Glomeremus Species from the Mascarene Islands (Orthoptera, Gryllacrididae) with the Description of the Pollinator of an Endemic Orchid from the Island of Réunion

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Glomeremus species from the Mascarene islands (Orthoptera, Gryllacrididae) with the description of the pollinator of an endemic orchid from the island of Réunion

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Sylvain Hugel Ben H Warren French National Centre for Scientific Resea… Université des Antilles et de la Guyane

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Thierry Pailler Dominique Strasberg University of La Réunion University of La Réunion

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Available from: Thierry Pailler Retrieved on: 31 August 2015 TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited.

Glomeremus species from the Mascarene islands (Orthoptera, Gryllacrididae) with the description of the pollinator of an endemic orchid from the island of Réunion

S. HUGEL1, C. MICHENEAU2,3, J. FOURNEL2, B. H. WARREN2, A. GAUVIN-BIALECKI4, T. PAILLER2, M. W. CHASE5 & D. STRASBERG2 1INCI, CNRS, Université de Strasbourg, 67084 Strasbourg, France. E-mail: [email protected] 2UMR PVBMT, CIRAD-Université de La Réunion, 97415 Saint-Denis, La Réunion 3Current address: Université Libre de Bruxelles, 1050 Brussels, Belgium 4LCSNSA, Université de La Réunion, 97415 Saint-Denis, La Réunion 5Jodrell Laboratory, Royal Botanic Gardens, Kew, Richmond, Surrey TW9 3DS, UK

Abstract

Glomeremus Karny, 1937 species from the Mascarene islands are examined. One new species is described from Réunion: G. orchidophilus n. sp.. Two new species are described from Mauritius: G. paraochidophilus n. sp. and G. tikasignatus n. sp.. The case of G. shel fordi (Griffini, 1909) supposedly described from Mauritius is discussed as well as the likely polyphyly of the genus Glomeremus. G. orchidophilus n. sp. is the only pollinator of the Orchid Angraecum cadetii on Réunion, representing the first supported case of regular pollination by an insect from the order Orthoptera in extant flowering plants. The possible involvement of Mauritian Glomeremus in A. cadetii pollination on Mauritius is discussed.

Key words: Mascarene Archipelago, raspy cricket, pollinator, Mauritius, Réunion

Introduction

In the present note, Glomeremus Karny, 1937 species from Mascarene islands are examined. Up to now, only one Glomeremus species was supposedly occurring on this archipelago: Glomeremus shelfordi (Griffini, 1909). This species was described after one female labelled “Mautitius?”. This type specimen is badly damaged. In spite of numerous field studies in the Mascarene archipelago, we failed to find this species, raising doubts over the real origin of this type specimen. However, these field studies allowed the recording of three new species: G. tikasignatus n. sp. and G. paraorchidophilus n. sp. occurring in Mauritius, and G. orchidophilus n. sp., a closely related species from Réunion. The likely paraphyly of Glomeremus is briefly discussed. Elements of the biology of Glomeremus from Réunion and Mauritius are given. Surprisingly, G. orchidophilus n. sp. was recorded as the only pollinator of the orchid Angraecum cadetii (Vandeae, Epidendroideae) on Réunion (Micheneau et al., 2010), representing the first clearly-supported case of orthopteran-mediated pollination in flowering plants. As A. cadetii occurs also on Mauritius, the possible involvement of Mauritian Glomeremus in A. cadetii pollination on Mauritius is discussed.

Material and methods

Species authorship. The author of the three species described in the present article is Hugel. Specimens examined. The new taxa described in the present paper are based on specimens recently collected on the islands of Réunion and Mauritius. All specimens have been collected by sight collecting during night hours using a halogen headlamp and a net.

58 Accepted by D. Rentz: 4 Jul. 2010; published: 22 Jul. 2010 TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited.

Morphological terminology. Abbreviations. FW, fore wing; HW, hindwing; Meso, mesonotum; Meta, metanotum; O, ovipositor; Pro, pronotum; Thx, thorax. Legs: T: tibia; F: femora, followed by leg number (I, II or III). Measurements. The measurements have been performed on dry specimens or specimens stored in ethanol 70°. L: length; W: width; H: height (of pronotum lateral lobe); all in mm. Head height: from the labrum to the top of the vertex; head width: maximal width; pronotum width: maximal width (including the lateral lobes); hind femora width: maximal width; ovipositor width: near the basis; ovipositor length: linear distance from the basis (distal end of the subgenital plate) to the distal tip. Material repository. CIRAD Réunion, Centre de coopération internationale en recherche agronomique pour le développement (Réunion); MHNR, Muséum d’Histoire Naturelle de la Réunion; MNHN, Muséum national d’Histoire naturelle (France); MSIRI, Mauritius Sugar Industry Research Institute (Mauritius); NHM, Natural History Museum, London, UK; OUMNH, Oxford University Museum of Natural History, Oxford, UK. Coll. S.H., Collection Sylvain Hugel, Strasbourg, France. Scan electron microscopy. The scan electron microscope used was a JEOL JSM-6360 in low vacuum mode, with an acceleration of 18 MeV. Specimens were directly imaged, without metallization.

Results

Glomeremus Karny, 1937 Glomeremus shelfordi (Griffini, 1909) [Mauritius ?, not confirmed] Glomeremus orchidophilus n. sp. [Réunion] Glomeremus paraorchidophilus n. sp.[Mauritius] Glomeremus tikasignatus n. sp. [Mauritius]

Glomeremus shelfordi (Griffini, 1909)

Eremus sphinx var. shelfordi Griffini, 1909: 333 (original description). Glomeremus shelfordi, Karny, 1937: 115.

