). The Orthopterists' Society at the Academy of Natural Science of Philadelphia, 164pp. Navarro, N. (2010) Región Oriental. In: Viña Bayés, N. (Ed.), Cuba. Un encuentro fotográfico con su naturaleza. Graphicom, Italy, pp. 252–361. Yong, S. (2018) Updating the taxonomy and distribution of Liparoscelis pallidispina Stål, 1873 (Orthoptera: Tettigoniidae: Pseudophyllinae). Ecologica Montenegrina, 17, 80–99. Yong, S. & Perez-Gelabert, D. E. (2014) Grasshoppers, crickets and katydids (Insecta: Orthoptera) of Cuba: an annotated checklist. Zootaxa, 3827(4), 401–438. Zayas, F. de (1976) Entomofauna Cubana. Tomo III. Introducción a la sección Polyneoptera. Editorial Científico-Técnica, Havana, pp. 1–130.

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