Studies in Australian : The Mecopodine Katydids Part 2 (: Tettigoniidae; ; Sexavaini) Queensland Palm Katydid Author(s) :D. C. F. Rentz, You Ning Su, Norihiro Ueshima Source: Transactions of the American Entomological Society, 132(3):229-241. 2006. Published By: The American Entomological Society DOI: URL: http://www.bioone.org/doi/ full/10.3157/0002-8320%282006%29132%5B229%3ASIATTM %5D2.0.CO%3B2

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Studies in Australian Tettigoniidae: The Mecopodine Katydids Part 2 (Orthoptera: Tettigoniidae; Mecopodinae; Sexavaini) Queensland Palm Katydid

DCF RENTZ, YOU NING SU AND NORIHIRO UESHIMA

[DCFR] 19 Butler Dr, Kuranda, Queensland, Australia 4881 [email protected] [YNS] 90 Mugga Way, Red Hill, Australian Capital Territory 2603 [email protected] [NU] 1435-1 Kubo-cho, Matsusaka, Mie 515-004, Japan [email protected]

ABSTRACT

Two tribes of Mecopodinae (Mecopodini and Sexavaini) are represented in Australia. Both are confined to the tropics. The Mecopodini have been covered recently (see Rentz et al., 2006) and that group comprises a single genus with four species that live in grassy situations, often adjacent to rainforests. The Sexavaini is represented by a single species of Segestidea I. Bolívar, a genus with several species to the north of Australia. Some Segestidea species are well known to cause damage to coconuts. The single species, S. queenslandica, the Queensland Palm Katydid, is apparently facultatively parthenogenetic and is described with notes on its biology and distribution.

SEXAVAINI: Karny 1924

This is a contuation of the Monograph of the Australian Tettigoniidae begun by Rentz (1985, 1993, 2001). A recent contribution in this respect was Rentz et al. (2005), a revision the pseudophylline genus Phricta Redtenbacher in Austra- lia. Approximately 70% of the Australian tettigoniid fauna remains to be described. The Sexavaini of the Mecopodinae has been informally dealt with in the literature as “Group Sexavae”. (See Willemse 1977), for example. It is a grouping of at least three genera of large katydids that are known in the South Pacific as Coconut Treehoppers. They are major pests of coconut and oil palm, but some species also feed on banana, karuka, manila hemp, and sago palm (see Young, 1985). Of the three genera included in this tribe, only one, Segestidea I. Bolívar, is known from Australia. The other genera are Stal (12 species) and Sexava Walker (3 species). Segestidea is known from some 10 species. F. Willemse (1977) provides the latest review of the group but did not know about the Austra- lian species. L. Willemse (2001) reviewed the economic status of species in the tribe. The Australian species is not known to cause economic damage. The species of Segestidea described here has been known to the authors for many years (see specimens examined section). An interesting property of this species is that it is facultatively parthenogenetic. Only a single male has been found in the field. However, captive virgin females in the first author’s lab laid eggs that hatched and developed into females (see specimens examined). Re- 230 AUSTRALIAN MECOPODINAE: 2

cently Alan Henderson of the Melbourne Zoo reported (in litt.) that a female he had collected on the Morris Dam road to Copperlode Dam, near Cairns, Queensland, produced eggs that eventually hatched into females. At this writing the nymphs continue to mature. Parthenogenicity is known in one Australian rainforest species, the , Phricta spinosa (Redtenbacher), (see Rentz et al. 2005). Parthenogenicity is rare in the Tettigoniidae. The reasons for this odd biological strategy in rainforest species are presently unclear.

SEGESTIDEA I. Bolîvar

Segestidea I. Bolîvar, 1903: 166. Eumossula C. Willemse, 1957: 38. (Type species: Eumossula gracilis C. Willemse, by monotypy. Type species: Segestidea princeps I. Bolîvar,1903 designated by Kirby (1906: 359) (= Segestidea novaeguineae (Brancsik, 1897).

As noted above, this tribe was considered as a “group” by F. Willemse (1977) with the suggestion that the subfamily was in serious need of revision. To date, there has been no taxonomic revision of this group. Distinction among the three genera of the subfamily was based on the presence or absence of a posterior dorso- apical spine on the fore and middle tibiae along with the comparative measure- ments of the body, tegmen, head and pronotum. In Sexava and Segestidea the apex of the fore and middle tibia bears a small spine on the dorsal margin of the posterior side (Fig. 9). In Segestes, the dorso-apical spines are absent. (A small number of specimens prove to be the exception!) Comparison of the male teg- mina of Sexava, Segestidea and Segestes demonstrate that the tegmina overlap basally for a short distance in Segestes and Segestidea but in Sexava the overlap is conspicuous for almost the entire length. Details of the differences on the tegminal venation and its probable effect on the stridulatory mechanism can be found in Willemse (1977). If the Australian examples of Segestidea are keyed in Willemse’s paper (1977), they key near rufopalpis (Willemse). The Australia species of Segestidea has a relatively small geographic range, occurring in rainforest south of Cairns and north to the Daintree and west in the Kuranda Range and at localities on the Atherton Tableland. Where it occurs, the species is fairly common and a well-known garden resident where palms are grown. Elsewhere the genus is confined to Obi (Moluccas), mainland Papua New Guinea and the Louisade and Bismarck Archipelagoes.

