Review of the Madagascan Orphninae (Coleoptera: Scarabaeidae) with a Revision of the Genus Triodontus Westwood

Zootaxa 4207 (1): 001–093 ISSN 1175-5326 (print edition) http://www.mapress.com/j/zt/ Monograph ZOOTAXA Copyright © 2016 Magnolia Press ISSN 1175-5334 (online edition) http://doi.org/10.11646/zootaxa.4207.1.1 http://zoobank.org/urn:lsid:zoobank.org:pub:F2875582-31E2-496F-AEEF-1D657DD86C33 ZOOTAXA

4207

Review of the Madagascan Orphninae (Coleoptera: Scarabaeidae) with a revision of the genus Triodontus Westwood

ANDREY V. FROLOV1,2,3,4, OLIVIER MONTREUIL2 & LILIA A. AKHMETOVA1,3 1Laboratory of Insect Systematics, Zoological Institute, Russian Academy of Sciences, Universitetskaya nab., 1, 199034 Sankt-Petersburg, Russia. E-mail: [email protected] 2MECADEV, UMR 7179 MNHN/CNRS, Muséum national d'Histoire naturelle, Entomologie, CP 50, 45 rue Buffon, 75231 Paris cedex 05, France. 3Universidade Federal de Mato Grosso, Instituto de Biociencias, Departamento de Biologia e Zoologia, Av. Fernando Correa da Costa, 2367, 78060–900 - Cuiaba, MT, Brazil 4Corresponding author

Magnolia Press Auckland, New Zealand

Accepted by A.B.T. Smith: 8 Nov. 2016; published: 13 Dec. 2016 ANDREY V. FROLOV, OLIVIER MONTREUIL & LILIA A. AKHMETOVA Review of the Madagascan Orphninae (Coleoptera: Scarabaeidae) with a revision of the genus Triodontus Westwood ( Zootaxa 4207) 93 pp.; 30 cm. 13 Dec. 2016 ISBN 978-1-77670-048-6 (paperback) ISBN 978-1-77670-049-3 (Online edition)

FIRST PUBLISHED IN 2016 BY Magnolia Press P.O. Box 41-383 Auckland 1346 New Zealand e-mail: [email protected] http://www.mapress.com/j/zt

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ISSN 1175-5326 (Print edition) ISSN 1175-5334 (Online edition)

Abstract ...... 4 Introduction ...... 4 Material and methods ...... 5 External morphology of adult Orphninae ...... 6 Natural history of the Madagascan Orphninae ...... 12 Key to the Madagascar Orphninae genera (both sexes) ...... 12 Genus Pseudorphnus Benderitter, 1913 ...... 12 Key to the Pseudorphnus species (males) ...... 13 Pseudorphnus carinatus Frolov, 2011...... 13 Pseudorphnus olsoufieffi Paulian, 1977 ...... 15 Pseudorphnus hiboni Paulian, 1959 ...... 17 Pseudorphnus coquerelii (Fairmaire, 1868) ...... 19 Genus Madecorphnus Paulian, 1992 ...... 22 Key to the Madecorphnus species (males) ...... 23 Madecorphnus falculoides (Paulian, 1977)...... 24 Madecorphnus montreuili Frolov, 2010 ...... 26 Madecorphnus barclayi Frolov, 2012 ...... 26 Madecorphnus tuberculatus Frolov, 2014 ...... 29 Madecorphnus fisheri Frolov, 2014 ...... 30 Madecorphnus pauliani Frolov, 2010 ...... 32 Madecorphnus niger Frolov, 2010 ...... 32 Madecorphnus punctatus Frolov, 2010 ...... 35 Madecorphnus dentatus Frolov, 2010 ...... 35 Madecorphnus saintemariensis Frolov, 2014 ...... 38 Madecorphnus simplex Frolov, 2010 ...... 40 Madecorphnus aquilonius Frolov, 2012 ...... 42 Madecorphnus falcatus Paulian, 1992 ...... 43 Madecorphnus falciger (Lansberge, 1886) ...... 45 Madecorphnus hanskii Frolov, 2012 ...... 46 Madecorphnus cuccodoroi Frolov, 2010...... 48 Madecorphnus brunneus Frolov, 2010 ...... 50 Madecorphnus perinetensis Frolov, 2010 ...... 50 Madecorphnus peyrierasi Frolov, 2010 ...... 54 Genus Renorphnus Frolov & Montreuil, 2009 ...... 54 Renorphnus clementi (Petrovitz, 1971) ...... 55 Genus Triodontus Westwood, 1845...... 57 Key to Triodontus species (males) ...... 58 Triodontus hova (Fairmaire, 1868) ...... 58 Triodontus nitidulus (Guérin-Méneville, 1844) ...... 61 Triodontus bicavatus (Fairmaire, 1905) ...... 66 Triodontus ankarafantsikae Frolov, Montreuil & Akhmetova, new species ...... 68 Triodontus lemoulti Frolov, Montreuil & Akhmetova, new species ...... 70 Triodontus modestus (Benderitter, 1914) ...... 72 Triodontus owas Westwood, 1852...... 74 Triodontus viettei Frolov, Montreuil & Akhmetova, new species ...... 76 Triodontus hildebrandtii (Fairmaire, 1883) ...... 78 Triodontus alticola Paulian, 1977 ...... 80 Triodontus fairmairei Frolov, Montreuil & Akhmetova, new species ...... 82 Triodontus itremoi Paulian, 1977 ...... 83 Triodontus copridoides Paulian, 1977 ...... 86 Triodontus hanskii Frolov, 2010 ...... 88 Triodontus inexpectatus Frolov, Montreuil & Akhmetova, new species ...... 88 Acknowledgments ...... 91 References cited ...... 91

The subfamily Orphninae (Coleoptera: Scarabaeidae) is reviewed from Madagascar. A total of four genera and 39 species were found, all being endemic to the island. The following five new species are described: Triodontus ankarafantsikae, Triodontus lemoulti, Triodontus viettei, Triodontus fairmairei, and Triodontus inexpectatus. The following new synonymies are proposed: Orphnus nigrita Brancsik, 1893 is synonym of Triodontus hova (Fairmaire, 1868); Triodontus occidentalis Paulian, 1977 and Orphnus obsoletus Brancsik, 1893 are synonyms of Triodontus nitidulus (Guérin-Méneville, 1844); Triodontus vadoni Paulian, 1977 and Triodontus perrotorum Paulian, 1977 are synonyms of Triodontus owas West- wood, 1852. Lectotypes are designated for the following names: Orphnus nitidulus Guérin-Méneville, 1844 and Orphnid- ius modestus Benderitter, 1914. Keys, descriptions, illustrations of habitus and male genitalia, and distributional records maps are given for all species.

Key words: scarab beetles, Pseudorphnus, Madecorphnus, Renorphnus, Madagascar

Introduction

The subfamily Orphninae (Coleoptera: Scarabaeidae) comprises small to medium-sized scarab beetles with the appearance somewhat intermediate between the “dung beetles” and “chafers”. Orphnines are uniformly colored, brown to black. The majority of the genera exhibit distinct sexual dimorphism with the larger males having variably developed frontoclypeal and prothoracic processes, sometimes species specific, but these characters are subject to allometric variability and some males are superficially similar to females. There are about 200 described species of the Orphninae, which are grouped into 17 genera (the up-to-date catalogue is available at http://www.scarabaeoidea.com/taxa/orphninae/catalogue). Orphnines are widely distributed in the tropical and subtropical regions of the southern continents except for Australia, and are the most diverse in the Afrotropics. The Afrotropical fauna is the richest and comprises over 100 species most of which are placed in the genus Orphnus MacLeay, 1819. The three other large regional Orphninae faunas (Frolov 2012a)— Madagascan, Neotropical, and Mediterranean—all comprise about 40 species yet the Madagascan fauna is limited to a much smaller area than the other two. Madagascar has long been recognized as a biodiversity hotspot and is one of the global biodiversity conservation priorities within the framework of both ‘‘irreplaceability’’ and ‘‘vulnerability’’(Brooks et al. 2006). Endemism of the Madagascan biota is very high in all groups of animals and plants, yet the biodiversity of the island is largely underestimated (Vieites et al. 2009). Almost every taxonomic work on the scarab beetles of the island reveals previously unknown taxa (e.g. Dechambre 1986; Lacroix 1988, 1989, 1993, 1997; Huchet 2003; Montreuil 2005, 2008, 2010, 2013; Montreuil & Viljanen 2007; Akhmetova & Montreuil 2010; Montreuil & Frolov 2014; Montreuil et al. 2014). The earliest records of the Orphninae from Madagascar are dated from the mid-19th century and until the mid- 20th century the literature on the group was mostly limited to the isolated descriptions of the species. Guérin- Méneville (1844) described the first Madagascan orphnine species, Orphnus nitidulus Guérin-Méneville, 1844, in the Afrotropical genus Orphnus. The next year Westwood (1845) erected Triodontus Westwood, 1845 for O. nitidulus and later described T. owas Westwood, 1852, from Madagascar without exact type locality. Fairmaire (1868) described O. hova Fairmaire, 1868 and O. coquerelii Fairmaire, 1868. The same year Lansberge (1868) described Drepanognathus falciger Lansberge, 1868. Brancsik (1893) described O. obsoletus Brancsik, 1893 and O. nigritus Brancsik, 1893 from Nosy Be Island. Fairmaire described O. hildebrandtii Fairmaire 1883 from Madagascar without more precise locality and O. bicavatus Fairmaire 1905 from northern Madagascar (Fairmaire 1883, 1905). Benderitter (1913) erected two genera, Orphnidius Benderitter, 1913 and Pseudorphnus Benderitter, 1913, for O. nitidulus and O. coquerelii, respectively, and later (Benderitter 1914a) described O. modestus from central Madagascar. Petrovitz (1971) described O. clementi Petrovitz, 1971 from eastern Madagascar. Renaud Paulian started revisionary taxonomic work on the Madagacan Orphninae and made the most significant contribution to the classification of the group. In one of his works (Paulian 1977), based mostly on the collection of the National Museum of Natural History in Paris, France, he treated 20 species including nine new. Paulian noted the high diversity of the Madagascan Orphninae and that all the species are endemic to the island although he treated two species, Sissantobius falciger and Orphnus clementi, as the members of the corresponding

4 · Zootaxa 4207 (1) © 2016 Magnolia Press FROLOV ET AL. Afrotropical genera. Later Paulian (1992) erected the genus Madecorphnus Paulian, 1992 to separate Madagascar species of Sissantobius from African S. mandibularis (Lansberge 1886). Previous authors did not use characters of the internal sac of the aedeagus, which allowed us to distinguish a number of cryptic species in the genus Madecorphnus (Frolov 2010a, 2010c, 2012b, 2014). Previous authors also underestimated intraspecific allometric variation in Triodontus, partly due to the rather scarce material available to them. A large material of the Madagascan Orphninae has been accumulated in the past two decades. The two major sources of it were the surveys of the Metapopulation Research Group led by Ilkka Hanski (University of Helsinki, Finland) and researchers from California Academy of Sciences (San Francisco, California, United States of America) led by Brian Fisher. Much of this material was untreated until now. In the present work we summarize all the data about the Madagascan Orphninae accumulated to date and revise the genus Triodontus. For all taxa we provide illustrations, collecting locality maps, and identification keys. General morphology of the adult Orphninae is described with the emphasis on the characters of diagnostic and taxonomical importance.

Material and methods

The material used in this work is housed in the following organizations and private collections (curators in brackets):

ABCB Alberto Ballerio private collection, Brescia, Italy (Alberto Ballerio) BMNH Natural History Museum, London, United Kingdom (Maxwell Barclay) CASC California Academy of Sciences, San Francisco, California, United States of America (Jere Schweikert, Norman Penny) FMNH Field Museum of Natural History, Chicago, Illinois, United States of America (Alfred Newton, Margaret Thayer) HNHM Natural History Museum, Budapest, Hungary (Otto Merkl) IRSNB Belgian Royal Institute of Natural Sciences, Brussels, Belgium (Alain Drumont) MHNG Natural History Museum, Geneva, Switzerland (Giulio Cuccodoro) MNHN National Museum of Natural History, Paris, France (Olivier Montreuil) MRAC Royal Museum for Central Africa, Tervuren, Belgium (Marc De Meyer) NHMB Natural History Museum, Basel, Switzerland (Eva Sprecher) NMPC National Museum of Natural History, Prague, Czech Republic (Jiří Hájek) NRS State Museum of Natural History, Stockholm, Sweden (Bert Viklund) OMCP Olivier Montreuil private collection, Fleury-les-Aubrais, France (Olivier Montreuil) SMTFD State Museum of Zoology, Dresden, Germany (Olaf Jäger) UHHF Finnish Museum of Natural History, University of Helsinki, Helsinki, Finland (Heidi Viljanen) ZIN Zoological Institute of Russian Academy of Sciences, Saint-Petersburg, Russia (Andrey Frolov)

Specimens were prepared by standard methods used in entomological research. Genital structures were cleared in 10% KOH, rinsed in distilled water and either air dried or placed in glycerol. Photographs were taken with a Canon D100 camera equipped with a EF-S 60 macro lens and a Leica MZ9.5 stereo microscope equipped with a Leica DFC290 digital camera from dry specimens (habitus, aedeagus) or from specimens in glycerol (internal sacs of aedeagus). Partially focused serial images were combined in Helicon Focus software (Helicon Soft Limited) to produce completely focused images. Helicon Focus software was used with default settings and a number of stack images varied from 20–40. Minor stacking artifacts were not retouched, only general image enhancing (levels, background elimination and slight sharpening) was applied. Distributional records maps were prepared with ArcGIS software (ESRI Incorporated). Co-ordinates of the localities were taken from the specimen labels, if available, or from the United States National Geospatial-Intelligence Agency GEOnet Names Server (GNS, http:// geonames.nga.mil/gns/html). Labels are cited verbatim and separated by slash and our comments are in square brackets.