Material examined. Unpublished high quality pictures of female holotype (OUMNH). This type specimen is badly damaged: only a part of the head, the thorax, four abdominal tergites, the right foreleg and the left fore femora are remaining. Type locality. As specified in Griffini’s original description, the holotype is labelled “Mauritius?”. In accordance with the original description, this specimen displays 3 ventral subapical spurs on both sides of the remaining fore tibia (according to Griffini, both fore tibia and mid tibia had the same spur formula) and it is wingless. These characters clearly distinguish G. shelfordi from the three other Glomeremus species recorded in the archipelago. In addition, the ovipositors of G. shelfordi and of the other species recorded in the archipelago are clearly distinct: 19 mm for G. shelfordi (vs. <14 mm in the three species we recorded on the archipelago); almost straight for G. shelfordi (vs. clearly falciform in the three other species we recorded in the archipelago). In spite of the numerous nights spent in the field in Mauritius, no specimen corresponding to G. shelfordi was recorded during our study, suggesting either that this island is not the type locality, or that the species is very rare or extinct. Diagnosis. This species differs from all other Glomeremus species by the following character combination: face and thorax without black patterns; TI and TII with 3 ventral subapical spurs on both sides; wingless; ovipositor more than twice as long as hind femora (this latter character by itself distinguishes G. shelfordi from all other Glomeremus).

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FIGURE 1. Glomeremus orchidophilus n. sp. Male paratype. Right side view. Bar: 10 mm.

Glomeremus orchidophilus n. sp. (Figs. 2, 4, 7, 10, 13, 16, 19, tab. 1)

Holotype. Male. Indian Ocean, Mascarene archipelago, la Réunion, Saint Philippe, hauts de Mare Longue, 421 m alt., 21°20’47’’S 55°44’22’’E; 13.XII.2005, S. Hugel, MNHN (MNHN-ENSIF2642). Allotype. Female. Same as holotype, 23.IV.2007, S. Hugel, MNHN (MNHN-ENSIF2643). Paratypes. Males. 1% la Réunion, Saint Benoît, Takamaka, Alt. 800 m, 21°05’26’’S 55°37’09’’E, 20.IV.2002, S. Hugel, MHNR. 1%, same locality, 22.XI.2009, REU2009176, S. Hugel leg & coll. 1%, la Réunion, Saint-Philippe, Basse Vallée, Vallée Heureuse, rempart Est, sentier du Puys Ramond, Alt. 896 m, 21°20’14’’S 55°42’15’’E; 16.IV.2007, S. Hugel, BMNH. 1%, la Réunion, Saint-Philippe, Basse Vallée, Vallée Heureuse, rempart Est, Alt. 700–900 m, 03.IV.2005, N. Cliquennois, S. Hugel coll. 1%, la Réunion, Saint- Philippe, Basse Vallée, Vallée Heureuse, sentier de la source Repiquet, 805 m alt., 21°19’33’’S 55°41’52’’E, 11.XII.2005, S. Hugel, S. Hugel Coll. 1%, same locality, 15.XI.2009, REU2009089, S. Hugel, S. Hugel Coll. 1%, la Réunion, Saint Benoît, Grand Etang, chemin de l’Etang, Alt. 545 m, 21°05’46’’S 55°38’51’’E, 19.XI.2009, REU2009135, S. Hugel, MSIRI. 4%, la Réunion, Ste Suzanne, forêt Dugain, Alt. 900 m, 20°59’14’’S 55°34’34’’E, 10.XII.2005, S. Hugel, CIRAD (1 spn); S. Hugel leg & coll (3 spn). Females. 1&, la Réunion, Saint Philippe, hauts de Mare Longue, 500 m alt., 21°20’S 55°44’E, 29.XII.2002, S. Hugel leg & coll. 1&, la Réunion, Saint Philippe, hauts de Mare Longue, 421 m alt., 21°20’47’’S 55°44’22’’E, 23.IV.2007, S. Hugel leg & coll. 1&, la Réunion, Bras-Panon, Plaine des Lianes, forêt de la cascade du Chien, 800 m Alt., 21°02’’S 55°36’E, 09.04.2002, N. Cliquennois, MHNR. 1&, la Réunion, Saint-Joseph, Hauts de Jacques Payet, Alt. 800 m, 21°20’23’’S 55°41’41’’E; 11.XII.2005, S. Hugel leg & coll. 1&, la Réunion, Saint Benoît, Grand Etang, chemin de l’Etang, Alt. 545 m, 21°05’46’’S 55°38’51’’E, 21.XII.2005, S. Hugel, CIRAD. 1&, la Réunion, Saint Benoît, Takamaka, Alt. 800 m, 21°05’26’’S 55°37’09’’E, 22.XI.2009, REU2009177, S. Hugel leg & coll. 1&, la Réunion, Ravine de la Grande Chaloupe, petit Bon Dieu (descente sur la crête), Alt. 620 m, 20°55’48’’S 55°23’’24’E, 04.XI.2009, REU2009002, S. Hugel leg & coll. 1&, la Réunion, Saint-Philippe, Basse Vallée, Vallée Heureuse, rempart Est, Alt. 900 m, 16.V.2002, N. Cliquennois, S. Hugel Coll. Diagnosis. Glomeremus orchidophilus n. sp. is very close to G. paraorchidophilus n. sp from Mauritius. Both species differ from all other Glomeremus species by the following characters: TI and TII with 4 ventral