Segestidea queenslandica Rentz, Su & Ueshima, sp. nov. Figs. 1-14; Table 1 Queensland Palm Katydid

ANIC Number Segestidea sp. 1

Holotype male.—Label data: “Segestidea sp. male Tettigoniidae: Sexavaini upland rainforest 16°44’00"S 145°33’55"S Black Mt. Road, Kuranda (335m elev.) NQ March 2003 C. Wilson”. Holotype in Australian National Collection, Canberra. Size large for genus, form robust (Table 1). Head protruding well in front of RENTZ, SU, AND UESHIMA 231 pronotum (Figs 1, 2, 5-7), strongly slanting; occiput smooth, fastigium with short, prominent spike protruding only slightly beyond extended antennal scape; eye small, round; frons smooth, flat, but with minute wrinkles laterally; clypeus pro- truding. Mandible (Fig. 12) unique, short, robust, incisor not produced as in other tettigoniids, but seemingly modified as a grinding structure. Antennal scape cy- lindrical, somewhat constricted in the middle, pedicel short, cylindrical, about 1/ 3 length of pedicel; flagellum thickened in proximal 1/3, tapering to apex, total length slightly longer than length of itself. Pronotum (Fig. 7) weakly pro- duced forward in the middle; surface of prozona and metazona similarly punctate and rough, surface with 2 prominent transverse shallow sulci; lateral lobe (Fig. 6) with ventral margin only feebly produced. Prosternum with a pair of elongate, widely spaced spikes; meso and metasterna cup-shaped. Tegmen (Fig. 8A) with stridulatory region highly modified with several quadrate cells; costa, short about 1/3 length of tegmen itself, subcosta and radius parallel, flaring apically, media extending nearly to apex of tegmen, Cu1 extending nearly to apex of tegmen; stridulatory file (Fig. 14) straight, teeth broad. Forelegs elongate, tibia quadrate in cross-section, feebly flaring apically, dorsal surface with 3 spines on each side, positioned in the middle 1/3, apex with a spine on each side, ventral surface with 10 spines on each side and an apical spine on each side; tarsus elongate, third article flap-like; fore femur laterally compressed, not modified, with a single spine on the internal margin of the ventral surface, positioned in apical 1/4; middle leg short then fore leg, tibia as in fore tibia, somewhat expanded at proximal end with surface sulcate, carinae more prominent near apex, ventral surface sulcate, with 10 spines on each side, apical spines not much larger than those adjacent, dorsal surface 2 spines on anterior margin, 3 spines on posterior margin and a prominent apical spine; femur without armature, surface smooth. Hind leg with femur not surpassing tegmina at rest; femur smooth, ventral surface with 4 spines on each side; tibia with apex expanded, dorsal surface with many prominent elongate spines, positioned along entire length, on each side; ventral surface with a sparse number of spines; apex with a pair of elongate spines on each surface. Genicular lobes of all femora with an elongate spine on each side. All tarsi elongate, apical segment with prominent claws. Abdomen smooth, without modification; cercus (Fig. 13) simple, feebly incurved, without internal tooth; subgenital (Fig. 13) with shallow V-shaped median incision, styles curved, less than the length of one side of incision. No sclerotised internal genital structures. Female.—Size much larger than male (Table 1) tegmina not dorsally accen- tuated as in male (Fig. 8B). Subgenital plate (Fig. 10) simple, with shallow me- dian incision. Cercus elongate, apex narrowed, abruptly curved inwards. Ovi- positor short, (Table 1), straight, without any armature. Type Locality.—The type locality is a patch of rainforest, near the residence of the first author, in which there is a variety of introduced and native palms, the dominant native palms of which are Lawyer Palms, Calamus spp. The collector of the male holotype states that it was brought to his attention by its loud stridula- tions on understorey vegetation where it was found in the company of a female. Nymph.—Early instars are pale white, seemingly resembling bird droppings (Fig. 2). At the second instar stage, a few speckles appear on the dorsal surface of the insect. A distinctive divided, subapical dark brown spot on the dorsal surface of the hind femur appears at the second instar stage. Following the second instar stage, a spot appears at the base of the hind femur. 232 AUSTRALIAN MECOPODINAE: 2