REVIEW OF THE MADAGASCAN ORPHNINAE Zootaxa 4207 (1) © 2016 Magnolia Press · 5 Diagnostic keys are provided for all taxa, but the species keys are suitable for males only. Differentiation of the females of closely related species by external morphology may be difficult or impossible in some cases as in many other scarab beetle groups. Also, females of many species are yet unknown.

External morphology of adult Orphninae

Below the main diagnostic and taxonomic characters of the adult Orphninae are described and illustrated based mostly on Triodontus itremoi Paulian, 1977. General appearance. Madagascan Orphninae are small to medium-sized, moderately convex beetles. The body length of the smallest specimen of Madecorphnus is about 4.5 mm, whereas the largest Triodontus is about 17 mm. Body coloration is formed by melanins and varies from brown to almost black. Orphnines show no coloration patterns and the color of the dorsal side is normally uniform. Madagascan taxa do not show any metallic tint of the body. The dorsal side is mostly shiny with sparse punctation. Males of some taxa may have frontoclypeal and prothoracic processes, which are genus and species specific and discussed in more detail below. Head. The head is wider than long, with the clypeus, frons, and vertex not separated from each other by sutures. For the descriptive purposes, we defined the following parts on the head dorsally (Fig. 1): anterior margin, anterior border, disc of frontoclypeus, canthus, eye, eye tubercle, and frontoclypeal process (horn or carina). In the majority of taxa of the Madagascan Orphninae, the head shape is subject to sexual dimorphism and allometric variability in males. The anterior margin of the frontoclypeus is more or less semicircular in most taxa. The frontoclypeus has a marked continuous border; the border is narrow in most taxa but can be wide and upturned medially in Renorphnus Frolov & Montreuil, 2009. The frontoclypeus is symmetrical in most taxa except for some males of Madecorphnus. In the latter case, the clypeal asymmetry correlates with the mandible asymmetry. The frontoclypeal suture is indistinct. The border between the clypeus and frons may be marked with a horn-shaped process (males of Triodontus and Pseudorphnus) or a low transverse ridge (Madagascan Pseudorphnus carinatus and some African Orphnus) but the comparative morphological data on this account are not available and in this study we do not separate frons and clypeus. The frontoclypeal process of the males is subject to the reasonable allometric variability and is almost absent in some specimens. The material available is not sufficient for a morphometric analysis, but we think that the variability is continuous rather than the males being dimorphic, i.e., the so-called “minor” and “major” males cannot be rigorously distinguished. The frontoclypeal process is always absent in females, although the females of some species of Triodontus may have a minute tubercle in the center of the frontoclypeus. The canthus is small, incompletely divides the eye into smaller dorsal and larger ventral parts. The males of Triodontus and Pseudorphnus have a small tubercle mediad of each eye; development of these ocular tubercles seems to correlate with the development of the frontoclypeal process. The antennae with 10 antennomeres, externally similar in all taxa. No cases of symphysocery, known in some African Orphnus (Frolov & Akhmetova 2016), have been encountered in Madagascan taxa. The antennal club has three compact antennomeres. The labrum is about two times wider than long, rounded laterally, sinuate anteriorly, densely setose. The apical part of the labrum protrudes past frontoclypeus and is visible in dorsal view. The mandibles are mostly symmetrical, about the same length, normally with 2–4 well-developed scissorial teeth. Exception to this are the males of Madecorphnus, which may have highly asymmetrical mandibles with the right mandible being up to two times or more longer than the left. The maxillae have separate lacinia and galea, which normally bear thick spinules along with the thin setae. The prementum is trapezoidal, densely setose, and somewhat rugose to almost smooth; a few species of Madecorphnus have two conical tubercles in the middle of the prementum. The mentum is about two times shorter than the prementum, concave. The gula is longer than wide, convex, having a shallow longitudinal groove in basal 3/4, and somewhat concave and setose apically. The glossae are feebly sclerotized and finely setose. The basal labial palpomere is about two times shorter than second and third palpomeres. Prothorax. The pronotum is trapezoidal, more or less rounded laterally. Shape and sculpture of pronotum are subject to sexual dimorphism and allometric variability in males of most taxa. We distinguish the following parts of the pronotum (Fig. 1): anterior margin, usually with a wide, flat or slightly convex border and membranous very anterior part adjacent to the head; base, mostly bordered and with a row of punctures; lateral margins, coarsely punctate and with long dense setae, anterior and posterior angles, disc of pronotum. The pronotal armature consists

6 · Zootaxa 4207 (1) © 2016 Magnolia Press FROLOV ET AL. of anterolateral and posterolateral pronotal processes, sometimes forming lateral pronotal ridges, and a central pronotal bulge, characteristic of the Triodontus males. Pronotal armature, when well developed (in so-called “major males”) is species specific, however such forms are rather rare in the collections and majority of specimens belong to the forms intermediate between “major” and “minor” males. Shape of the pronotal armature is also genus specific. The most complex pronotal armature is found in the males of the genus Triodontus. The prothoracic legs of the majority of Madagascan Orphninae are of typical scarabaeoid shape, with three rather large outer teeth to facilitate digging in soil. Protibiae of the males lack apical spurs but have a few apical setae on protibia (in the place of the absent spur) that are thicker than the others (Fig. 3A). Females have well- developed, rather long apical spurs. The procoxae of both sexes have a longitudinal hollow on anterior surface of procoxa (Fig. 3E); the hollow is concealed in the coxal cavities while a beetle is walking; it opens only when the prothoracic legs are appressed to the pronotum. Pterothorax. The elytra are convex, elongate, with marked humeral and apical humps. The first (sutural) elytral stria is well marked in most taxa except for Pseudorphnus; basally it is joined with basal elytral border which is similar to the stria in some species or appears as a row of contiguous punctures in others. The first elytral interval is more or less convex in most taxa. Other striae are distinct in some taxa (either as impressed lines or rows of punctures) or unclear; elytral intervals, except for sutural interval, are flat in most taxa. Punctation of elytra varies. The wings are fully developed in all Madagascan species. The wing venation can be described as of advanced

Scarabaeidae type (Fedorenko 2009) with reduced costal window, anal vein AA1+2 not reaching the wing margin, open cell 1a with strongly angulate posterior border, and well developed jugal sclerotization almost reaching the wing margin (Fig. 3F). The scutellum is exposed, semicircular to triangular, with rounded apex. The metepisternum has a slightly widened anterodorsal angle slightly overlapping the epipleuron, which is somewhat concave in this place. The orifice between the mesocoxal cavities, found in the Neotropical orphnines, is absent. The mesothoracic and metathoracic legs are similar in shape; the metafemora and metatibiae are usually about 1/8 longer than the mesofemora and mesotibiae. Tibiae somewhat triangular with two apical spurs; the inner margin only slightly concave and with one transverse keel. Metatibial spurs are separated by basal tarsomeres (Fig. 3C); metatibial spurs are adjoining (Fig. 3D). Longer tibial spur as long as the two basal tarsomeres. Claws 1/3 length of last tarsomere. The stridulatory apparatus is of typical orphnine shape. Stridulatory field is situated basally on the dorsal surface of metacoxa and consists of straight fine keels (Fig. 3B). Abdomen and external genitalia. The abdomen of the Orphninae has eight tergites and seven or eight sternites (Figs. 4A, B). Tergites 1–6 are weakly sclerotized, flexible, hidden under the wings and elytra. Tergites 7– 8 (propygidium and pygidium) are strongly sclerotized, more or less exposed (although not visible in dorsal view). Sternites 1–3 are modified, partly hidden under metacoxa. Sternite 1 is strongly reduced and its delimitation is difficult. Sternite 2 bears plectra and irregular, mostly longitudinal ridges. The plectra are triangular to somewhat rounded, with the apex being highly sclerotized and somewhat upturned. Sternite 3 is the widest, features a few secondary transversal ridges forming metacoxal cavities. Sternites 4–7 are mostly of the same width, relatively narrow medially. Sternite 8 is wider than sternites 4–7 and different in males and females. The male external genitalia of Madagascan Orphninae have subsymmetrical phallobase, symmetrical parameres, and variably armed internal sac of the aedeagus (endophallus). The phallobase in the Madagascan taxa is of the same type (Frolov 2013b) as in the Afrotropical taxa: it is strongly sclerotized dorsally and with a thin membrane ventrally. The shape of the parameres varies reasonably and in the most cases is species specific. The parameres of Triodontus are of complex shape: they are divided into inner and outer lobes, which may have processes and excavations. The internal sac consists of sclerites of different shape and number. The combination of the characters of the internal sac armature and the shape of the parameres reliably diagnose all species and in many cases, especially in Triodontus and Madecorphnus, are the only characters to separate the otherwise similar species. However, in Triodontus, the shape of the parameres is highly specific while the shape of the large symmetrical sclerite of the internal sac is similar in some species. In Madecorphnus, as opposed to Triodontus, the shape of the parameres is similar in some species while the shape of the internal sac armature is highly specific. Spiculum gastrale is triangular (Fig. 4D).

REVIEW OF THE MADAGASCAN ORPHNINAE Zootaxa 4207 (1) © 2016 Magnolia Press · 7 FIGURE 1. Triodontus itremoi, male in dorsal view (left legs removed): aa. prn—anterior angle of pronotum, acly—anterior margin of clypeus, al. prn. proc—anterolateral pronotal process, an. prn—anterior pronotal margin, bs. elyt—elytron base, bs. prn—base of pronotum, canth—canthus, d. prn.—disc of pronotum, elyt. ap. umb—apical elytral hump, istr. st—sutural interstria, labr—labrum, lm. prn—lateral margin of pronotum, mand—mandible, ocu—eye, pa. prn—posterior angle of pronotum, pl. prn. proc—posterolateral pronotal process, proc. ceph—frontoclypeal horn or tubercle, scut—scutellum, st. elyt—sutural margin of elytron, str. elyt I—elytral stria 1, tub. ocu—eye tubercle, umb. cpr—central pronotal bulge, umb. hum—humeral elytral hump.

8 · Zootaxa 4207 (1) © 2016 Magnolia Press FROLOV ET AL. FIGURE 2. Triodontus itremoi, male in ventral view (left legs and abdomen removed): ant—antenna, cav. mscox—mesocoxal cavity, cav. prcox—procoxal cavity, epipl—epipleuron, gul—gula, mand—mandible, max—maxilla, ment—mentum, mscox— mesocoxa, msepim—mesepimeron, msfem—mesofemur, msst—mesosternum, mstar—mesotarsus, mstib—mesotibia, mstr— mesotrochanter, mtcox—metacoxa, mtepis—metepisternum, mtfem—metafemur, mtst—metasternum, mttar—metatarsus, mttib—metatibia, mttr—metatorchanter, ocu—eye, prfem—profemur, propl—propleurae, protib—protibia, prst—prosternum, prstnl—prosternellum, prtar—protarsus, prtr—protrochanter, str. propl—propleural stria.

REVIEW OF THE MADAGASCAN ORPHNINAE Zootaxa 4207 (1) © 2016 Magnolia Press · 9 FIGURE 3. Triodontus itremoi. Protibia in ventral view (A), metacoxa in dorsal view (B), metatibia in apical view (C), mesotibia in apical view (D), procoxa in ventral view (E), wing (F); pars. str—stridulatory field, procx. fv—hollow of procoxa;

1a and 2a—anal cells; AA1+2, AAP, AA3b+(AA4+AP1+2), AP4—anal veins; aas—antero-apical sclerotization; cas—costo-apical sclerotization; CuA—anterior cubital vein; js—jugal sclerotization; MP3+4—posterior medial vein; rc—radial cell; RMP— fused posterior radial (RP3+4), anterior medial and posterior medial (MP1+2) veins; SV2—secondary “vein”. Wing venation terminology follows Fedorenko (2009). Not to scale.

10 · Zootaxa 4207 (1) © 2016 Magnolia Press FROLOV ET AL. FIGURE 4. Triodontus itremoi, abdomen (A—dorsal, B—ventral view): cav. mtcox—metacoxal cavity, plec—plectrum, propyg—propygidium, pyg—pygidium, stern2–stern8—abdominal sternites 2–8, terg1–terg6—abdominal tergites 1–6; Triodontus lemoulti, spiculum gastrale (C); Triodontus itremoi, female genitalia (D): burs. cop—bursa copulatrix, coxt—coxite, gl. acc—vaginal gland, gl. rec—spermatheca gland, ovd—oviduct, parapr—paraproct, rec. sem—spermatheca, scoxt— subcoxite, styl—stylus. Not to scale.

Female external genitalia consist of a few sclerites connected by membranes. More defined and sclerotized are the subcoxites, covering large vaginal accessory glands, and coxites, bearing styli. All three pairs of sclerites bear long setae apically and have a sensitive function. Other sclerites, proctiger (hemitergites IX), and paraprocts (epipleurites IX), are less distinct. Sexual dimorphism. The character immediately separating males from females in the orphnines is the absence

REVIEW OF THE MADAGASCAN ORPHNINAE Zootaxa 4207 (1) © 2016 Magnolia Press · 11 of the protibial spur in the former. This character is always distinct except, sometimes, for the males of Madecorphnus (see diagnosis of this genus). The two other characters are more general and valid for the majority of scarab beetles: 1) females usually have relatively smaller pronotum, distinctly narrower than elytra, and, 2) the head and pronotal armature is developed only in the males (but is subject to allometric variability: see diagnoses of the genera). Abdominal sternites differ in length. Sternite 8 in males is shorter and sometimes clearly sinuate medially; in some Triodontus species, it has a tubercle or concavity medially. The sexes do not differ in body coloration or pubescence. Also, all the specimens examined, both males and females, had fully developed wings.