60 · Zootaxa 2545 © 2010 Magnolia Press HUGEL ET AL. TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited. subapical spurs on both sides, in addition to the apical spur (3 in G. sh e lf ordi ); FW reduced, but clearly distinct (all other Glomeremus species are wingless except G. tikasignatus n. sp., G. mediopictus Uvarov, 1957, and G. feanus (Griffini, 1908)); occiput with a black triangle pointing the fastigium frontis (Fig. 2; similar in G. falcifer (Sjöstedt, 1909)); head entirely black in G. feanus; some black patterns on the face in G. glomerinus glomerinus (Gerstaecker, 1860); black pattern below the scrobus in G. marginatus (Brunner v. Wattenwyl, 1888); occiput and area below the scrobus black in G. nitidus (Karsch, 1893); male SGP with a distinct process between the styli, hind margin of the process with a median emargination (similar in G. nitidus, but emargination wider and deeper separating rounded lobes; SGP posterior margin broadly rounded with a shallow median emargination in G. mediopictus Uvarov, 1957 and G. pileatus (Krauss, 1902); no emargination in G. kilimandjaricus (Sjöstedt, 1909), G. glomerinus knothae (Griffini, 1909), G. chima era (Griffini, 1911); male cerci not particularly extended (extended and thin in G. brevifalcatus brevifalcatus (Brunner v. Wattenwyl, 1888); female SGP posterior margin shallowly concave or almost strait (with a shallow median emargination in G. brevifalcatus brevifalcatus; with a distinct emargination in G. sphinx (Gerstaecker, 1860); with an obtuse “V”-shaped notch in G. sphingoides (Karny, 1929); broadly rounded, without emagination in G. marginatus; without emargination in G. c hi m a er a ). Ovipositor length < 7.5 mm, < 0.8 times FIII length (> 2 times FIII length G. shelfordi and G. sphingoides; 1.2–1.7 times FIII length in G. marginatus, G. c h im a e ra , G. obtusus (Karny, 1929); 0.9–1.1 times as long as FIII length in G. tikasignatus n. sp. and G. glomerinus); ovipositor dorsal valves with distinct ventrally directed protrusion near the basis (Fig. 16–17; absent in all other species with known females); ovipositor falciform (Fig. 16, 17; ovipositor roughly strait after the basis in G. feanus, G. glomerinus, G. sphinx, G. mediopictus, G. pileatus, G. sh e lf ordi , G. sphingoides). Glomeremus orchidophilus n. sp. and Glomeremus paraorchidophilus n. sp. are distinguished by: the size (smaller in the latter, tables 1 & 2); male tergite X sclerites forming a downward/inward-directed lamellar process in the former (Fig. 10, 13), forming an upwards/sideward-directed lamellar process in the latter (Fig. 11, 14); male tergite IX distal end with a shallow emargination separating two bulges in the former (ventral/ posterior view; Fig. 13), with a deeper emargination separating two lobes in the latter (ventral/posterior view; Fig. 14); male SGP with a shallow notch on the terminal process and with styli exceeding the process in the former (Fig. 7), with a deeper notch and with styli hardly exceeding the process in the latter (Fig. 8); female SGP with a posterior bulge in the former (Fig. 16, 19), with an anterior bulge in the latter (Fig. 17, 20).

TABLE 1. Measurements of Glomeremus orchidophilus n. sp.

Body Head Pronotum TI TII TIII FI FII FIII FIII FW O L L W L W H L L L L L L W L L W ♂ Holotype 16.5 2.7 4.7 3.9 4.1 2.3 5.3 5.4 7.9 5.0 4.8 8.5 3.0 2.0 Average 15.8 2.5 4.4 3.7 4.0 2.3 5.0 5.1 7.6 4.6 4.3 8.2 2.8 1.8 (n = 12) Min 14.9 2.3 4.2 3.5 3.8 2.0 4.6 4.7 7.1 4.1 4.1 7.8 2.7 1.6 Max 17.2 2.7 4.7 3.9 4.2 2.5 5.3 5.4 8.1 5.0 4.8 8.5 3.0 2.0 ♀ Allotype 15.7 2.9 4.7 3.9 4.0 2.2 5.5 5.6 8.5 5.1 5.0 8.7 3.1 1.7 6.4 1.3 Average 18.0 2.9 4.8 4.0 4.2 2.5 5.5 5.7 8.4 5.0 5.0 9.2 3.1 1.8 6.4 1.3 (n = 9) Min 15.7 2.6 4.5 3.9 4.0 2.2 5.3 5.4 7.9 4.6 4.7 8.7 2.9 1.6 6.0 1.2 Max 21.2 3.0 5.1 4.3 4.4 2.8 5.8 6.1 9.0 5.2 5.2 10.0 3.3 2.0 6.6 1.4

Description. In addition to generic characters. Body length: size small for the genus. Head: slightly broader than the thorax; fastigium frontis much wider than scapus; median and lateral ocelli weakly distinct. Thorax. Pronotum: maximal width (including lateral lobes) slightly wider than long; anterior margin weakly convex (in dorsal view), arched; posterior margin strait or weakly concave (in dorsal view), almost horizontal; sulci not distinct on the discus, converging on lateral lobes. Wings. FW well distinct, at most half as long as Pro. Legs. TI and TII with usual spurs formula (4 pairs of subapical + 1 pair of apical spurs

