Copiphora, Serville, 1831

Piotr Naskrecki, 2000, Katydids of Costa Rica / Vol. 1, Systematics and bioacoustics of the cone-head katydids (Orthoptera: Tettigoniidae: Conocephalinae sensu lato)., Philadelphia, PA: The Orthopterists Society at the Academy of Natural Sciences of Philadelphia, : 28-30

COPIPHORA Serville, 1831

Common name: Spear bearers

1831 Serville, Ann. Sci. nat. Paris 22:147; type species: Locusta cornuta De Geer, 1773

1999 Naskrecki and Otte, Illustr. Cat. Orthop. I (CD ROM) >>references; types illustrated

[syn.] Copidophora = 1846 Agassiz, Zool. Nomencl. Ind. Univ.: 98

[syn.] Copiophora = 1838 Burmeister, Handb. Ent. 2: 702

Diagnosis

Slender to robust; both sexes macropterous ( Figs. 5 A, 9A). Head with fastigium of vertex strongly produced, 2-3.5 times as long as diameter of eye, sharply conical ( Fig. 5 A) or laterally flattened ( Fig. 8 C), rarely fastigium short ( Fig. 9 A). Head robust, frons flat or weakly convex; genae with moderately to strongly developed lateral carinae; eyes globular, moderately protruding. Mid tibia armed dorsally. Male cerci thick, blunt apically, with one or two inner spines; ovipositor straight or weakly downcurved, sometimes extremely long.

Description (male except where specified)

Head.— Fastigium of vertex strongly produced, 1.5 to 5.5 times as long as diameter of eye ( Figs. 5 A, 8C, 9A); fastigium sharply conical, laterally flattened, or dorsoventrally expanded subapically; rarely fastigium short, trituberculate; fastigium usually with well developed ventral keel; dorsal surface of fastigium smooth or tuberculate; fastigium without well developed lateral carinae; lateral ocelli well developed, sometimes very prominent, situated on large protuberances; apex of fastigium sharply tapered or with small hook, rarely blunt; basal portion of fastigium of vertex with a prominent knob ventrally, well separated from fastigium of frons. Antennal sockets separated by distance equal to 0.6-1.0 diameter of eye. Eyes globose, moderately projecting; frons flat or weakly convex; genae with well developed, granulose lateral carinae. Labrum and mandibles symmetrical.

Thorax and wings.— Dorsal surface of pronotum smooth to weakly rugose, flat or metazona weakly raised; anterior margin of pronotum straight, posterior margin straight or weakly convex; lateral lobes with posterior angle rounded, with weakly developed humeral sinus. Thoracic auditory spiracle large, elliptical, completely hidden under lateral lobes of pronotum. Prosternum unarmed, mesosternum with a pair of small, widely separated, vertical spine-like lobes; metasternum unarmed.

Wings in both sexes fully developed, surpassing apices of hind femora . Stridulatory apparatus of male well developed; stridulatory file straight or weakly curved ( Figs. 40 B-G, 43A-C), teeth variable, from thin and wide to thick and narrow, sometimes both types of teeth present in file; stridulatory area of left wing covered with secondary venation; mirror of right wing approximately rectangular to nearly circular, with or without thin veinlet parallel to AA 1. Posterior margin of front wing straight or weakly convex; apex of front wing narrowly to broadly rounded.

Legs.— Fore coxa with an elongate, sometimes laterally flattened, forward projecting spine dorsally; middle and hind coxa without spine; all trochanters unarmed. All femora unarmed dorsally but with prominent spines on anterior ventral margins, posterior ventral margins unarmed; sometimes femoral spines large, triangular, nearly lobe-like; genicular lobes of all femora armed with long spines, sometimes posterior lobes of fore and/or mid femora unarmed. Front tibia unarmed dorsally, both ventral margins with short, immovable spines; tympanum on fore tibia bilaterally closed, tympanal slits facing forward, tympanal area moderately swollen; middle tibia armed dorsally with 1-3 dorsal spines, ventrally armed on both margins; hind tibia armed on all four dorsal and ventral margins; apex of hind tibia with two pairs of ventral and one pair of dorsal movable spurs.

Abdomen.— Dorsal surface of abdominal terga smooth, unmodified. Male 10th tergite with surface smooth, its posterior margin weakly to distinctly emarginated; supraanal plate in both sexes small, broadly rounded apically; male cercus stout, blunt apically, usually with one, blunt, subapical internal spine ( Figs. 5 E-F, 6D-E); sometimes apex of cercus with sharp, heavily sclerotized, triangular lobe ventrally ( Fig. 7 D); paraprocts unmodified; female cercus, simple, narrowly conical. Subgenital plate of male with a very shallow apical emargination; styli short, 1.5-2.0 times as long as thick, to very short, peg-like; female subgenital plate with triangular apical incision or nearly straight apically. Male internal genitalia with weakly sclerotized inner margins of phallic lobes but without well developed titillators ( Figs. 5 D, 7E). Ovipositor narrow, straight or slightly downcurved, its margins parallel; both dorsal and ventral margins of ovipositor smooth; ovipositor often exceedingly long, ratio ovipositor/hind femur 1.4 - 3.5.