Natural history of the Madagascan Orphninae

There are no direct data about feeding and nesting behavior of the adult Madagascan Orphninae. The shape of the mouthparts suggests that they are generalist saprophages, although the strongly asymmetrical mandibles in some Madecorphnus species probably prevents them from feeding. A number of specimens of different species were collected from pitfall traps baited with fish, chicken intestine, or dung. Although the rather short trap exposures in some cases (up to 48 hours) might suggest attraction to the baits, it is also possible that these were accidental captures due to the rather dense populations of the species. Available distribution records suggest that orphnines prefer forest biotopes and the distribution of all genera is biased to the areas with the remnants of the indigenous rain forests. The majority of the species are probably forest litter dwellers at all development stages. Exception to this might be the secondary adaptation to feeding on crops as has been reported for the larvae of Triodontus nitidulus (Randriamanantsoa et al. 2010).

Key to the Madagascar Orphninae genera (both sexes)

1. Protibiae with 2 outer teeth apically and 1 much smaller outer tooth basally (Fig. 7D)...... Pseudorphnus Benderitter - Protibiae with 3 outer teeth in apical half (Figs. 10H, 30C, 28F) ...... 2 2. Body length smaller (usually less than 7 mm); lateral margin of pronotum with 4 long setae: 2 near anterior margin, 1 in the middle, and 1 near posterior margin; some or all these setae may be abraded, but 4 chaetiferous punctures are always distinct; left and right mandibles may differ strongly in length in males, and frontoclypeus is somewhat asymmetrical in such cases; disc of pronotum smooth, without processes or excavations ...... Madecorphnus Paulian - Body length larger (usually longer than 7 mm); lateral margin of pronotum with numerous setae (at least more than 10) evenly distributed along the margin, setae may differ strongly in length; left and right mandibles of about same length, frontoclypeus symmetrical; disc of pronotum may be with more-or-less developed tubercles, ridges, and excavations in males ...... 3 3. Eyes smaller (width of eye in dorsal view more than 7 times smaller than distance between eyes); propleurae with a longitudinal ridge bearing a row of setae parallel to lateral margin of pronotum; disc of pronotum with more-or-less developed tubercles, ridges, and excavations in males, and smooth in females; apical outer tooth of protibia in males directed laterally, at reasonable angle to internal margin of tibia (Fig. 30C) ...... Triodontus Westwood - Eyes larger (width of eye in dorsal view less than 4 times smaller than distance between eyes); propleurae without a longitudinal ridge, only with a sparser row of setae; disc of pronotum with 2 feebly developed tubercles in middle or without tubercles; apical outer tooth of protibia almost parallel to internal margin of tibia (Fig. 28F) ...... Renorphnus Frolov & Montreuil

Genus Pseudorphnus Benderitter, 1913

Pseudorphnus Benderitter, 1913: 83. Type species. Orphnus coquerelii Fairmaire, 1868, by monotypy.

Diagnosis. Body length 6.5–13 mm, brown to dark brown. Mandibles subsymmetrical. Frontoclypeus with rounded anterior margin, with tubercle or longitudinal ridge in males; females have frontoclypeus without tubercles or ridges. Pronotum with deep excavation in the middle and with 2 horn or ridge-shaped lateral processes bordering excavation near anterior margin in males; females with pronotum convex or slightly impressed anteromedially. The shape of the head and prothoracic armature in males is subject to moderate allometric variability: they vary from fully developed as described above to only a small tubercle on the frontoclypeus and a shallow fossa on the disc of the pronotum anteriorly. Propleurae smoothly convex, without carinae separating anterolateral areas from basal area. Scutellum triangular, narrowly rounded apically, about 1/10 length of elytra. Elytra convex, with feebly

12 · Zootaxa 4207 (1) © 2016 Magnolia Press FROLOV ET AL. marked humeral humps. Sculpture of elytron varies. Wings fully developed. Metepisternum triangular, tapering caudally, with slightly widened anterodorsal angle slightly overlapping epipleuron. Protibiae with 2 strong outer teeth and 1 small tooth located basally. Phallobase strongly sclerotized dorsally and with a thin membrane ventrally. Parameres relatively long apices tapering or curved downwards, without setae. Internal sac of aedeagus with species-specific armature. Diagnostic characters. The most reliable characters to differentiate Pseudorphnus species are the shape of the parameres and the shape of the frontoclypeal and prothoracic armature (especially when fully developed). Body size also varies among the species. Species composition, distribution, and habitat. Pseudorphnus is endemic to Madagascar. Four species of the genus are known so far. The species are distributed chiefly along the eastern slopes of the Central Escarpment corresponding to the distribution of the Malagasy tropical wet forests. All species are apparently forest litter dwellers feeding on different rotten substances. One species, P. coquerelii, is quite common in the collections, which may indicate a secondary adaptation to the developing in the crop fields (similar to Triodontus nitidulus, see below). Nesting behavior and preimaginal stages are unknown.

Key to the Pseudorphnus species (males)

1 Elytra without distinct striae, punctate on disc; elytral intervals almost flat ...... 2 - Elytra with distinct striae, punctate or impunctate on disc; elytral intervals convex or flat ...... 3 2 Frontoclypeus with transverse carina (Figs. 5E–F), sculpture of frontoclypeus smooth; parameres with less acute apices (in lateral view, Fig. 5C) ...... P. carinatus Frolov - Frontoclypeus with conical tubercle medially (Figs. 6D–E), sculpture of frontoclypeus rugose; parameres with more acute apices somewhat depressed laterally (in lateral view, Fig. 6C)...... P. olsoufieffi Paulian 3 Elytra with densely punctate striae (Figs. 7A, E); elytral intervals somewhat convex; pronotal horns of male directed upward and forward (Figs. 7A–B); apices of parameres finger shaped (Fig. 7C); length 11–13 mm ...... P. hiboni Paulian - Elytra almost impunctate on disc, with striae visible as slightly elevated lines (Figs. 8A–C); elytral intervals almost flat; pronotal horns of male wider and directed upward (Figs. 8B, D); apices of parameres hook-shaped (in lateral view (Fig. 8G); length 6.5–8.5 mm...... P. coquerelii (Fairmaire)

Pseudorphnus carinatus Frolov, 2011 (Figs. 5A–G)

Pseudorphnus carinatus Frolov, 2011: 65.

Type material examined. Holotype (Figs. 5A–F), male, “MADAGASCAR: Province d'Antsiranana, Ampasindava, Foret d'Ambianivy, 3.9 km 181 S Ambaliha, elev. 600 m, 4–9 March 2001 / 13 47′55″S, 48 9′42″E coll. Fisher, Griswold et al. California Acad. of Sciences, pitfall trap in rainforest, collection code: BLF3250 / CASENT 8013699 / Holotypus Pseudorphnus carinatus Frolov det. 2010”. Paratype: single male with the same locality label as the holotype. Differential diagnosis. From other Pseudorphnus species, the males of P. carinatus can be separated by the characteristic shape of the frontoclypeal ridge. It is similar to P. olsoufieffi in the body size, sculpture of pronotum, and elytra, and in the shape of prothoracic ridges, but the males of P. olsoufieffi have a rather large, conical tubercle situated in the center of the frontoclypeus. The two species also differ in the sculpture of frontoclypeus (smooth in P. carinatus and rugose in P. olsoufieffi) and in the shape of the parameres (having more acute apices somewhat depressed laterally in P. olsoufieffi. Description. Male: body elongate, oval, strongly shiny (Figs. 5A–B). Color blackish brown, elytra and underside of body slightly lighter. Frontoclypeus slightly convex anteriorly, obtuse laterally, anterior margin setose and crenulate in dorsal view. Eyes relatively large (diameter larger than the distance between eye and gula in ventral view), incompletely divided by canthus into smaller dorsal and larger ventral parts. Frontoclypeus with a long (about 8/10 the width of frontoclypeus) transverse low ridge near anterior margin (Figs. 5E–F). Dorsal surface of head impunctate. Labrum bilobate, slightly sinuate in the middle and relatively feebly protruding past frontoclypeus. Length in the middle is 1/8 width (in dorsal view).

REVIEW OF THE MADAGASCAN ORPHNINAE Zootaxa 4207 (1) © 2016 Magnolia Press · 13 Pronotum 1.5 times wider than long, widest medially. Anterior margin with wide border, base with fine border. Lateral margins densely punctate, appearing crenulate in dorsal view. Disc of pronotum with deep excavation in the middle, with 2 slender longitudinal somewhat triangular ridges bordering the excavation (Figs. 5B, F). Surface of disc between the ridges smooth, without punctures. Sides of pronotum rugose posteriorly. Lateral margins with long, brown setae.

FIGURE 5. Pseudorphnus carinatus. Habitus (A—dorsal, B—lateral view), aedeagus in lateral view and parameres in dorsal view (C), labels of holotype (D), head in dorsal view (E), head and pronotum in dorso-lateral view (F), distributional records (G).

14 · Zootaxa 4207 (1) © 2016 Magnolia Press FROLOV ET AL. Scutellum triangular, narrowly rounded apically, about 1/8 length of elytra. Elytra convex, with marked humeral humps. Maximum width approximately at the middle. Only sutural stria is distinct and reaching apex of elytron. Elytra covered with sparse narrow and somewhat curved punctures, each bearing a short, yellow seta. Epipleura with long, sparse, brown setae. Base of elytra bordered and densely punctate with punctures similar to those in striae but more rugose and irregular. Protibiae of typical shape for Pseudorphnus. Lateral margin basad of outer teeth not crenulate. Apices with 3 robust, spur-like setae and a number of smaller setae. Protarsi well developed, about 4/5 length of protibiae. Claws 1/3 length of apical tarsomere. Apical protarsomere as long as protarsomeres 3 and 4 combined, as thick as other tarsomeres. Ventral surface of protibiae smooth with 2 rows of setae along sides and sparse, longer setae in the middle. Ventral surface of femora sparsely punctate, with 1 raised longitudinal line. Mesothoracic and metathoracic legs similar in shape; metafemora and metatibiae about 1/8 longer than mesofemora and mesotibiae. Tibiae somewhat triangular, with 2 apical spurs, with inner margin only slightly concave and with 1 transverse keel. Longer tibial spur as long as 2 basal tarsomeres. Claws 1/3 length of apical tarsomere. Femora almost impunctate, with 2 rows of long setae. Abdominal sternites irregularly punctate, pubescent; with dense, long setae. Sternite 8 medially as long as sternites 4–5 combined. Pygidium transverse, irregularly punctate, hidden under elytra. Aedeagus with short, acute apices (Fig. 5C). Female. Unknown. Variation . The two known specimens, the holotype and the paratype, are very similar to each other and differ only in the body size (length 10.5 and 11.0 mm). Distribution and habitat. This species is known from a single locality in northern Madagascar (Fig. 5G) situated in the subhumid forest region.

Pseudorphnus olsoufieffi Paulian, 1977 (Figs. 6A–G)

Pseudorphnus olsoufieffi Paulian, 1977: 1205; Frolov 2011: 66; Frolov & Montreuil 2006: 30.

Type material examined. Holotype (Figs. 6A–F), male, “Madacascar Antsianaka et lac Alaotra [Toamasina Province, Alaotra Lake] 2e Trimestre 1889 Perrot Freres / Pseudorphnus olsouffiefi n. sp. R.Paulian det. / HOLOTYPE” (MNHN). Additional material examined. MADAGASCAR: Toamasina: one female, “MADAGASCAR EST ANKASOKA ROUTE DE LAKATO [Ankasoka, road to Lakato] 1-[...]-1972 R.VIOSSAT REC.” (MNHN). Diagnosis. Pseudorphnus olsoufieffi is similar to P. cari natus in the body size, sculpture of the pronotum and elytra, and in the shape of the prothoracic ridges, but the males of the species differ in having a rather large, conical tubercle situated in the center of the frontoclypeus, as opposed to having a long transverse low ridge near the anterior margin in the later species. The two species also differ in the sculpture of the frontoclypeus (rugose in P. olsoufieffi and smooth in P. carinat us ) and in the shape of the parameres (having less acute apices not depressed laterally in P. carinatus). Description. Male. Elongate, shiny (Figs. 6A, B), colored uniformly brown beetle. Body length 9.5 mm. Frontoclypeus slightly convex anteriorly, obtuse laterally, anterior margin setose and crenulate in dorsal view. Eyes relatively large (diameter larger than the distance between eye and gula in ventral view), incompletely divided by canthus into slightly smaller dorsal and larger ventral parts. Frontoclypeus with a short conical tubercle in center, rugosely punctate (Figs. 6D, E). Labrum bilobate, slightly sinuate in the middle and relatively feebly protruding past frontoclypeus. Length in the middle is 1/8 width (in dorsal view). Pronotum 1.6 times wider than long, widest medially. Anterior margin with wide border, base with fine border. Lateral margins densely punctate, appearing crenulate in dorsal view. Disc of pronotum with deep excavation in the middle, with 2 slender longitudinal somewhat triangular ridges bordering the excavation (Figs. 6B, E). Surface of disc between the ridges smooth, without punctures. Sides of pronotum rugose posteriorly. Lateral margins with long, brown setae.