GLOMEREMUS SPECIES FROM THE MASCARENE ISLANDS Zootaxa 2545 © 2010 Magnolia Press · 61 TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited. ventrally, and one single apical spur on TII posterior dorsal angle); subapical spurs of usual length (roughly at least as long as T width). Hind legs: femora 2.8–3.3 times as long as wide (Fig. 4); FIII with 6–7 ventral anterior spines; FIII with 5–10 ventral posterior spines; TIII with 6–7 dorsal anterior spines; with 5–7 dorsal posterior spines. Abdomen. Stridulatory files present. Sexual dimorphism: apart from primary sexual characters, %% and && are similar, males are smaller. Male: terminalia (Fig. 7, 10, 13). Tergite VIII slightly extended posteriorly (compared to preceding tergites); posterior margin forming a 45° angle (in side view). Tergite IX vertically oriented (side view); distal end of tergite pointing ventrally (side view) with a shallow emargination separating two bulges (ventral/ posterior view; Fig. 13). Tergite X sclerites (see Hugel, 2009) forming a downward/inward-directed lamellar process (Fig. 10, 13). Cerci not elongated (Fig. 13). SGP distal end with a distinct process between the styli; lateral margins of the process converging; process with a shallow posterior emargination (Fig. 7); styli distinctly exceeding the SGP distal process; styli not thickened (Fig. 8). Female: sternite before SGP (VII) not modified; SGP (Figs. 16, 19); with posterior bulge (side view, Fig. 16) posterior margin shallowly concave (Fig. 19); with two distal lateral sclerifications. Ovipositor regularly arched, regularly narrowing towards the apex; pointing distally, 0.65 – 0.75 times as long as FIII, dorsal valves with a distinct ventrally directed protrusion near the basis. Colour: general coloration brown yellowish; occiput with a black pattern forming a triangle pointing on the fastigium frontis (as in G. paraorchidophilus n. sp.; Fig. 2); pronotum with a black pattern on the anterior margin, forming a triangle pointing backwards, sometimes with a black sagittal line; legs without black pattern; knees sometimes darkened; anterior part of abdominal tergites darkened.

Glomeremus paraorchidophilus n. sp. (Figs. 5, 8, 11, 14, 17, 20, tab. 2)

Holotype. Male. Indian Ocean, Mascarene archipelago, Maurice, Savanne District, Parc National de Rivière Noire, Cocotte, 745 m Alt., 20°26’29’’S 57°28’20’’E, 22.IV.2005, S. Hugel, MNHN (MNHN-ENSIF2644). Allotype. Female. Same as holotype, 22.IV.2005, S. Hugel, MNHN (MNHN-ENSIF2645). Paratypes. Males. 2%, same as holotype, 22.II.2008, BIOTAS2008212, S. Hugel, MHNR; 22.IV.2005, S. Hugel leg & coll. 1%, Maurice, Savanne District, Parc National de Rivière Noire, Plaine Champagne, point de vue sur les Gorges, 681 m alt., 20°25’39’’S 57°25’43’’E, 22.IV.2005. S. Hugel, MSIRI. 1%, Maurice, Plaines Wilhems District, Petrin, 604 m alt., 20°24’27’’S 57°28’21’’E, 23.II.2008, BIOTAS2008122, S. Hugel leg & coll. 1%, Maurice, Port Louis District, le Pouce (dernier plateau), 733 m alt., 20°12’13’’S 57°31’43’’E, 08.V.2009, BIOTAS2009217, S. Hugel leg & coll. 2%, Maurice, Plaines Wilhems District, Brise Fer vieille parcelle, 648 m alt., 20°23’02’’S 57°26’23’’E, 24.II.2008, BIOTAS2008176, S. Hugel, CIRAD Réunion; BIOTAS2008177, S. Hugel, CIRAD Réunion. 1%, Maurice, Black River District, Parc National de Rivière Noire, Piton Brise Fer, 617 m alt., 20°23’02’’S 57°25’51’’E, 13.V.2009, BIOTAS2009349, S. Hugel leg & coll. Females. 1&, same as holotype, 07.V.2009, BIOTAS2009164, S. Hugel leg & coll. 1&, Maurice, Savanne District, Parc National de Rivière Noire, Cocotte, 745 m Alt., 20°26’29’’S 57°28’20’’E, 22.IV.2005, S. Hugel leg & coll. 3&, Maurice, Plaines Wilhems District, Florin, 610 m alt., 20°23’28’’S 57°27’37’’E, 25.II.2008, BIOTAS2008191, S. Hugel leg & coll; BIOTAS2008190, S. Hugel leg & coll; BIOTAS2008192, S. Hugel, MSIRI. 4&, Maurice, Plaines Wilhems District, Petrin, 604 m alt., 20°24’27’’S 57°28’21’’E, 23.II.2008, BIOTAS2008130, S. Hugel, BMNH; BIOTAS2008131, S. Hugel, BMNH; BIOTAS2008123, S. Hugel leg & coll. BIOTAS2008124, S. Hugel leg & coll. 1&, Maurice, Port Louis District, le Pouce (dernier plateau), 733 m alt., 20°12’13’’S 57°31’43’’E, 08.V.2009, BIOTAS2009206, S. Hugel leg & coll. 2&, Maurice, Plaines Wilhems District, Brise Fer vieille parcelle, 648 m alt., 20°23’02’’S 57°26’23’’E, 04.V.2009, BIOTAS2009188, S. Hugel leg & coll; 24.II.2008, BIOTAS2008178, S. Hugel, CIRAD Réunion. Diagnosis. See the diagnosis of G. orchidophilus n. sp.. Description. In addition to generic characters. Body length: size small for the genus. Head: slightly broader than the thorax; fastigium frontis wider than scapus; median and lateral ocelli weakly distinct.