Coloration.— General coloration light to dark green ( Figs. 36 E, G); facial markings variable and often species-specific; fastigium of vertex often bright yellow.

Dorsum of the head and pronotum sometimes with darker, brown or reddish markings. Sometimes posterior margins of tegmina red, resulting in distinct red dorsal stripe; stridulatory file and femoral spines often contrastingly white.

Remarks.— The genus Copiphora currently includes 24 described species, distributed throughout Central and South America [ C. subulata (Stoll) described from South Africa undoubtedly belongs to a different genus]. Two additional new species and one subspecies from Costa Rica are described below.

Members of the genus Copiphora are some of the most spectacular katydids in Central America. Some species of the genus possess large, horn-like fastigia, which gave rise to the common name “ rhinoceros katydids.” Females of many species also possess some of the longest ovipositors among the Tettigoniidae , and for this reason I propose a common name “spear bearers” for such species.

All Costa Rican species of Copiphora are forest insects, occurring primarily in the canopy and high understory levels. Copiphora rhinoceros , C. cultricornis , and C. hastata frequently descend to low, shrubby vegetation, and females of probably all species come to the ground to lay eggs. At La Selva Biological Station, females of C. rhinoceros have been repeatedly observed laying eggs in the fronds at the base of the palm Calyptrogyne ghiesberghiana (Linden and H.A. Wendl.) H.A. Wendl. Copiphora brevicauda costaricensis seems to be restricted to the canopy level and has never been seen on lower vegetation. All species of the genus are strictly nocturnal. They feed on a variety of organic material, including seeds, fruits, and insects. Copiphora hastata seems to specialize on hard seeds but has also been seen feeding on flowers and fruits of a palm Calyptrogyne ghiesberghiana , caterpillars, and egg clutches of a frog Agalychnis callidryas (Cope) . The closely related C. cultricornis has not been seen feeding in the wild, but in captivity individuals of this species accepted both insects (although they seemed incapable of catching live, fast moving insects) and various fruits, seeds, and vegetables (including leaves of lettuce, not accepted by any other species of the genus). Copiphora rhinoceros has been seen feeding on fruits of Psychotria sp. ( Rubiaceae ), C. ghiesberghiana , unidentified alate seeds of several tree species, several species of caterpillars (including ones covered with dense, urticating hairs), snails ( Fig. 36 G), other katydids, and even small lizards ( Norops sp.). Copiphora brevicauda costaricensis seems to be primarily predaceous, actively hunting and capable of capturing even such fast moving prey as cockroaches. Upon seeing an insect, they jump on it, using the first two pairs of legs to grasp the victim, at the same time inflicting damaging bites with their extremely powerful mandibles.

Figures

FIG. 5. Copiphora rhinoceros. A. male - habitus, B. male face, C. male head and pronotum, dorsal view, D. titillators, posterior view (everted), E. male cerci, posterior view, F. male cerci, dorsal view.

FIG. 7. Copiphora hastata. A. male - habitus, B. male face, C. male head and pronotum, dorsal view, D. titillators, posterior view (everted), E. male cerci, posterior view, F. male cerci, dorsal view.

FIG. 8. Costa Rican Copiphora spp. A-D: C. cultricornis; A. male face, B. male head and pronotum, dorsal view, C. male head and pronotum, lateral view, D. male cerci, dorsal view; E-H: C. brevicauda costaricensis; E. male head and pronotum, lateral view, F. male face, G. ovipositor, H. male cerci.

FIG. 9. Copiphora ottei. A. male - habitus, B. male face, C. male head and pronotum, dorsal view, D. male cerci, dorsal view.

FIG. 36. Costa Rican species of Conocephalinae s. l. A. Conocephalus angustifrons - male, B. Caulopsis cuspidata - female, C. Acantheremus colwelli - male, D. Vestria punctata - male and female, in copula, E. Copiphora hastata - male, F. Podacanthophorus vargasi - female, G. Copiphora rhinoceros - male eating a snail, H. Subria sylvestris - male, brown form.

FIG. 40. Stridulatory files of species of Conocephalus. A. C. magdalenae, B. C. ictus, C. C. cinereus, D. C. fasciatus.