REVIEW OF THE MADAGASCAN ORPHNINAE Zootaxa 4207 (1) © 2016 Magnolia Press · 15 FIGURE 6. Pseudorphnus olsoufieffi. Habitus (A—dorsal, B—lateral view), aedeagus in lateral view and parameres in dorsal view (C), head in dorsal view (D), head and pronotum in dorsolateral view (E), labels of holotype (F), distributional records (G).

Scutellum triangular, narrowly rounded apically, about 1/10 length of elytra. Elytra convex, with marked humeral humps. Maximum width approximately at the middle. Elytral striae are feebly distinct. Elytra covered with sparse narrow and somewhat curved punctures, bearing a short, yellow seta; the

16 · Zootaxa 4207 (1) © 2016 Magnolia Press FROLOV ET AL. setae are mostly abraded on disc. Epipleura with long, sparse, brown setae. Base of elytra bordered and densely punctate with punctures similar to those in striae but more rugose and irregular. Lateral margin of protibia basad of outer teeth not crenulated. Apices with 3 robust, spur-like setae and a number of smaller setae. Mesothoracic and metathoracic legs are similar in shape; metafemora and metatibiae about 1/8 longer than mesofemora and mesotibiae. Tibiae somewhat triangular, with 2 apical spurs, with inner margin only slightly concave and with 1 transverse keel. Longer tibial spur as long as 2 basal tarsomeres. Claws 1/3 length of apical tarsomere. Femora almost impunctate, with 2 rows of long setae. The holotype lacks the abdomen. Aedeagus with parameres having short, acute, somewhat depressed laterally apices (Fig. 6C). Female. The only female specimen available, putatively identified as P. olsoufieffi, has the body length of 10.5 mm. It differs from the male in the rugose surface of the head, absence of the frontoclypeal horn or tubercle, presence of a distinct spur on protibiae, densely punctate disc of the pronotum, and absence of the pronotal excavations and ridges. The sexes are similar in general body shape and color. Distribution. This species is known from two localities in the East Madagascar (Fig. 6G). Exact habitat is unknown but the localities are situated in the rain forest region.

Pseudorphnus hiboni Paulian, 1959 (Figs. 7A–H)

Pseudorphnus hiboni Paulian, 1959: 143; Paulian 1977: 1207; Frolov & Montreuil 2006: 28.

Type material examined. Holotype (Figs. 7F–G), female, “Fort Dauphin [Toliary Province, Tôlanaro] (P. Hybon) 1943 / Pseudorphnus hiboni n. sp. Type / Type” (MNHN). Additional material examined. MADAGASCAR: Fianarantsoa: two females (UHHF) and three females (ZIN), “Madagascar Ranomafana NP, Vatoharana wet forest, 19.2.2002, fish bated pitfall trap, [21°18′00″S, 47°30′00″E] I. Hanski leg.”; five males, five females, "Madagascar, Province Fianarantsoa, Ranomafana National Park, Talatakely area, 900 m, mixed tropical forest, 4–16 January 2001 / 21.25041S 47.41945E D.H.&K.M.Kavanaugh, R.L.Brett, E.Elsom, and F.Vargas colls.pitfal traps COL-DHK-2001-001TN / CASENT 8006675" (CASC); three males, “Madagascar Ranomafana NP, Vatoharana wet forest, 2.2.2005, chicken int. trap [21°18′00″S, 47°30′00″E], H. Viljanen leg.” (ZIN); one male (MNHN) and one female (ZIN), “Madagascar Ranomafana NP, wet forest, 12.2.2003, dung trap, Ilkka Hanski leg.”; one male and one female, “Madagascar env. Fianarantsoa Ranomafana 900m 5–15.I.2001 S.Murzin leg” (ABCB). Diagnosis. Pseudorphnus hiboni is easily recognized by its large size: the body is longer than 11.0 mm, whereas the specimens of the other species are mostly shorter than 10.5 mm. It also differs in having the relatively slender horns in either sides of the excavation near the anterior margin of the pronotum in males, finger-shaped apices of the parameres, and the base of the elytra bordered and densely punctate with punctures similar to those in the striae and forming a “tile” pattern. Description. Male. Body elongated, strongly shiny (Figs. 7A, F). Color brown, elytra, and underside of body slightly lighter. Frontoclypeus slightly convex anteriorly, rounded laterally, anterior margin setose and crenulate in dorsal view. Eyes relatively large (diameter larger than the distance between eye and gula in ventral view), incompletely divided by canthus into smaller dorsal and larger ventral parts. Frontoclypeus with relatively long horn rounded apically. Dorsal surface of head impunctate. Labrum bilobate, slightly sinuate in the middle and relatively feebly protruding past frontoclypeus. Length in the middle is 1/8 width (in dorsal view). Pronotum 1.5 times wider than long; widest medially. Anterior margin with wide border, base with fine border. Lateral margins densely punctate, appearing crenulate in dorsal view. Disc of pronotum with deep, somewhat transverse excavation in the middle, with 2 horns bordering excavation near anterior margin. Pronotal horns as long as frontoclypeal horn but more slender, directed upward and forward (Fig. 7B). Surface of disc between the horns smooth, without punctures. Sides of pronotum rugose, somewhat tuberculate posteriorly. Lateral margins with long, brown setae.

REVIEW OF THE MADAGASCAN ORPHNINAE Zootaxa 4207 (1) © 2016 Magnolia Press · 17 FIGURE 7. Pseudorphnus hiboni. Habitus in dorsal view (A—male, F—female), head and pronotum in dorsolateral view (B), aedeagus in lateral view and parameres in dorsal view (C), protibia (D, male left, female right), base of elytra, SEM (E), labels of holotype (G), distributional records (H).

Scutellum triangular, narrowly rounded apically, about 1/10 length of elytra. Elytra convex, with feebly marked humeral humps. Maximum width approximately at the middle. Elytra with 7 distinct striae on disc. Striae with characteristic, narrow, semicircular punctures. Each puncture with a short,

18 · Zootaxa 4207 (1) © 2016 Magnolia Press FROLOV ET AL. yellow seta. Only sutural stria reaches elytral apex. Striae 2–5 about 2/3 length of elytron, striae 6–7 about 1/3–1/4 length of elytron and do not reach elytral base. Epipleura with long, sparse, brown setae. Base of elytra bordered and densely punctate with punctures similar to those in striae and forming a “tile” pattern (Fig. 7E). Protibiae of typical shape of Pseudorphnus, with 2 strong outer teeth and a small 1 located basally. Lateral margin basad of outer teeth not crenulate. Apices with 3 robust, spur-like setae and a number of smaller setae. Protarsi well developed, about 4/5 length of protibiae. Claws 1/3 length of apical tarsomere. Apical protarsomere as long as tarsomeres 3 and 4 combined, as thick as other tarsomeres. Ventral surface of protibiae smooth with 2 rows of setae along sides and sparse longer setae in the middle. Ventral surface of femora sparsely punctate, with 1 raised longitudinal line. Mesothoracic and metathoracic legs similar in shape; metafemora and metatibiae about 1/8 longer than mesofemora and mesotibiae. Mesotibiae and metatibiae somewhat triangular, with 2 apical spurs, with inner margin only slightly concave and with 1 transverse keel. Longer tibial spur as long as 2 basal tarsomeres. Claws 1/ 3 length of apical tarsomere. Femora almost impunctate, with 2 rows of long setae. Abdominal sternites irregularly punctate, pubescent with dense, long setae. Sternite 8 medially as long as sternites 4–5 combined. Pygidium transverse, irregularly punctate, hidden under elytra. Aedeagus (Fig. 7C) with slender, finger-shaped apices of parameres, feebly tapering apically. Female. The female (Fig. 7F) differs from the male in having rugose surface of the head, pronotum slightly impressed anteriorly with sparse punctures in the middle, absence of the frontoclypeal horn or tubercle, presence of the distinct spur of protibiae, and absence of the pronotal excavations and ridges. The sexes are similar in general body shape and color. Variation. All examined male specimens had well-developed prothoracic ridges and frontoclypeal horns, thus showing a relatively low allometric variability. Sculpture of the elytra varies with some specimens having no distinct rows of the punctures on the elytra, except for the first (sutural) stria. Body length of the examined specimens varied from 11.0 to 12.5 mm (males) and from 11.0 to 13.0 mm (females). Distribution and habitat. The species was described from Tôlanaro (formerly Fort Dauphin) in southern Madagascar. All the other findings were made in a few localities in the Ranomafana National Park (Fig. 7H). The park is situated in Fianarantsoa Province, about 100 km west of the Indian Ocean on the east-facing escarpment of Madagascar's central high plateau and covers about 43 000 ha of tropical wet forest with elevations varying from 374–400 m. Some of the examined specimens of P. hiboni were captured by pitfall traps baited with fish and chicken intestine or dung and exposed for approximately 45 hours including two nights (Heidi Viljanen, personal communication). Short-time exposures of the traps suggest that the beetles were attracted to the carrion rather than accidental captures.

Pseudorphnus coquerelii (Fairmaire, 1868) (Figs. 8A–H)

Orphnus coquerelii Fairmaire, 1868: 784. Pseudorphnus coquereli (Fairmaire): Paulian 1937: 14, 1977: 1206; Frolov & Montreuil 2006: 31.

Type material examined. Holotype (Figs. 8A, E), female, “madag: / TYPE / MUSÉUM PARIS 1906 Coll. Léon FAIRMAIRE / Pseudorphnus coquereli (Fairm.) Frolov det. 2010 / SYNTYPE / MNHN EC2303” (MNHN). Additional material examined. MADAGASCAR: Antsiranana: one female, “Madagascar Est m-if du Marojejy env-ons Manantenina 150 m. [Marojejy Range, environs of Manantenina, 150 m. a.s.l., 14°28′59″S, 49°49′00″E], II.1974 A.Peyrieras” (MNHN); one female, “Nosy Mitsio [Nosy Mitsio Island, 12°54′00″S, 48°36′00″E, 13–14.I.1960, R. Paulian leg.]” (MNHN); one unsexed specimen, “Antalaha [14°52′59″S, 50°16′59″E]” (MNHN); one male, three females, “Madagascar R-on Antalaha XI.35 Vadon! [Antalaha District, Vadon leg.]” (MNHN); one female, “Madagascar R-on Antalaha XII.35 Vadon!” (MNHN); two males, one female, “Sambirano [Sambirano River] N.O. Madagasc.” (MNHN); two males, one female, “Sambirano N.O. Madagasc.” (MHNG); one female, “Museum Paris Madagascar Nossi-Be [Nosy Be Island] H. Pierron 1885” (MNHN); two females, “Andranovolo [15°01′59″S, 50°07′00″E] / Madagascar R-on Antalaha XII.38 Vadon” (MNHN); one male, “Mad. Vohemar, Ampanefena [13°52′00″S, 49°58′00″E] XII.36 Vadon” (MNHN); one female, “Vohemar