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FIGURES 2–21. Glomeremus of the Mascarene islands. 2, 4, 7, 10, 13, 16, 19, Glomeremus orchidophilus n. sp. 5, 8, 11, 14, 17, 20, Glomeremus paraorchidophilus n. sp. 3, 6, 9, 12, 15, 18, 21, Glomeremus tikasignatus n. sp. 2, 3, head in frontal view. 4–6, left hind femora in side view. 7–9, male subgenital plate in ventral view. 10–12, scan electron microscope picture of male tergite X right sclerite. 13–15, male terminalia (SGP removed) in ventral/posterior view. 16– 18, female terminalia in right side view. 19–21, female subgenital plate and basis of ovipositor in ventral view. Bars: 1 mm.

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Thorax. Pronotum: maximal width (including lateral lobes) slightly wider than long; anterior margin weakly convex (in dorsal view), arched; posterior margin strait (in dorsal view), almost horizontal; sulci not distinct on the discus, converging on lateral lobes. Wings. FW well distinct, half as long as Pro. Legs. TI and TII with usual spurs formula (4 pairs of subapical + 1 pair of apical spurs ventrally, and one single apical spur on TII posterior dorsal angle); subapical spurs of usual length (roughly at least as long as T width). Hind legs: femora 2.5–3.0 times as long as wide (Fig. 5); FIII with 2–8 (usually 6) ventral anterior spines; FIII with 3–7 ventral posterior spines; TIII with 6–8 dorsal anterior spines; with 6–7 dorsal posterior spines. Abdomen. Stridulatory files present.

TABLE 2. Measurements of Glomeremus paraorchidophilus n. sp.

Body Head Pronotum TI TII TIII FI FII FIII FIII FW O L L W L W H L L L L L L W L L W ♂ Holotype 14.3 2.2 4.1 3.3 3.6 1.8 4.7 4.6 6.7 4.0 4.0 7.0 2.8 1.9 Average 14.8 2.1 4.2 3.2 3.8 1.9 4.5 4.5 6.7 4.1 4.1 7.2 2.6 1.8 (n = 9) Min 13.4 1.8 3.8 3.0 3.4 1.5 4.1 4.2 6.0 3.8 3.7 6.6 2.4 1.6 Max 16.5 2.2 4.6 3.5 4.2 2.1 4.7 4.8 7.0 4.5 4.4 8.1 2.8 1.9 ♀ Allotype 18.6 2.5 4.7 3.8 4.2 2.5 5.6 5.8 8.3 5.1 5.2 9.1 3.0 2.3 7.1 1.3 Average 17.0 2.5 4.6 3.7 4.2 2.3 5.4 5.4 7.9 4.8 4.7 8.6 3.0 2.1 6.9 1.2 (n = 13) Min 13.5 2.3 4.4 3.5 3.9 2.1 4.9 5.0 7.5 4.2 4.2 8.0 2.9 1.8 6.6 1.1 Max 19.6 2.8 4.8 3.9 4.6 2.7 5.7 5.8 8.3 5.2 5.2 9.2 3.2 2.4 7.1 1.3

Sexual dimorphism: apart from primary sexual characters, %% and && are similar, males are smaller. Male: terminalia (Fig. 8, 11, 14). Tergite VIII slightly extended posteriorly (compared to preceding tergites); posterior margin forming a 45° angle (in side view). Tergite IX vertically oriented (side view); distal end of tergite pointing ventrally (side view) with a distinct emargination separating two lobes (ventral/ posterior view; Fig. 14). Tergite X sclerites (see Hugel, 2009) forming an upwards/sideward-directed lamellar process (Fig. 11, 14). Cerci shortened (Fig. 14). SGP distal end with a distinct process between the styli; lateral margins of the process weakly converging; process with a distinct posterior notch (Fig. 8); styli hardly exceeding the SGP distal process; styli not thickened (Fig. 8). Female: sternite before SGP (VII) not modified; SGP (Figs. 17, 20) with rounded lateral margins; with anterior bulge (side view, Fig. 17) posterior margin with a shallow emargination (Fig. 20); with two distal lateral sclerifications. Ovipositor regularly arched, regularly narrowing towards the apex; pointing distally, 0.7 – 0.8 times as long as FIII, dorsal valves with a distinct ventrally directed protrusion near the basis. Colour: general coloration brown yellowish; occiput with a black pattern forming a triangle pointing on the fastigium frontis (as in G. orchidophilus n. sp.; Fig. 2); pronotum with a black pattern on the anterior margin, forming a triangle pointing backwards, sometimes with a black sagittal line and black on the posterior margin; legs without black pattern; knees sometimes darkened; anterior part of abdominal tergites darkened.