Egg.—Egg elongate, (Fig. 11) more acutely pointed at one end than the other; surface minutely evenly rugose and bearing a longitudinal groove, on both sides, extending for the full length of the egg itself; pointed end with a minute hook, the direction of the hook relative to the groove arbitrary; micropyles located within the groove on the bowed side. Five eggs measured (in mm) averaged 9.2 x 1.8. Specimens Examined.—Paratypes, Queensland: 15O50’S. 145°20’E., Gap Creek, 5 km ESE. Of Mt Finnigan, nr Cooktown, 13-16v.1981 (DCF Rentz, stop 37, 1 female, ANIC). 16°13’S. 145°25’E., 5 km W. by S. of Balley Pt., “Cow Bay Hotel”, 1.xi.1993 (DCF Rentz, L. Lowe, stop 10, 1 female, died as last instar 12.i.1994, ANIC). 17 O00’S. 145°50’E., Pine Creek, nr CSIRO Tower, 11 km SE by E. of Cairns, 88.iii.1988 (DCF Rentz stop A-18, 1 female, reared from eggs laid in lab, hatched 21.vii.1988, matured 1.v.1989, ANIC). 16°48’S. 145°38’E., Kuranda (Top of the Range) 19 Butler Dr., 15.iv.2003 31.iii.2004, 15.ii.2006, 28.ii.2006, 15.iii.2006 31.iii.2006 (DCF Rentz, 9 females, 1 prey of Black Butcherbird, Cracticus quoyi ANIC, DPIM). Kuranda, 23.xii.1982 (N. Quick, 1 female, ANIC). 16°45’S. 146O37’E., Kuranda, 2 km SW on Kennedy Hwy, (“Arona”), 3.i.2005 (DCF Rentz, 1 female, ANIC). 16°45’S. 145O36’E., Black Mtn Rd., 11.4 road km from Kennedy Highway, 8 km NNW of Kuranda, 19.ii.1988 (DCF Rentz, stop A-27, 2 females, ANIC). 17°07’S. 145°38’E., Mt Haig Rd., 10 km ENE of Tinnaroo Dam Wall, Atherton Tableland, 3.ii.1988 (DCF Rentz, stop A-14, 1 female, ANIC). 17°28’S. 146°0’E., Polly Creek, Garradunga, nr Innisfail, 19.i.1998 (J. Hasenpusch, 1 adult female, 2 female nymphs, matured in lab. on 22.x.1998, ANIC); same locality & collector, 18.i.1999 (1 female, DPIM); same locality & collector (1 second last instar 12.xii.2003. Paratypes will be deposited in the Australian National Insect Collection, Canberra (ANIC), the Queensland Museum Brisbane, Dept of Primary Indus- tries, Mareeba (DPIM), Queensland, the Academy of Natural Sciences of Phila- delphia and the California Academy of Sciences, San Francisco. Discussion.—S. queenslandica is not easily confused with any other tettigoniid in Australia. It ranks among the largest of Australian katydids with only some females of the pseudophylline, Phricta spinosa (Redtenbacher) being larger. Curiously, it shares with some Phricta species the reddish ventral surface of the hind femur. It has not been observed to flip up and display the hind legs when alarmed as do Phricta species. In fact, it shows little defense when it is encountered. S. queenslandica is a nocturnal species adults of which spend the daylight hours hidden in foliage where they remain motionless and resembling dead leaves. After dark the katydids emerge to feed on a variety of palms includ- ing Lawyer Palms (Calamus spp.) (Fig. 4) and a range of introduced ornamental palms. At no time has this katydid been seen to cause serious damage to the host plants. It does not generally occur outside rainforest and is observed in gardens only if they are adjacent to natural rainforest. S. queenslandica also shares with P. spinosa facultative parthenogenicity. Both live in similar habitats and indeed are sympatric and synchronic in many localities. Females of both, although arboreal for most of their lives, must de- scend to the ground at night to oviposit. At this time they are probably vulnerable to predation. Could this be the explanation for such an unusual reproductive strat- egy? Or will parthenogenicity be found to be more widespread amongst rainforest Tettigoniidae? At this point we know very little about the biology of other species in other higher taxa. RENTZ, SU, AND UESHIMA 233

The shape of the mandible of S. queenslandica (Fig. 12) is quite different from most tettigoniids. Gangwere (1965) surveyed the orthopteroid relat- ing mandible shape with feeding habits, where known. Rentz (1985, 1993 and 2001) illustrated the mandibles of many Australian tettigoniids and attempted to relate their shape to diet. The mandible of this species lacks an elongate incisor. Instead it is blunt and more of a grinding structure. This is probably directly re- lated to the tough structure of the palm leaves it eats. . ACKNOWLEDGEMENTS

We would like to thank Mr. Michael Cermak, owner of the single known male specimen of the species, for permitting its description and subsequent placement in the Australian National Insect Collection, CSIRO, and Canberra. He contributed the photograph of the living holotype. We also thank the CSIRO for permission to study material in their care and are grateful to Mr Alan Henderson, Melbourne Zoo, for his comments on the culture of the species in the living collection of the zoo.