REVIEW OF THE MADAGASCAN ORPHNINAE Zootaxa 4207 (1) © 2016 Magnolia Press · 19 [13°21′36″S, 50°00′00″E] Madagascar” (NMPC); one unsexed specimen, “Nosy Mitsio [Nosy Mitsio Island, 12°53′56″S, 48°35′56″E]” (MNHN); Toamasina: one female, “Madagascar Andasibe [18°55′59″S, 48°25′00″E] X.1978–II.1980 leg. Wojnarovich” (HNHM); one male, “Museum Paris Moramanga [18°55′59″S, 48°12′00″E]” (MNHN); one female, “Madagascar R-on Maroantsetra Vadon! [Maroantsetra District]” (MNHN); one male, “Nandihizana [15°45′00″S, 49°19′00″E] / Madagascar R-ne Maroantsetra XII.1937 Vadon!” (MNHN); two males, “Museum Paris Madagascar Fenerive [17°22′00″S, 49°25′00″E] F.Genot 1904” (MNHN); three males, one female, “Madagascar Ambodivoangy [15°17′49″S, 49°36′47″E] I.1946 Vadon!” (MNHN); one male, same label but XII.1946, (MNHN); one male, same label but XI.1945 (MNHN); one female, “MADAGASCAR: TAM Station Alaotra 27.xii.1990 A.Pauly col. jardin bac j. [Alaotra Lake, TAM Station , 17°40′59″S, 48°27′00″E]” (MRAC); one male, two females, “Madagascar Antongil (Sud) Mocquerys 1897 [Antongil Bay]” , (MNHN); one male, one female, “Museum Paris Madagascar Baie D'Antongil A. Mocquerys 1898 [Antongil Bay]” (MNHN); one male and one female, “S. Baie Antongil [southern Antongil Bay]”, (MNHN), two females and two unsexed specimens, (SMTFD); two females, “MADAGASCAR: Toamasina Ambatovy, 12.4 km NE Moramanga elev 1080 m pitfall trap, 4–7 March 2007 18˚50'22"S 048˚18'30"E California Acad. of Sciences coll. B.L.Fisher et al. montane rainforest BLF16917” (CASC); one unsexed specimen, “MADAGASCAR: Toamasina Ambatovy, 12.4 km NE Moramanga elev 1080 m pitfall trap, 4–7 March 2007 18˚50'22"S 048˚18'30"E California Acad. of Sciences coll. B.L.Fisher et al. montane rainforest BLF16917” (MNHN); two males, one female, “Antongil-Bai [Antongil Bay] Madagaskar” (SMTFD); Fianarantsoa: one male, one female, “Madagascar Vohilava 60 m. [60 m, 21°46′00″S, 47°55′00″E] Faraony” (MNHN); one female, “Madagascar Region de Mananjary [21°13′00″S, 48°19′59″E] A. Mathiaux” (MNHN); Mahajanga: one female, “Madagascar Manditsara / Amboaboa XI.35 Vadon [15°55′59″S, 48°43′00″E]” (MNHN); one female, “Madagascar Mananarara Nd. XI.35 Vadon [15°55′59″S, 48°43′00″E]” (MNHN); one unsexed specimen, “Mahizina [15°52′59″S, 48°04′00″E]” (MNHN); one female, “Maromandia [14°13′00″S, 48°04′59″E] / Madagascar R-on Antalaha XII.38 Vadon” (MNHN); one male, “Museum Paris Madagascar Prov D'Analalava Maromandia [Analalava District, Maromandia, 14°13′00″S, 48°04′59″E] R. Decary 1923” (MNHN); one female, “Madagascar Majunga [15°43′00″S, 46°19′00″E] Ch.Alluaud 1897” (MNHN); one male, one female, “Ste Marie de Madagascar [Sainte Marie de Madagascar, 16°06′00″S, 46°37′59″E]” (MNHN); Toliara: one female, “Betioky (Tulear) Mdgk. lg. Clement [Betioky, 23°43′00″S, 44°22′59″E]” (MHNG); Madagascar (no exact locality): one male (MHNG); one female, 21.X, Kaudern leg. (NRS); one male (HNHM); nine males, four females (MNHN); one female (SMTFD). Diagnosis. Pseudorphnus coquerelii differs from the other species of the genus in having hook-shaped parameres (in lateral view) and in the shape of the prothoracic longitudinal ridges in males, which are somewhat bimodal in lateral view (although this character may be unreliable in the specimens with poorly developed prothoracic ridges). From P. hiboni, it also differs in its smaller size and sculpture of the elytra. Description. Male. Body elongated, strongly shiny (Figs. 8A–C). Color brown, elytra and underside of body sometimes lighter. Frontoclypeus slightly convex anteriorly, rounded laterally, anterior margin setose and crenulate in dorsal view. Eyes relatively large (diameter larger than the distance between eye and gula in ventral view), incompletely divided by canthus into somewhat smaller dorsal and larger ventral parts. Frontoclypeus with relatively short carina-like horn, which is 1/3 the width of frontoclypeus basally. Dorsal surface of head impunctate. Labrum bilobate, slightly sinuate in the middle and relatively feebly protruding past frontoclypeus. Length in the middle is 1/6 width (in dorsal view). Pronotum 1.5 times wider than long; widest medially. Anterior margin with wide border, base with fine border. Disc of pronotum with deep excavation in the middle, with 2 relatively high slender longitudinal lateral ridges, somewhat bimodal in lateral view, bordering the excavation (Fig. 8D). Surface of disc between the ridges smooth, without punctures. Lateral sides with elongate irregular punctures. Lateral margins with long, brown setae. Scutellum triangular, rounded apically, about 1/10 length of elytra. Elytra convex, with feebly marked humeral humps. Maximum width approximately at the middle. Elytra with a distinct sutural stria and 2–3 more-or-less distinct longitudinal lines apparently corresponding to 3, 5, and 7 striae. Lines with minute, elongate punctures. Only sutural stria reaches elytral apex. Epipleura with long, sparse, brown setae. Base of elytra densely punctate with punctures similar to those in striae (Fig. 8F). Protibiae with 2 strong outer teeth and a small tooth located basally. Lateral margin basad of outer teeth not crenulate. Apices with 3 robust, spur-like setae and a number of smaller setae. Protarsi well developed, about 4/5

20 · Zootaxa 4207 (1) © 2016 Magnolia Press FROLOV ET AL. length of protibiae. Claws 1/3 length of apical tarsomere. Apical protarsomere 2/3 the length of tarsomeres 3 and 4 combined, as thick as other tarsomeres. Ventral surface of protibiae smooth with 2 rows of setae along sides and sparse longer setae in the middle. Ventral surface of femora sparsely punctate, with 1 raised longitudinal line.

FIGURE 8. Pseudorphnus coquerelii. Habitus (A—female holotype in dorsal view, B—male in dorsal view, C—female in dorsal view, D—male in lateral view), labels of holotype (E), base of elytra, SEM (F), aedeagus in lateral view and parameres in dorsal view (G), distributional records (H).

REVIEW OF THE MADAGASCAN ORPHNINAE Zootaxa 4207 (1) © 2016 Magnolia Press · 21 Mesothoracic and metathoracic legs similar in shape; metafemora and metatibiae slightly longer than mesofemora and mesotibiae. Tibiae somewhat triangular, with 2 apical spurs, with inner margin only slightly concave and with 1 transverse keel. Longer tibial spur as long as 2 basal tarsomeres. Claws 1/3 length of last tarsomere. Femora almost impunctate, with 2 rows of long setae. Abdominal sternites irregularly punctate, pubescent with dense, long setae. Sternite 8 medially as long as sternites 4–5 combined. Pygidium transverse, irregularly punctate, hidden under elytra. Parameres with slender, hook-shaped apices (Fig. 8G). Female. The female (Fig. 8C) differs from the male in the rugose surface of the head, absence of the frontoclypeal horn or tubercle, presence of the distinct spur on protibiae, densely punctate disc of the pronotum, and absence of the pronotal excavations and ridges. The sexes are similar in general body shape and color. Variation. Some male specimens have poorly-developed frontoclypeal horns and prothoracic ridges and are similar to females. Body length of the examined specimens varied from 6.5–9.0 mm (males) and from 6.0–7.5 mm (females). Distribution and habitat. Pseudorphnus coquerelii is one of the most widespread Madagascar orphnine species. It is known from relatively large number of localities in the northern and eastern Madagascar as well as from the smaller islands near the northwestern coast (Fig. 8H). Most specimens originate from the regions predominantly covered with wet lowland forests but a few were collected from distant localities in western Madagascar (dry deciduous forests and succulent woodlands). As in the case of T. nitidulus (see below), wide distribution and high abundance of this species may be a result of the secondary adaptation to feeding on the cultivated crops.

Genus Madecorphnus Paulian, 1992

Madecorphnus Paulian, 1992: 171; Frolov 2010a, 2010c. Drepanognathus Lansberge, 1886: 92 (in part: Paulian 1977; synonym of Orphnus MacLeay: Frolov 2005). Sissantobius Ritsema, 1888: 217 (in part: Paulian 1937, 1977; synonym of Orphnus MacLeay: Frolov 2005). Type species. Drepanognathus falciger Lansberge, 1886, designated by Paulian (1977).

Diagnosis. Madecorphnus are small-sized (4.5–7.0 mm) beetles with mostly uniform, brown to black coloration. Upper side of the body has a few setae with distinct locations: 1 seta on the elytral base near epipleuron, 1 seta on the posterior angle of the pronotum, 1 seta approximately in the middle of the lateral margin of the pronotum, 2 setae on the anterior angle of the pronotum, 2 setae on the lateral sides and 2 setae on the anterior margin of the frontoclypeus. Mandibles asymmetrical, subequal in length in females and subequal to strongly unequal in males. Frontoclypeus symmetrical to asymmetrical in males, wide, bordered anteriorly, smooth, without tubercles or ridges. Pronotum wider than long, smooth, similar in both sexes, without any depressions, tubercles, or ridges. Propleurae smooth, convex, without carinae separating anterolateral areas from basal area. Scutellum triangular, narrowly rounded apically, about 1/10 the length of the elytra. Elytra convex, with feebly marked humeral humps, smooth, with only first stria distinct. Wings fully developed. Metepisternon triangular, tapering caudally, with slightly widened anterodorsal angle slightly overlapping epypleuron. Protibiae with 3 outer teeth. Phallobase strongly sclerotized dorsally and with a thin membrane ventrally. Parameres relatively long, their apices tapering or curved downwards, without setae. Internal sac of aedeagus has species-specific armature. Sexual dimorphism in Madecorphnus is weaker than in the other Orphninae taxa. The sexes mostly differ in the shape of the mandibles. Males with the relatively feebly-developed mandibles are similar to females. In Madecorphnus, the sexual dimorphism character common for all Orphninae genera, the absence of distinct protibial spur in males, is less expressed. Often, the apical seta is longer and more robust than the others and is similar to the spur (Fig. 10H). However, examination of this seta at higher magnification shows that it bears minute, feebly visible setae while the true spurs are always smooth. Madecorphnus males also lack any frontoclypeal horns and prothoracic ridges, tubercles or excavations, which are found in the majority of species of almost all genera of the Orphninae. Diagnostic characters. The shape of the mandibles is species specific at least in some species, but it is subject

22 · Zootaxa 4207 (1) © 2016 Magnolia Press FROLOV ET AL. to allometric variability and therefore cannot always be a reliable diagnostic character. The most useful characters to separate species proved to be the number and shape of the internal sac sclerites and the shape of the parameres. Species composition, distribution, and habitat. Madecorphnus is endemic to Madagascar and the most speciose genus of the Madagascan Orphninae with 19 described species. The species are distributed throughout the island but mostly in its eastern part. The notable exception is M. falculoides, which was found in the western part. The majority of the localities agree well with the current distribution of the remnants of indigenous forests. Little is known about bionomy of Madecorphnus. Some specimens were collected in carrion baited pitfall traps. However, it is unknown if they were attracted to carrion or captured accidently. A few specimens were collected in flight intercept traps or sifted out of forest litter. Madecorphnus species are probably generalist saprophagous litter dwellers but the labels of most of the collected specimens lack any data about the way the beetles were collected. Nesting behavior and preimaginal stages are so far unknown. Remarks. The first known species of the genus Madecorphnus and its type species, M. falciger (Lansberge, 1886), was originally described in the genus Drepanognathus Lansberge, 1886, along with D. mandibularis Lansberge, 1886 (currently Orphnus mandibularis). Since the generic name Drepanognathus was preoccupied, the replacement name Sissantobius Ritsema, 1888 was used for the two species. The name Madecorphnus was proposed by Paulian (1977) to accommodate M. falciger and S. falculoides Paulian from Madagascar and to separate these two species from African Sissantobius mandibularis (sensu Paulian), the type species of the genus Sissantobius. Synonymy of the names Drepanognathus, Sissantobius, and Madecorphnus is discussed in more detail in Paulian (1992) and Frolov (2005). Paulian (1977: 1204) illustrated the peculiar aedeagus and evaginated internal sac of M. falciger (sensu Paulian), but these illustrations do not agree with the aedeagus of the type of M. falciger. The shape of the parameres is similar to that of M. niger Frolov, 2010, but the internal sac with numerous short spinules having wide round bases is unknown in any described Madecorphnus species. We failed to find this specimen, apparently belonging to an undescribed species, in the collection of MNHN.

Key to the Madecorphnus species (males)

1. Right mandible with a tooth on inner side of scissorial part (Fig. 9A) ...... 2 - Right mandible without a tooth on inner side of scissorial part (Fig. 13A) ...... 5 2. Mentum without tubercles; parameres slender in dorsal view (Fig. 9B); internal sac of aedeagus with 2 large bifurcated sclerites and a few dozen spinules (Figs. 9E–F)...... M. falculoides (Paulian) - Mentum with 2 tubercles (Fig. 11F); parameres and internal sac not as above ...... 3 3. Internal sac of aedeagus with 3 sclerites, 2 larger and 1 smaller (Figs. 10E–F) ...... M. montreuili Frolov - Internal sac of aedeagus with 2 bifurcated sclerites (Figs. 11D–E, 12C) ...... 4 4. Sclerites of internal sac of aedeagus smaller (Figs. 11D–E); body narrower and eyes smaller (Fig. 11A) . . . M. barclayi Frolov - Sclerites of internal sac of aedeagus larger (Fig. 12C); body wider and eyes larger (Fig. 12A) ...... M. tuberculatus Frolov 5. Internal sac of aedeagus with long sclerites but without fields of small spinules (Fig. 13D)...... M. fisheri Frolov - Internal sac armature not as above ...... 6 6. Apices of parameres without lateral teeth (Fig. 14C) ...... 7 - Apices of parameres with more-or-less developed lateral teeth (Fig. 16C) ...... 8 7. Parameres without lateral excavations, about 1.5 times shorter than basal sclerite of aedeagus (Fig. 14C) . .M. pauliani Frolov - Parameres with lateral excavations, about 2 times shorter than basal sclerite of aedeagus (Fig. 15B) ...... M. niger Frolov 8. Head and pronotum with fine, dense punctation; internal sac of aedeagus with 3 large sclerites and an area with smaller spinules (Figs. 16E, F)...... M. punctatus Frolov - Head and pronotum with indistinct to sparse punctation (punctures separated by at least 3 times their diameter); internal sac of aedeagus with different armature ...... 9 9. Parameres with large lateral teeth (Fig. 17B) ...... 10 - Parameres with small lateral teeth (Fig. 19B) ...... 11 10. Internal sac of aedeagus with 1 long, spinule-shaped sclerite and with 3 small, short spinules (Figs. 17D–E) ...... M. dentatus Frolov - Internal sac of aedeagus with 2 tooth-shaped sclerites, 1 longer sclerite, and a few fields of minute spinules, 1 of which forms a finger-shaped process of the sac (Fig.18 C) ...... M. saintemariensis Frolov 11. Internal sac of aedeagus with 1 sclerite composed of 2 long slender spinules connected basally (Figs. 19D–E) ...... M. simplex Frolov - Internal sac of aedeagus with 2 or more separate sclerites ...... 12 12. Internal sac of aedeagus with 2 rather large sclerites: a spur-like sclerite, and an asymmetrical, claw-shaped sclerite (Figs. 20D–E) ...... M. aquilonius Frolov