Glomeremus tikasignatus n. sp. (Figs. 3, 6, 9, 12, 15, 18, 21, tab. 3)

Holotype. Male. Indian Ocean, Mascarene archipelago, Maurice, Savanne District, Parc National de Rivière Noire, Plaine Champagne, point de vue sur les Gorges, 681 m alt., 20°25’39’’S 57°25’43’’E, 22.IV.2005. S. Hugel, MNHN (MNHN-ENSIF2646). Allotype. Female. Same as holotype, 13.VIII.2001. S. Hugel, MNHN (MNHN-ENSIF2647). Paratypes. Males. 2%, Maurice, Savanne District, Parc National de Rivière Noire, Cocotte, 745 m Alt., 20°26’29’’S 57°28’20’’E, 22.II.2008, BIOTAS2008210, S. Hugel leg & coll.; 07.V.2009, BIOTAS2009163,

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S. Hugel leg & coll. Females. 1&, Maurice, Black River District, Parc National de Rivière Noire, Piton Brise Fer, 617 m alt., 20°23’02’’S 57°25’51’’E, 13.V.2009, BIOTAS2009286, S. Hugel, MSIRI. 5&, Maurice, Savanne District, Parc National de Rivière Noire, Cocotte, 745 m Alt., 20°26’29’’S 57°28’20’’E, 22.II.2008, BIOTAS2008211, S. Hugel leg & coll.; 06.III.2008, BIOTAS2008412, S. Hugel leg & coll.; 22.IV.2005, S. Hugel leg & coll.; 22.IV.2005, S. Hugel, CIRAD Réunion; 19.IV.2005, S. Hugel leg & coll. Diagnosis. This species differs from the other Glomeremus in the following characters: TI and TII with 4 ventral subapical spurs on both sides, in addition to the apical spur (3 in G. s he l fo rd i); FW reduced, but clearly distinct (all other Glomeremus species are wingless except G. orchidophilus n. sp., G. paraorchidophilus n. sp., G. mediopictus Uvarov, 1957, and G. feanus (Griffini, 1908)); the thorax and head without black pattern; male cerci not particularly extended (extended and thin in G. brevifalcatus); male tergite X sclerites forming an upwards directed lamellar process (Fig. 12, 15); male SGP with a shallow emargination (Fig. 9); female SGP with a “W”-shaped end margin (Fig. 21); ovipositor falciform, pointing apically, as long as FIII, dorsal valves without ventrally directed protrusion (Fig. 20; ovipositor roughly straight after the base in G. feanus, G. glomerinus, G. s p hi n x, G. mediopictus, G. pileatus, G. shelfordi, G. sphingoides). Description. In addition to generic characters. Body length: size medium to large for the genus. Head (Fig. 3): as broad or slightly broader than the thorax; fastigium frontis wider than scapus; median and lateral ocelli clearly distinct (less obvious on specimens dried from alcohol), median ocellus circular, less wide than scapus. Thorax. Pronotum: maximal width (including lateral lobes) as wide as or slightly wider than long; anterior margin convex (in dorsal view), arched; posterior margin strait (in dorsal view), almost horizontal; anterior sulcus deep on lateral lobes and well distinct on discus; posterior sulcus shallower and less distinct on discus; both sulci converging on lateral lobes; prozona elevated. Wings. FW well distinct, half as long as Pro. Legs. TI and TII with usual spurs formula (4 pairs of subapical + 1 pair of apical spurs ventrally, and one single apical spur on TII posterior dorsal angle); subapical spurs of usual length (at least as long as T width). Hind legs: femora 3.4–3.7 times as long as wide (Fig. 6); FIII with 5–7 ventral anterior spines; FIII with 6–11 ventral posterior spines; TIII with 5–7 dorsal anterior spines; with 6–8 dorsal posterior spines. Abdomen. Stridulatory files present.

TABLE 3. Measurements of Glomeremus tikasignatus n. sp.

Body Head Pronotum TI TII TIII FI FII FIII FIII FW O L L W L W H L L L L L L W L L W ♂ Holotype 24.4 3.3 5.5 4.8 4.7 2.5 6.8 6.8 10.4 6.3 6.3 11.0 3.1 2.0 Average 26.5 3.4 6.0 4.9 5.4 2.5 7.4 7.4 11.2 6.8 6.5 11.9 3.3 2.2 (n = 3) Min 24.4 3.2 5.5 4.8 4.7 2.4 6.8 6.8 10.4 6.3 6.3 11.0 3.1 2.0 Max 27.5 3.8 6.6 5.2 5.7 2.7 8.2 8.1 12.2 7.2 6.6 13.2 3.6 2.4 ♀ Allotype 23.5 2.7 6.4 5.4 5.8 2.3 7.7 8.1 11.8 7.2 7.2 12.0 3.5 3.4 13.9 1.7 Average 25.3 3.3 6.4 5.4 6.0 2.8 8.1 8.1 12.4 7.4 7.3 12.8 3.6 3.4 13.1 1.6 (n = 7) Min 21.8 2.7 6.1 5.1 5.8 2.3 7.7 7.9 11.5 7.1 6.9 12.0 3.4 3.1 12.3 1.5 Max 29.8 3.6 7.0 5.8 6.5 3.6 8.7 8.6 13.4 7.6 7.7 13.6 3.8 4.0 13.9 1.7

Sexual dimorphism: apart from primary sexual characters, %% and && are similar. Male: terminalia (Fig. 9, 12, 15). Tergite VIII slightly extended posteriorly (compared to preceding tergites); posterior margin forming a 45° angle (in side view). Tergite IX vertically oriented (side view); distal end of tergite pointing ventrally (side view) with a shallow emargination separating two small bulges (ventral/ posterior view; Fig. 15). TX sclerites (see Hugel, 2009) forming an upwards directed lamellar process (Fig. 12, 15). Cerci not particularly elongated (Fig. 15). SGP distal end with an indistinct emargination (Fig. 9); styli exceeding distinctly the SGP distal end, not thickened (Fig. 9). Female: sternite before SGP (VII) not

GLOMEREMUS SPECIES FROM THE MASCARENE ISLANDS Zootaxa 2545 © 2010 Magnolia Press · 65 TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited. modified; SGP (Figs. 18, 21) with strait lateral margins, posterior margin pointing ventrally (side view, Fig. 18), the posterior end corresponds to the SGP dorsal surface, with a shallow “W” shape (ventral view, Fig. 21). Ovipositor regularly arched, pointing distally, as long as FIII, dorsal valves without ventrally directed protrusion. Colour: general coloration brown; head, legs pronotum without black pattern (knees sometimes darkened); anterior part of abdominal tergites darkened; median and lateral ocelli white.