REFERENCES

Gangwere, S. K. 1965. The structural adaptations of mouthparts in Orthoptera and allies. Eos, Revista Espanola de Entomologia 41: 67-85. Rentz, D. C. F. 1985. A monograph of the Tettigoniidae of Australia.. Volume 1. The Tettigoniinae. CSIRO Pub. East Melbourne. Rentz, D. C. F. 1993. A Monograph of the Tettigoniidae of Australia. Volume 2. The Phasmodinae, Zaprochilinae and Austrosaginae. CSIRO Pub. East Melbourne. Rentz, D. C. F. 2001. Tettigoniidae of Australia. Volume 3, The Listroscelidinae, Tympanophorinae, Meconematinae and Microtettigoniinae. CSIRO Pub. Collingwood, Vic. Rentz, D. C. F., Su, Y. N., Ueshima, N. 2005. Studies in Australian Tettigoniidae. The genus Phricta Redtenbacher (Orthoptera: Tettigoniidae; Pseudophyllinae; Phrictini) Transactions of the American Entomological Society, 131: 131-158. Willemse, F. 1977. Classification and distribution of the Sexavae of the Melanesian subre- gion (Orthoptera, Tettigonioidea, Mecopodinae) Tijdschrift voor Entomologie, 120: 213-277. Willemse, L. P. M. 2001. Guides to the pest Orthoptera of the Indo-Malayan region. Fauna Malesiana Field and Study Guide series. Pp. 1-150. Backhuys publ., Leiden, Nether- lands. Young, G. R. 1985. Observations of the biology of Segestes decoratus Redtenbacher (Or- thoptera, Tettigonioidea, Mecopodinae). General and Applied Entomology, 17: 57- 64. 234 AUSTRALIAN MECOPODINAE: 2

Table 1. Measurements (in mm) of Segestidea queenslandica Rentz & Su, sp. nov.

Locality Length Length Width Length Length Length body Pronotum Pronotum Tegmen Hind Ovipositor Femur

Holotype male 48.7 7.9 5..1 61.7 39.4 FemalesA-37 63.5 8.6 6.6 46.0 73.0 28.0 Kuranda 63.5 9.3 7.5 46.0 77.0 30.0 “ 63.0 9.5 6.8 47.6 78..3 31.2 “ 59.0 8.7 8.3 46.0 78.0 31.0 Black Mt 60.0 9.6 8.1 49.0 77.0 31.4 Mt Haig 59.2 9.3 8.2 46.7 77.0 29.0 Garradunga 56.4 9.1 7.8 43.5 71.5 31.0 RENTZ, SU, AND UESHIMA 235

Fig. 1. Segestidea queenslandica sp. nov. in natural habitat. A, Holotype male; B, female. 236 AUSTRALIAN MECOPODINAE: 2

Fig. 2. Female nymph of Segestidea queenslandica , sp. nov.

Fig. 3. Known distribution of Segestidea queenslandica sp. nov. RENTZ, SU, AND UESHIMA 237

Fig. 4. Habitat where Segestidea queenslandica sp. nov. occurs. Nymphs are found on the low-growing Lawyer Vines, Calamus spp., at night. During the day, they lie motionless resembling bird droppings. 238 AUSTRALIAN MECOPODINAE: 2

Fig. 5. Lateral view pinned specimens Segestidea queenslandica sp. nov. A, Holotype male; B, paratype female.

Fig. 6. Lateral view pronotum, female. RENTZ, SU, AND UESHIMA 239

Fig. 7. Dorsal view head and pronotum female Segestidea queenslandica sp. nov.

Fig. 8. Left tegmina (see Table 1 for measurements). A, Holotype male; B, paratype female. 240 AUSTRALIAN MECOPODINAE: 2

Fig. 9. Internal view left foreleg, Segestidea queenslandica sp. nov. Note apical spine on both sides and short apical carina on dorsal surface of tibia.

Fig. 10. Subgenital plate, female, ventral view. RENTZ, SU, AND UESHIMA 241

Fig. 11. Laid egg Segestidea queenslandica sp. nov. Note median groove present on both surfaces.

Fig. 12. Left mandible Segestidea queenslandica sp. nov.. adult female. Note robust structure and blunt tip of the incisor. This is atypical of most tettigoniids.

Fig. 13. Segestidea queenslandica sp. nov. male genitialia, holotype, dorsal view.

Fig. 14. Segestidea queenslandica sp. nov. stridulatory file, holotype male.