REVIEW OF THE MADAGASCAN ORPHNINAE Zootaxa 4207 (1) © 2016 Magnolia Press · 23 - Internal sac of aedeagus with different armature ...... 13 13. Parameres with small teeth at extreme apex (Fig. 21B, lateral view); body length greater than 7.0 mm. . . . M. falcatus Paulian - Parameres of different shape; body length less than 6.0 mm ...... 14 14. Internal sac of the aedeagus with 2 large, curved paired sclerites and 1 long, spur-shaped sclerite (Figs. 22D–E). …………. 15 -. Internal sac armature different ...... 16 15 Curved paired sclerites larger, semicircular; spur-shaped sclerite shorter ...... M. falciger (Lansberge) - Curved paired sclerites smaller, comma-shaped; spur-shaped sclerite longer ...... M. hanskii Frolov 16 Internal sac of aedeagus with 2 large, separate sclerites and numerous small spinules, some of which are robust and almost tooth-shaped and may be arranged into 2 indistinctly separated clusters ...... M. cuccodoroi Frolov - Internal sac of aedeagus with 2 or more separate sclerites and numerous small spinules; the latter are fine and are all approximately the same size ...... 17 17 Parameres less acute in lateral view (Fig. 25B); internal sac of the aedeagus with 1 longer sclerite and 1 smaller, somewhat bifurcated sclerite (Figs. 25D–E) ...... M. brunneus Frolov - Parameres more acute in lateral view (Fig. 26B); internal sac with 2 longer (1 doubled) and 2 smaller sclerites (Figs. 26D–E) ...... 18 18 Parameres less tapering apically, lateral teeth closer to paramere apices (Fig. 26B); eyes larger, rounded in dorsal view (Fig. 26A)...... M. perinetensis Frolov - Parameres more tapering apically, lateral teeth not so close to paramere apices (Fig. 27B); eyes smaller, narrowly elongate in dorsal view (Fig. 27A) ...... M. peyrierasi Frolov

Madecorphnus falculoides (Paulian, 1977) (Figs. 9A–G)

Sissantobius falculoides Paulian, 1977: 1203. Madecorphnus falculoides: Paulian 1992: 169; Frolov 2010c: 1100.

Type material examined. Holotype (Figs. 9A–B, D–F), male, “MADAGASCAR Ouest : Antsalova, Antsingy. R.N. 9 I-1975 A. Peyrieras / Sissantobius falculoides n. sp. R.Paulian det. / HOLOTYPE” (MNHN). Paratypes: one male and four females with the same locality data as the holotype (MNHN). Diagnosis. This species can be distinguished from other Madecorphnus species in having the right mandible with acute tooth on inner side of scissorial part (Fig. 9A), distinctive shape of the parameres (Fig. 9B) and internal sac armature (Figs. 9E–F). Description. Male. Body length 5.0 mm. Color uniformly dark brown to black. Right mandible slender, about 1.4 times longer than left, slightly curved, with a slender, long tooth in the middle (Fig. 9A). Labrum trapezoidal, feebly visible in dorsal view. Frontoclypeus distinctly asymmetrical, slightly convex anteriorly, obtuse, with 2 long and a number of smaller setae. Frontoclypeus slightly depressed in the middle anteriorly, finely punctate. Pronotum 1.5 times wider than long, widest medially. Margins with relatively wide border, lateral margins with 4 long setae. Pronotum evenly convex, finely punctate. Elytra convex, with distinct humeral and apical tubercules. Maximum width approximately at basal 1/3. First stria distinct and reaching the apex of elytron, other striae indistinct. Epipleura with long, sparse setae. Base of elytra with border from scutellum to humeral callus. Elytra punctate with sparse, relatively large punctures with minute punctures in between. Elytra with 2 longitudinal stria each, appearing as double lines on the disc of elytron; lateral striae less distinct as medial stria. Protibiae with 3 outer teeth, lateral margin basad of outer teeth not crenulate. Apex with robust, spur-like seta and a few smaller setae basally. Mesothoracic and metathoracic legs similar in shape. Longer tibial spur shorter than 2 basal tarsomeres in mesothoracic legs and shorter than 3 tarsomeres in metathoracic legs. Parameres symmetrical, narrow and tapering in dorsal view, with 2 small lateral teeth not visible in dorsal view (Fig. 9B). Internal sac with 2 large double sclerites and large area of relatively robust adjacent spinules (Figs. 9E– F). Female. Female (Fig. 9C) differs from the male in having short subsymmetrical mandibles, long protibial spur, and relatively wider elytra. Distribution and habitat. This species is known from one locality in western Madagascar (Fig. 9G).

24 · Zootaxa 4207 (1) © 2016 Magnolia Press FROLOV ET AL. FIGURE 9. Madecorphnus falculoides. Habitus in dorsal view (A—male, C—female), aedeagus in lateral view and parameres in dorsal view (B), labels of holotype (D), internal sac of aedeagus (E—photograph, F—schematic representation of armature), distributional records (G).

Madecorphnus montreuili Frolov, 2010c: 1102.

Type material examined. Holotype (Figs. 10A–B, D–F), male, “MADAGASCAR ANTANAMBE 25/01/90 P.E.C. Foret PROG. MAG188-007 / Holotypus Madec. montreuili Frolov 2009” (MNHN). Additional material examined. Five males and two females with the same data as the holotype (MNHN). Diagnosis. This species is similar to M. barclayi Frolov, 2012 and M. tuberculatus Frolov, 2014 in having 2 distinct tubercles on the mentum and bidentate apex of right mandible, but differs from them in having internal sac of aedeagus with 3 conical to spur-shaped sclerites, 2 larger and 1 smaller. Description. Male. Body length 5.8–6.0 mm. Color uniformly blackish brown. Right mandible about 1.5 times longer than left, strongly curved, with a tooth behind apex (Fig. 10A). Labrum trapezoidal, its length about 1/5 width (in dorsal view). Frontoclypeus slightly asymmetrical, slightly convex anteriorly, obtuse, with 2 long and a number of smaller setae. Frontoclypeus somewhat depressed in the middle anteriorly, smooth. Pronotum 1.8 times wider than long, widest medially. Margins with relatively wide border, lateral margins with 4 long setae. Pronotum evenly convex, minutely punctate. Elytra convex, with distinct humeral and apical tubercules. Maximum width approximately at basal 1/3. First stria distinct and reaching the apex of elytron, other striae indistinct. Epipleura with long, sparse, brown setae. Base of elytra with border from scutellum to humeral callus. Elytra punctate with sparse relatively large punctures in the middle of each elytron near stria 1. Protibiae with 3 outer teeth, lateral margin basad of outer teeth not crenulate. Apex with robust, spur-like seta and a few smaller setae basally. Mesothoracic and metathoracic legs similar in shape. Longer tibial spur shorter than 2 basal tarsomeres in mesothoracic legs and shorter than 3 tarsomeres in metathoracic legs. Parameres symmetrical, relatively wide in dorsal view, with 2 distinct teeth laterally (Fig. 10B). Internal sac with 2 large and 1 smaller sclerites (Figs. 10E–F). Female, The female (Fig. 10C) differs from male in having short subsymmetrical mandibles, long protibial spur, and relatively wider elytra. Distribution and habitat. Madecorphnus montreuili is known from one locality in eastern Madagascar (Fig. 10G). This area, south of Antongil Bay, has a few patches of indigenous rain forest inland of Antanambe, where the beetles were collected.

Madecorphnus barclayi Frolov, 2012 (Figs. 11A–G)

Madecorphnus barclayi Frolov, 2012b: 162.

Type material examined. Holotype (Figs. 11A–F), male, “MADAGASCAR E. Masoala, 50–470 m, R. Antsamanarana, 3–7/xii/1993 Flight Intercept Trap BMNH(E) 1994-138 / Holotypus Madec. barclayi Frolov 2009” (BMNH). Paratypes: two specimens with the same locality label as the holotype, male (ZIN) and female (BMNH). Diagnosis. This species is most similar to M. tuberculatus in having internal sac of aedeagus with 2 sclerites but the shape of the sclerites is different: they are much smaller and less sclerotized. It also differs in having narrower body with smaller eyes. Description. Male. Body length 5.6–6.0 mm. Color uniformly castaneous, elytra somewhat paler. Right mandible about 1.5 times longer than left, strongly curved, with a tooth behind apex (Fig. 11A). Labrum trapezoidal, with rounded anterior angles, its length about 1/5 width (in dorsal view). Mentum with 2 conical tubercles (Fig. 11F). Frontoclypeus slightly asymmetrical, almost flat anteriorly, obtuse, with 1 long and a few smaller setae. Head dorsally with minute punctures separated by more than 5 puncture diameters. Pronotum 1.5 times wider than long, widest medially. Margins with relatively wide border, lateral margins with 4 long setae. Punctation on pronotum similar to that on head.

26 · Zootaxa 4207 (1) © 2016 Magnolia Press FROLOV ET AL. FIGURE 10. Madecorphnus montreuili. Habitus in dorsal view (A—male, C—female), aedeagus in lateral view and parameres in dorsal view (B), labels of holotype (D), internal sac of aedeagus (E—photograph, F—schematic representation of armature), distributional records (G), protibia (H, male left, female right).

REVIEW OF THE MADAGASCAN ORPHNINAE Zootaxa 4207 (1) © 2016 Magnolia Press · 27 FIGURE 11. Madecorphnus barclayi. Habitus in dorsal view (A), aedeagus in lateral view and parameres in dorsal view (B), labels of holotype (C), internal sac of aedeagus (D—photograph, E—schematic representation of armature), mentum (F), distributional records (G).

Elytra convex, with distinct humeral humps. Maximum width approximately at basal 1/3. First stria distinct and reaching the apex of elytron, other striae indistinct. Epipleura with long, sparse, brown setae. Base of elytra with border connected to first elytral interval. Elytra punctate with sparse relatively large punctures on disc.

28 · Zootaxa 4207 (1) © 2016 Magnolia Press FROLOV ET AL. Protibiae with 3 outer teeth, lateral margin basad of outer teeth not crenulate. Apex with robust, spur-like seta and a few smaller setae basally. Mesothoracic and metathoracic legs similar in shape, metathoracic legs 1.3 times longer than mesothoracic legs. Longer tibial spur shorter than 2 basal tarsomeres in mesothoracic legs and almost as long as 3 tarsomeres in metathoracic legs. Parameres symmetrical, relatively wide in dorsal view, with 2 distinct teeth laterally (Fig. 11B). Internal sac with 2 slender bifurcated sclerites (Figs. 11D–E). Female. The female differs from male in having short mandibles and long protibial spur. Variation. Body length of examined specimens 5.6 mm (male) and 5.5 mm (female). Apart from the body length the male paratype is very similar to the holotype. Distribution and habitat. Madecorphnus barclayi is known from one locality in the northern part of the Masoala Peninsula (Fig. 11G). It is the only species of the genus recorded from the peninsula so far. The Masoala Peninsula is one of the largest forest blocks in Madagascar and a top conservation priority due to its lowland humid forests, now rare elsewhere in Madagascar (Du Puy & Moat 1996, Kremen et al. 1999).

Madecorphnus tuberculatus Frolov, 2014 (Figs. 12A–D)

Madecorphnus tuberculatus Frolov, 2014: 38.

Type material examined. Holotype (Figs. 12A–C), male, “MADAGASCAR: Toamasina Prov., Reserve Betampona Camp Vohitsivalana, 37.1 km 338° Toamasina elev 520 m 1–3 Dec 2005 / 17° 53' 12" S 049° 12' 09" E California Acad. of Sciences coll. Brian L. Fisher et al. sifted litter rainforest collection code: BLF13242” (CASC). Diagnosis. This species is similar to M. barclayi Frolov in having the internal sac of the aedeagus with 2 sclerites but the shape of the sclerites is different: they are much larger and heavily sclerotized. It also differs from M. barclayi in having a relatively wider body with relatively larges eyes. Description. Male. Body length 5.8 mm. Color uniformly dark brown, legs somewhat paler (Fig. 12A). Right mandible about 1.5 times longer than left, strongly curved, with a tooth behind apex. Labrum trapezoidal, with rounded anterior angles, its length about 1/8 width (in dorsal view). Mentum with 2 conical tubercles. Frontoclypeus slightly asymmetrical, almost flat anteriorly, obtuse, with a group of 4 setae displaced to the left side. Canthus and frontal suture indistinct. Head dorsally with minute punctures separated by more than 4 puncture diameters. Pronotum 1.7 times wider than long, widest medially. Disc of pronotum convex, without any depressions, tubercles, or ridges. Punctation on pronotum finer than on head, almost indistinct. Margins with relatively wide border, lateral margins with 4 long setae: 1 seta on basal angle, 1 seta approximately in the middle of lateral margin, and 2 setae on the apical angle. Base with a row of feebly elongate punctures (smaller and sparser than in M. fisheri). Elytra convex, with distinct humeral and apical tubercules. Maximum width approximately in the middle. First stria distinct and reaching apex of elytron, other striae indistinct. Epipleura with long, sparse, brown setae. Base of elytra with border connected to first elytral interval. Elytra with double punctation: sparse, large punctures on disc and minute, feebly visible punctures throughout elytron. Protibiae with 3 outer teeth, lateral margin basad of outer teeth not crenulate. Apex with robust, spur-like seta and a few smaller setae basally. Mesothoracic and metathoracic legs similar in shape, metathoracic legs slightly longer than mesothoracic legs. Longer tibial spur shorter than 2 basal tarsomeres in mesothoracic legs and as long as 2 tarsomeres in metathoracic legs. Parameres large, about 1.3 time shorter than phallobase, wide and rounded apically (in dorsal view), with small lateral teeth and large excavations between teeth and paramere apices (Fig. 12B). Internal sac of the aedeagus with 2 tooth-shaped sclerites (Fig. 12C). Female unknown. Distribution and habitat. Madecorphnus tuberculatus is known from one locality in the Betampona Reserve, a patch of primary low elevation rain forest about 25 km inland of the eastern coast (Fig. 12D).