Discussion

Definition of Glomeremus. Karny (1937) defined many Gryllacrididae genera after wing venation characters. For African wingless or micropterous species lacking other characters, he erected the genus Glomeremus. As wing condition (macro-, brachy-, micro- and apterism) can differ in closely related orthopteroid species (e.g. Whiting et al. 2003) and within Gryllacrididae genera (e.g Niphetogryllacris brevipennis Chopard, 1958 vs. other Niphetogryllacris species having long wings) and as the genus is defined by lack of characters rather than by clear synapomorphic characters, this genus is likely to be polyphyletic. Although male genitalia of the three species described above are very similar, female characters of G. tikasignatus n. sp. on the one hand and G. orchidophilus n. sp. and G. paraorchidophilus n. sp. on the other hand are clearly differing. Females of the two latter species are clearly distinguished by the dorsal valves with a distinct ventrally directed protrusion near the basis. To our knowledge, this character is unique to these two species among the Gryllacrididae. Pollination of the orchid Angraecum cadetii Bosser. Observations conducted on the island of Reunion (using hard-disk camcorders) revealed that Glomeremus orchidophilus n. sp. regularly visited flowers of Angraecum cadetii during the flowering period of the orchid, typically from mid-end of January to March (austral summer). At the peak of the orchid flowering period, raspy crickets visited the same patch of plants with high fidelity night after night (from 7:50 pm to 4:40 pm), adopting the same behaviour each time, i.e. probing deeply into the orchid spur, the head totally hidden in the centre of the flower, and visiting the majority of “fresh-looking” flowers within reach. Importantly, we never observed Glomeremus orchidophilus n. sp. eating and/or damaging A. cadetii flowers. Pollen transfer (i.e. orchid pollinarium removal and deposition) was clearly observed, indicating that this species of Glomeremus is an effective pollinator of A. cadetii on Reunion. The average duration of a single flower visit was 16.5 ± 8.7 seconds (min = 3.1; max = 41.0; n = 75 flowers). Typically, raspy crickets reached flowers by climbing up leaves of the orchid or jumping across from neighbouring plants. In all successful pollination events, the raspy cricket positioned itself on the fleshy lip of the flower with its dorsal side orientated towards the orchid column and probed deep within the spur. Pollinaria of the orchid became stuck to the head of the crickets as they retreated from flowers. Orthopterans are well known for herbivory, and this insect order is not normally considered to be capable of regular pollination (e.g. Darwin, 1862; Knuth, 1909; van der Pijl and Dodson, 1966; Proctor et al., 1996; van der Cingel, 2001). Nevertheless, specialised orthopteran clades are feeding on nectar and pollen (Zaprochilinae) or even full flowers (Phasmodinae) (Rentz, 1993). Although a few studies have recorded orthopterans as flower visitors, occasionally carrying pollen after having consumed it, no evidence of regular pollination has ever been clearly documented. Casual cases of pollination by orthopterans have always been linked with herbivory and pollinivory and flower visits by orthopterans might even be deleterious (Schuster, 1974). Our observations conducted in Reunion (Pandanus forests) revealed that A. cadetii seems to rely on the unique services of Glomeremus orchidophilus n. sp. to achieve fruit set. Glomeremus orchidophilus n. sp. regularly visit orchid flowers, presumably for the nectar they contain (food reward), and the role of these insects in the pollination of A. cadetii does not appear to be occasional or incidental. Along with lack of pollination by all other recorded flower visitors (either by day or at night), and high level of fruiting success observed in natural populations (ranging from 11.9 % to 43.4 % depending of the sites sampled across the island), it appears that A. cadetti (unable to auto-pollinated without pollen vectors) is highly dependent on its orthopteran pollinator on Reunion for successful reproduction and species long-term survival (see Micheneau et al., 2010 for more details).

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The diet of Gryllacridinae ranges from plant material (seeds, fruits, frowers) to other arthropods depending on the species (Hale and Rentz 2001), some species being specialized (Rentz, 1996). In the stomach of Glomeremus orchidophilus n. sp., (5 specimens dissected) we found mostly plant material including pollen and seeds, and few insect body parts, indicating an omnivorous diet. Interestingly, gryllacridinae are central place foragers searching food at night and returning with a strong fidelity to the same nest each morning (Hale, 2000; Hale and Rentz, 2001). This nest fidelity depends on gryllacridid mnesic capacities, the use of spatial landmarks and the capability of measuring translational displacements (Hale and Bailey 2004). Together with possible attraction by chemical signals emitted during night hours by mature flowers, gryllacridinae abilities to relocalize their nest might be used to repeatedly visit food sources. Such repeated visits are suggested by the high pollination rate observed in A. cadetti spikes, where all flowers are not mature at the same time. Theses mnesic and navigation abilities might represent ‘preadaptations’ to orchid pollination that may have favored repeated nectar-feeding from Angraecum flowers, which form a highly localized food resource. We never recorded florivory of A. cadetii by G. orchidophilus n. sp., but other Mascarene Gryllacridids are frequently observed eating flowers (e.g. Niphetogryllacris sp.), suggesting that florivory might have been the first step of A. cadetii and G. orchidophilus n. sp. ancestors interaction. Interestingly, whereas Angraecum cadetii occurs both on Reunion and Mauritius, its only known pollinator is restricted to Reunion. However G. paraorchidophilus n. sp., a very close relative of G. orchidophilus n. sp., occurs on Mauritius. Both species display very similar ecological requirements and habits, but G. paraorchidophilus n. sp. is slightly smaller in size. Whether G. paraorchidophilus n. sp. ensures successful pollination of A. cadetii in Mauritius remains an open question. One specimen has been observed walking near an open flower, but without stopping, and without probing into the orchid spur.