REVIEW OF THE MADAGASCAN ORPHNINAE Zootaxa 4207 (1) © 2016 Magnolia Press · 29 FIGURE 12. Madecorphnus tuberculatus. Habitus in dorsal view (A), aedeagus in lateral view and parameres in dorsal view (B), internal sac of aedeagus (C), distributional records (D).

Madecorphnus fisheri Frolov, 2014 (Figs. 13A–E)

Madecorphnus fisheri Frolov, 2014: 35.

Type material examined. Holotype (Figs. 13A–D), male, “MADAGASCAR: Toliara Prov., Parc National d'Andohahela, Col du Sedro, 3.8 km 113° ESE Mahamavo, 37.6 km NNW Tolagnaro, 21–25 I 2002 / 24°45′50″S, 46°45′6″E coll. Fisher, Griswold et al. California Acad. of Sciences, sifted litter - montane rainforest elev 900m code: BLF5010 / CASENT 5504179 / Madecorphnus fisheri Frolov 2013 HOLOTYPUS” (CASC). Diagnosis. This species can be easily separated from other Madecorphnus species by its internal sac of the aedeagus without fields of spinules but with 1 long, straight sclerite, 1 shorter, folded sclerite, and 2 comma-shaped sclerites (Fig. 13D). Description. Male. Body length 6.0 mm. Color uniformly brown, disc of head and pronotum darker (Fig. 13A).

30 · Zootaxa 4207 (1) © 2016 Magnolia Press FROLOV ET AL. FIGURE 13. Madecorphnus fisheri, holotype. Habitus in dorsal view (A), aedeagus in lateral view and parameres in dorsal view (B), labels (C), internal sac of aedeagus (D), distributional records (E).

Right mandible slightly longer than left, without tooth behind apex. Labrum trapezoidal, with slightly rounded sides, length about 1/7 width (in dorsal view). Frontoclypeus slightly asymmetrical, apically obtuse, with 2 long and 4 shorter setae on the apical margin. Canthus and frontal suture indistinct. Frontoclypeus slightly depressed apicomedially. Head finely punctate with minute punctures separated by more than 4 times their diameter. Mentum without tubercles. Pronotum approximately 1.5 times wider than long, widest medially. Disc of pronotum convex, without any depressions, tubercles, or ridges. Punctation on pronotum finer than on head, almost indistinct. Margins with relatively wide border, lateral margins with 4 long setae: 1 seta on basal angle, 1 seta approximately in the middle of lateral margin, and 2 setae on the apical angle. Scutellum triangular, about 1/12 length of elytra. Elytra convex, with distinct humeral and apical, widest at basal third. First stria distinct and reaching the apex

REVIEW OF THE MADAGASCAN ORPHNINAE Zootaxa 4207 (1) © 2016 Magnolia Press · 31 of elytron, other striae indistinct. Disc of elytra sparsely punctate with relatively large punctures. Epipleura with long, sparse, brown setae. Base of elytron with border from scutellum to humeral callus. Protibiae with 3 outer teeth, lateral margin basad of outer teeth not crenulate. Apex with robust, spur-like seta and a few smaller setae basally. Mesothoracic and metathoracic legs similar in shape to each other. Longer tibial spur shorter than 2 basal tarsomeres in mesothoracic legs and as long as 2 tarsomeres in metathoracic legs. Parameres relatively wide in lateral view, with almost indistinct lateral teeth (Fig. 13B). Internal sac of the aedeagus with a characteristic long, almost straight sclerite, shorter folded sclerite, and 2 comma-shaped sclerites (Fig. 13D). Fields of spinules absent. Female unknown. Distribution and habitat. The type locality of M. fisheri is the southernmost locality of a Madecorphnus species to date. The specimen was collected in the mountain rainforest of the Andohahela National Park (Fig. 13E).

Madecorphnus pauliani Frolov, 2010 (Figs. 14A–F)

Madecorphnus pauliani Frolov, 2010c: 1102.

Type material examined. Holotype (Figs. 14A–E), male, “Madagascar Forêt Tanala [Fort-Carnot] Alluaud 1901 82 / Holotypus Madec. pauliani Frolov 2009 / MUSÉUM PARIS Coll. Ch. ALLUAUD” (MNHN). Diagnosis. This species differs from other Madecorphnus species in having 3 setae on the anterior angles of the pronotum and in distinct shape of the parameres. Description. Male. Body length 5.2 mm. Color dark brown, head and legs somewhat paler (Fig. 14A). Right mandible about 2 times longer than left, without tooth behind apex. Labrum trapezoidal, its length 1/4–1/ 5 width (in dorsal view). Frontoclypeus asymmetrical, slightly convex anteriorly, obtuse, with 2 long and a number of smaller setae. Canthus and frontal suture indistinct. Frontoclypeus somewhat depressed in the middle anteriorly, minutely punctate. Pronotum 1.8 times wider than long, widest medially. Margins with relatively wide border, lateral margins with 5 long setae (3 setae located on each anterior angle). Pronotum evenly convex and minutely punctate. Elytra convex, with distinct humeral and apical tubercules. Maximum width approximately at basal 1/3. First stria distinct and reaching the apex of elytron, other striae indistinct. Epipleura with long, sparse, brown setae. Base of elytra with border from scutellum to humeral callus. Elytra minutely punctate except for a few relatively large punctures in the middle of each elytron near stria 1. Protibiae with 3 outer teeth, lateral margin basad of outer teeth not crenulate. Apex with robust, spur-like seta and a few smaller setae basally. Mesothoracic and metathoracic legs similar in shape. Longer tibial spur as long as 2 basal tarsomeres in mesothoracic legs and as long as 3 basal tarsomeres in metathoracic legs. Parameres widely rounded apically, without lateral teeth (Fig. 14C). Internal sac with 2 large, feebly sclerotized sclerites (Figs. 14D–E). Female unknown. Distribution and habitat. Madecorphnus pauliani is known from one locality in southeastern Madagascar (Fig. 14F).

Madecorphnus niger Frolov, 2010 (Figs. 15A–F)

Madecorphnus niger Frolov, 2010c: 1103; Frolov 2013a: 2.

Type material examined. Holotype (Figs. 15A–C), male, “Madagascar Est Perinet A. Peyrieras X-1972 / Museum Paris / Holotypus Madec. niger Frolov 2009” (MNHN). Paratype, male: “Madagascar East Perinet A.Peyrieras XI-1973” (MNHN).

32 · Zootaxa 4207 (1) © 2016 Magnolia Press FROLOV ET AL. FIGURE 14. Madecorphnus pauliani, holotype. Habitus in dorsal view (A), labels (B), aedeagus in lateral view and parameres in dorsal view (C), internal sac of aedeagus (D—photograph, E—schematic representation of armature), distributional records (F).

Additional material examined. Male, “MADAGASCAR: Province d'Antananarivo 3 km 41° NE Andranomay, 11.5 km 147° SSE Anjozorobe 1300m 5–13 Dec 2000 18°28′24″S, 47°57′36″E coll. Fisher, Griswold et al. California Acad of Sciences ex rotten log in montane rainforest BLF2537” (CASC). Male, “MADAGASCAR: Toamasina Parc National de Zahamena, Onibe River elev 780 m 21–23 February 2009

REVIEW OF THE MADAGASCAN ORPHNINAE Zootaxa 4207 (1) © 2016 Magnolia Press · 33 17°45′33″S, 048°51′17″E Calif. Acad. of Sciences coll. B.L.Fisher et al. yellow pan trap rainforest coll code: BLF22170” (CASC).

FIGURE 15. Madecorphnus niger. Habitus in dorsal view (A), aedeagus in lateral view and parameres in dorsal view (B), internal sac of aedeagus (C—photograph, D—schematic representation of armature), distributional records (E), labels of holotype (F).

Diagnosis. This species can be separated from other Madecorphnus species by the shape of the parameres and from many species also by darker color of the body. Description. Male. Body length 5.5–5.7 mm. Color black, mouthparts and legs brown (Fig. 15A). Right mandible slightly longer than left, without tooth basad of apex. Labrum trapezoidal, length about 1/4–1/ 5 width (in dorsal view). Frontoclypeus asymmetrical, slightly convex anteriorly, obtuse, with 2 long and a number of smaller setae. Frontoclypeus slightly depressed in the middle anteriorly, minutely punctate (punctures separated by more than 10 their diameters). Pronotum 1.8 times wider than long, widest medially. Margins with relatively wide border, lateral margins with 4 long setae. Pronotum evenly convex, minutely punctate. Elytra convex, with distinct humeral and apical tubercules. Maximum width approximately at basal 1/3. First stria distinct and reaching the apex of elytron, other striae indistinct. Epipleura with long, sparse, brown setae. Base

34 · Zootaxa 4207 (1) © 2016 Magnolia Press FROLOV ET AL. of elytra with border from scutellum to humeral callus. Elytra minutely punctate except for a few relatively large punctures in the middle of each elytron near stria 1. Protibiae with 3 outer teeth, lateral margin basad of outer teeth not crenulate. Apex with robust, spur-like seta and a few smaller setae basally. Mesothoracic and metathoracic legs similar in shape. Longer tibial spur as long as 2 basal tarsomeres in mesothoracic legs and as long as 3 basal tarsomeres in metathoracic legs. Parameres curved and rounded in lateral view, without teeth but with concavities laterally (Fig. 15B). Endophallus armature consists of 3 similar highly-sclerotized, tooth-like sclerites (Figs. 15C, D). Female unknown. Variation. Except for body size, the examined specimens differ slightly in the length of the right mandible. Distribution and habitat. Madecorphnus niger is known from three localities in eastern Madagascar (Fig. 15E).

Madecorphnus punctatus Frolov, 2010 (Figs. 16A–G)

Madecorphnus punctatus Frolov, 2010c: 1104.

Type material examined. Holotype (Figs. 16A–F), male, “MADAGASCAR CENTRE Est du lac Mantasoa, frt Ambohiboatavo 1340 m, III.1973 P. Griveaud et A. Peyrieras / Holotypus Madec. punctatus Frolov 2009” (MNHN). Paratypes: two males with the same locality data as the holotype. Diagnosis. This species can be separated from other Madecorphnus species by the densely pubescent head and pronotum and by the shape of the parameres and internal sac armature. Description. Male. Body length 4.5–5.0 mm. Head and pronotum dark brown, elytra and legs brown (Fig. 16A). Right mandible as long as left, without tooth basad of apex. Labrum trapezoidal, length about 1/4–1/5 width (in dorsal view). Frontoclypeus symmetrical, slightly convex anteriorly, obtuse, with 2 long and a number of smaller setae. Frontoclypeus, slightly depressed in the middle anteriorly, rather densely, irregularly punctate (punctures separated by 1–3 puncture diameters) (Fig. 16B). Pronotum 1.8 times wider than long, widest medially. Margins with relatively wide border, lateral margins with 4 long setae. Pronotum evenly convex, densely, irregularly punctuate (punctures separated by 1–3 their diameters anteriorly, becoming sparser basally). Elytra convex, with distinct humeral and apical tubercules. Maximum width approximately at basal 1/3. First stria distinct and reaching the apex of elytron, other striae indistinct. Epipleura with long, sparse, brown setae. Base of elytra with border from scutellum to humeral callus. Elytra sparsely punctate with relatively large punctures (more densely near suture). Protibiae with 3 outer teeth, lateral margin basad of outer teeth not crenulate. Apex with robust, spur-like seta and a few smaller setae basally. Mesothoracic and metathoracic legs similar in shape. Longer tibial spur a bit shorter than 2 basal tarsomeres in mesothoracic legs and a bit shorter than 3 tarsomeres in metathoracic legs. Parameres tapering apically, somewhat arrow shaped, angulate laterally (Fig. 16C). Internal sac with 3 large sclerites and an area of short spinules (Figs. 16E, F). Female unknown. Variation. Except for body size variation, the examined specimens are similar. Distribution and habitat. Madecorphnus punctatus is known from one locality in central Madagascar (Fig. 16G).

Madecorphnus dentatus Frolov, 2010 (Figs. 17A–F)

Madecorphnus dentatus Frolov, 2010c: 1105.

REVIEW OF THE MADAGASCAN ORPHNINAE Zootaxa 4207 (1) © 2016 Magnolia Press · 35 FIGURE 16. Madecorphnus punctatus, holotype. Habitus (A) and head (B) in dorsal view, aedeagus in lateral view and parameres in dorsal view (C), labels (D), internal sac of aedeagus (E—photograph, F—schematic representation of armature), distributional records (G).

36 · Zootaxa 4207 (1) © 2016 Magnolia Press FROLOV ET AL. FIGURE 17. Madecorphnus dentatus, holotype. Habitus in dorsal view (A), aedeagus in lateral view and parameres in dorsal view (B), labels (C), internal sac of aedeagus (D—photograph, E—schematic representation of armature), distributional records (F).

Type material examined. Holotype (Figs. 17A–E), male, “Chaînes Anosyennes, S.O. du Trafonaomby plateau Andohahelo 1770–1950 m, V-1972 / MUSEUM PARIS Madagascar Est mission C.N.R.S. R.C.P. n°225 / Holotypus Madec. dentatus Frolov 2009” (MNHN).