Acknowledgements

We thank James Hogan (OUMNH) for the high quality pictures of G. shelfordi holotype. We thank the National Parks and the forestry services of both Mauritius/Rodrigues (National Parks and Conservation Service) and la Réunion (Parc National de la Réunion, Office National des Forêts) for giving access to the protected areas, and the Mauritian Wildlife Fundation (MWF, Mauritius & Rodrigues) for logistic support on the field. We are indebted to all the colleagues and friends from Mauritius, Rodrigues and la Réunion, who either helped us directly in the field and/or gave precious advices and support for the fieldwork: M. Allet (NPCS), C. Baider (Mauritius Herbarium MSIRI), A. Brondeau (ONF), N. Cliquenois (Majunga, Madagascar), N. Cole (MWF), G. d’Argent (Mondrain Nature Réserve), B. Devaux (ONF), V. Florens (Univ. Mauritius), A. Franck (CIRAD Réunion), S. Ganeshan (MSIRI), O. Griffiths (Mauritius), G. Huot-Marchand (ONF), the Lambelin/Walther family (Ste Suzanne), B. Lequette (PNR), P. Moollee (NPCS), A. Releaux (MWF), O. Robinet (PNR), J. Rochat and his team (Insectarium de la Réunion), K. Ruhomaun (NPCS), V. Tataya (MWF), J. Thomas (ONF), J. Triolo (ONF), N. Zuel (MWF/Univ. Zurich), all the volunteers and wards of the MWF, and S. H.'s father who joined most the fieldtrips. We acknowledge J.-H. Lignot for the use of the scan electron microscope of his laboratory (DEPE, UMR 7168, Strasbourg). The field trips in la Réunion in 2001 and 2005 have been supported by the Germaine Cousin grant (Société Entomologique de France), and by the CIRAD Réunion, respectively. The collecting trips in Mauritius and off shore islands (2008–2009) have been funded by the BIOTAS project, ANR-06-BDIV-002.

References

Darwin, C. (1862) On the various contrivances by which British and foreign orchids are fertilised by insects, and on the good effect of intercrossing; John Murray, London, 365 pp. Griffini, A. (1909) Studi sui Grillacridi del Museo di Oxford. Parte Ia, specie etiopiche, indo-malesi ed australiane. Atti della Societa Italiana di Scienze Naturali, 47, pp. 300–338.

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Hale, R.J. (2000) Nest utilisation and recognition by juvenile gryllacridids (Orthoptera: Gryllacrididae). Australian Journal of Zoology, 48, 643–652. Hale, R.J. & Bailey, W.J. (2004) Homing behaviour of juvenile Australian raspy crickets (Orthoptera: Gryllacrididae). Physiological Entomology 29, 426–435. Hale, R.J. & Rentz, D.C.F. (2001) The Gryllacrididae: An overview of the world fauna with emphasis on Australian examples. in: Field, L.H. (Ed). The Biology of Wetas, King Crickets and their Allies. Cabi Publishing, Wallingford. pp. 95–110. Hugel, S. (2009) Gryllacrididae and Tettigoniidae (Insecta, orthoptera, ensifera) from espiritu Santo, Vanatu. Zoosystema, 31 (3), pp. 525–576. Karny, H.H. (1937) Orthoptera Fam. Gryllacrididae Subfamiliae omnes. In: Wytsman, P (Ed). Genera Insectorum, V. Verteneuil & L. Desmet, Bruxelles. 317 pp. Knuth, P. (1909) Handbook of flower pollination, Volume III. Clarendon Press, Oxford. Micheneau, C., Fournel, J., Warren, B.H., Hugel, S., Gauvin-Bialecki, A., Pailler, T., Strasberg, D. & Chase, M.W. (2010) Orthoptera, a new order of pollinator. Annals of Botany, 105, pp. 355–364. Proctor, M., Yeo, P. & Lack, A. (1996) The Natural History of Pollination. Timber Press, Portland, 479 pp. Rentz, D.C.F. (1996) Grasshopper country. The abundant orthopteroid insects of Australia. Orthoptera; grasshoppers, katydids, crickets. Blattodea; cockroaches. Mantodea; mantids. Phasmatodea; stick insects. University of New South Wales, Sydney, pp284. Rentz, D.C.F. (1993) Tettigoniidae of Australia, Vol. 2, The Austrosaginae, Zaprochilinae and Phasmodinae. CSIRO, East Melbourne, 386pp. Schuster, J.C. (1974) Saltatorial Orthoptera as Common Visitors to Tropical Flowers. Biotropica, 6 (2), 138–140. van der Cingel, N.A. (2001) An atlas of orchid pollination: America, Africa, Asia and Australia. A.A. Balkema Publishers, Rotterdam, 260 pp. van der Pijl, L. & Dodson, C.H. (1966) Orchid flowers, their pollination and evolution. Coral Gables, University of Miami Press, 214 pp.