REVIEW OF THE MADAGASCAN ORPHNINAE Zootaxa 4207 (1) © 2016 Magnolia Press · 37 Paratypes: two specimens, female with the same data as the holotype and male, “Madagascar Est : N.-O. de Fort-Dauphin, massif de l'Andohahelo, forêt d'Andranomangara, R.N.I. n 11, 1750 m, 20/25-I-1974 A. Peyrieras” (MNHN). Diagnosis. This species can be separated from other Madecorphnus species by the distinct shape of the parameres and internal sac armature. Description. Male. Body length 6.0 mm. Color reddish brown, head, legs, and apices of elytra somewhat darker. Right mandible slightly longer than left, without tooth behind of apex. Labrum trapezoidal, length about 1/4–1/ 5 width (in dorsal view). Frontoclypeus asymmetrical, slightly convex anteriorly, obtuse, with 2 long and a number of smaller setae. Canthus and frontal suture indistinct. Frontoclypeus somewhat depressed in the middle anteriorly, sparsely punctate. Pronotum 1.8 times wider than long, widest medially. Margins with relatively wide border, lateral margins with 4 long setae. Pronotum evenly convex, punctate (punctures separated by 2–3 puncture diameters). Elytra convex, with distinct humeral and apical tubercules. Maximum width approximately at basal 1/3. First stria distinct and reaching the apex of elytron, other striae indistinct. Disc of elytra with sparse punctures. Epipleura with long, sparse, brown setae. Base of elytra with border from scutellum to humeral callus. Elytra minutely punctate except for a few relatively large punctures in the middle of each elytron near stria 1. Protibiae with 3 outer teeth, lateral margin basad of outer teeth not crenulate. Apex with robust, spur-like seta and a few smaller setae basally. Mesothoracic and metathoracic legs similar in shape. Longer tibial spur as long as 2 basal tarsomeres in mesothoracic legs and as long as 3 basal tarsomeres in metathoracic legs. Aedeagus. Parameres symmetrical, tapering apically in lateral view, with large teeth laterally (Fig. 17B). Internal sac of aedeagus with 1 sclerite composed of 2 long slender spinules connected basally and with 3 small spinules (Fig. 17D–E). Female. Except for shorter mandibles, females differ from males in having 2 setae on hind angles of pronotum and in robust protibial spur. Variation. Male paratype has shorter mandibles similar to those in female. Distribution and habitat. Madecorphnus dentatus is known from two localities in southeastern Madagascar (Fig. 17F).

Madecorphnus saintemariensis Frolov, 2014 (Figs. 18A–D)

Madecorphnus saintemariensis Frolov, 2014: 36.

Type material examined. Holotype (Figs. 18A–C), male, "MADAGASCAR: Toamasina Prov., Ile Sainte Marie, Foret Ambohidena, 22.8 km 44°NW Ambodifotatra, el 20m 21 November 2005 / 16°49′28″S, 049°57′51″E California Acad. of Sciences coll. Brian L. Fisher et al. sifted litter, littoral rainforest coll. code: BLF12840” (CASC). Paratype: female with the same data as the holotype (CASC). Diagnosis. Madecorphnus saintemariensis differs from other species of the genus in having the internal sac of the aedeagus with 2 tooth-shaped sclerites, 1 longer sclerite, and a few fields of minute spinules, 1 of which forms a finger-shaped process of the sac (Fig. 18C); and in the parameres having large lateral teeth (Fig. 18B). Description. Male. Body length 6.9 mm. Color uniformly dark brown, pronotum and disc of head almost black, legs somewhat paler (Fig. 18A). Right mandible almost 1.5 times longer than left, without tooth behind apex. Labrum trapezoidal, with broadly rounded sides, length about 1/6 width (in dorsal view). Frontoclypeus asymmetrical, apically obtuse, with 4 setae on the apical margin, 3 are displaced to the left side. Canthus and frontoclypeus depressed apicomedially. Head finely punctate with minute punctures separated by 5–10 puncture diameters. Pronotum approximately 1.6 times wider than long, widest medially. Disc of pronotum convex, without any depressions, tubercles, or ridges. Punctation on pronotum is similar to that on head. Margins with relatively wide

38 · Zootaxa 4207 (1) © 2016 Magnolia Press FROLOV ET AL. border, lateral margins with 4 long setae: 1 seta on basal angle, 1 seta approximately in the middle of lateral margin, and 2 setae on the apical angle. Base with a row of longitudinally elongate punctures (smaller and sparser than in M. fisheri).

FIGURE 18. Madecorphnus saintemariensis, holotype. Habitus in dorsal view (A), aedeagus in lateral view and parameres in dorsal view (B), internal sac of aedeagus (C), distributional records (D).

Scutellum triangular, right angled apically, about 1/11 length of elytra. Elytra convex, with distinct humeral humps, widest at basal third, glabrous. First stria distinct and reaching the apex of elytron, other striae indistinct. Disc of elytra with double punctation consisting of large, sparse punctures and minute punctures. Epipleura with long, sparse, brown setae. Base of elytron with border from scutellum to humeral humps. Protibiae with 3 outer teeth, lateral margin basad of outer teeth not crenulate. Apex with robust, spur-like seta and a few smaller setae basally. Mesothoracic and metathoracic legs similar in shape. Longer tibial spur longer than first basal tarsomere in mesothoracic legs and as long as 2 tarsomeres in metathoracic legs.

REVIEW OF THE MADAGASCAN ORPHNINAE Zootaxa 4207 (1) © 2016 Magnolia Press · 39 Parameres relatively large, about 1.4 time shorter than phallobase, wide, with large lateral teeth (Fig. 18B). Internal sac of aedeagus with 2 tooth-shaped sclerites, 1 longer sclerite, and a few fields of minute spinules, 1 of which forms a finger-shaped process of the sac (Fig. 18C). Female. Females differ from males in having mandibles of subequal length, and apical spur of protibia. Distribution and habitat. Madecorphnus saintemariensis is the only Madecorphnus species found outside Madagascar mainland, and the only orphnine species found in Sainte Marie Island (Nosy Boraha), a small island 6 km off the eastern coast of Madagascar (Fig. 18D). Primary vegetation of the island, consisting of low altitude rainforest and coastal forest, is largely disturbed (Du Puy & Moat 1996). Madecorphnus saintemariensis is one of the few orphnine species inhabiting coastal forests in Madagascar.

Madecorphnus simplex Frolov, 2010 (Figs. 19A–F)

Madecorphnus simplex Frolov, 2010c: 1106, Frolov 2013a: 5.

Type material examined. Holotype (Figs. 19D, E), male, “MADAGASCAR CENTRE AMBATOFITORAHANA 1800 M F D H M / 31.XII.1972 / A. PEYRIERAS / Holotypus Madec. simplex Frolov 2009” (MNHN). Additional material examined. Male, “Madagascar, Fianarantsoa Prov., Foret d'Atsirakambiaty, 7.6 km 285° WNW Itremo, elev 1550 m., 22–26 January 2003 / 20°35′36″S, 046°33′48″E colls. Fisher Griswold et al. Calif. Acad. of Sci. pitfall trap—montane rainforest collection code BLF7150” (CASC); male, “MADAGASCAR: Province Fianarantsoa, Parc National Ranomafana, at broken bridge, elev 1110 m 19–26 March 2002 / 21°13′57″S, 47°22′19″E collector R. Harin'Hala California Acad. of Sciences, malaise trap in high altitude rainforest, MA-02- 09A-21” (CASC). Diagnosis. This species can be separated from other Madecorphnus species by the distinct shape of its parameres and the internal sac armature. Description. Male. Body length 5.5 mm. Color reddish brown (Fig. 19A). Right mandible 1/5 longer than left, without tooth behind apex. Labrum trapezoidal, length about 1/4–1/5 width (in dorsal view). Frontoclypeus asymmetrical, slightly convex anteriorly, obtuse, with 2 long and a number of smaller setae. Frontoclypeus slightly depressed in the middle anteriorly, sparsely punctate (punctures separated by about 5 puncture diameters). Pronotum 1.8 times wider than long, widest medially. Margins with relatively wide border, lateral margins with 4 long setae. Pronotum evenly convex, sparsely punctate. Elytra convex, with distinct humeral and apical tubercules. Maximum width approximately at basal 1/3. First stria distinct and reaching the apex of elytron, other striae indistinct. Epipleura with long, sparse, brown setae. Base of elytra with border from scutellum to humeral humps. Elytra sparsely punctate except for a few relatively large punctures. Protibiae with 3 outer teeth, lateral margin basad of outer teeth not crenulate. Apex with robust, spur-like seta and a few smaller setae basally. Mesothoracic and metathoracic legs similar in shape. Longer tibial spur as long as 2 basal tarsomeres in mesothoracic legs and shorter than 3 tarsomeres in metathoracic legs. Metafemora with a few pectinate setae on the inner side apically. Parameres rounded apically (in lateral view), with small teeth laterally (Fig. 19B). Internal sac with 1 sclerite composed of 2 long slender spinules connected basally (Figs. 19D–E). Female. Unknown. Distribution and habitat. Madecorphnus simplex is known from 3 localities in central and eastern Madagascar (Fig. 19F).

40 · Zootaxa 4207 (1) © 2016 Magnolia Press FROLOV ET AL. FIGURE 19. Madecorphnus simplex, holotype. Habitus in dorsal view (A), aedeagus in lateral view and parameres in dorsal view (B), labels (C), internal sac of aedeagus (D—photograph, E—schematic representation of armature), distributional records (F).

Madecorphnus aquilonius Frolov, 2012b: 162.

Type material examined. Holotype (Figs. 20A–E), male, “MADAGASCAR, Province Diego-Suarez, Sakalava Beach, dwarf littoral forest elev 10 m, 13–16 May 2001 / 12°15′46″S, 49°23′51″E R.Harin'Hala collector malaise trap — across sandy trail MA-01-04B-09 / CASENT 8014036 / HOLOTYPUS Madecorphnus aquilonius Frolov 2011” (CASC).

FIGURE 20. Madecorphnus aquilonius, holotype. Habitus in dorsal view (A), aedeagus in lateral view and parameres in dorsal view (B), labels (C), internal sac of aedeagus (D—photograph, E—schematic representation of armature), distributional records (F).

42 · Zootaxa 4207 (1) © 2016 Magnolia Press FROLOV ET AL. Diagnosis. Madecorphnus aquilonius can easily be separated from other Madecorphnus species by having characteristic internal sac armature consisting of 2 rather large sclerites: a spur-like sclerite and an asymmetrical, claw-shaped sclerite (Figs. 20D–E). Description. Male. Body length 5.8 mm. Color uniformly brown (Fig. 20A). Right mandible about 1/2 longer than left, without tooth behind apex. Labrum trapezoidal, with rounded sides, length about 1/6 width (in dorsal view). Frontoclypeus distinctly asymmetrical, apically obtuse, with 4 relatively long setae on the apical margin. Canthus and frontal suture indistinct. Frontoclypeus slightly depressed apicomedially. Head without traces of frontoclypeal suture, finely punctate with minute punctures separated by greater than 4 times their diameter. Pronotum approximately 1.75 times wider than long, widest medially. Disc of pronotum convex, without any depressions, tubercles, or ridges. Punctation on pronotum as fine as that on head. Margins with relatively wide border, lateral margins with 4 long setae: 1 seta on basal angle, 1 seta approximately in the middle of lateral margin, and 2 setae on the apical angle. Scutellum triangular, angulate apically, about 1/11 length of elytra. Elytra convex, with distinct humeral humps, widest at basal third. First stria distinct and reaching the apex of elytron, other striae indistinct. Disc of elytra sparsely punctate with relatively large punctures. Epipleura with long, sparse, brown setae. Base of elytron with border connected to first elytral stria. Protibiae with 3 outer teeth, lateral margin basad of outer teeth not crenulate. Apex with robust, spur-like seta and 2 smaller setae basally. Mesothoracic and metathoracic legs similar in shape to each other, metathoracic legs 1.4 times longer than mesothoracic legs. Longer tibial spur as long as 2 basal tarsomeres in mesothoracic legs and longer than 2 tarsomeres in metathoracic legs. Parameres with small teeth laterally, narrowly rounded apically in dorsal view and curved downwards in lateral view (Fig. 20B). Internal sac with 2 rather large sclerites: a spur-like sclerite and an asymmetrical, claw-shaped sclerite (Figs. 20D–E). Female. Unknown. Distribution and habitat. This species is known from one locality in the northernmost Madagascar in a sandy littoral forest of the Madagascar dry deciduous forest ecoregion (Fig. 20F).

Madecorphnus falcatus Paulian, 1992 (Figs. 21A–F)

Madecorphnus falcatus Paulian, 1992: 1971; Frolov 2010c: 1107, 2013a: 3.

Type material examined. Holotype (Figs. 21A–E), male, “MADAGASCAR ANTANAMBE 25/01/90 P.E.C. Foret PROG. MAG188-007 / Sissantobius falcatus n.sp. R. Paulian det. / HOLOTYPE” (MNHN). Diagnosis. This is the largest known species of Madecorphnus with the body length reaching 7 mm. Also, it differs from other Madecorphnus species in having a distinctive shape of the parameres (Fig. 21B) and internal sac armature consisting of 2 small, bifurcate sclerites, 1 long spur-like sclerite, and numerous small spinules (Figs. 21D–E). Description. Male. Body length 6.9–7.1 mm. Color dark brown (Fig. 21A). Right mandible 2 times longer than left, without tooth behind apex. Labrum trapezoidal, feebly visible in dorsal view. Frontoclypeus asymmetrical, slightly convex anteriorly, obtuse, with 2 long and a number of smaller setae. Frontoclypeus slightly depressed in the middle anteriorly, almost smooth. Pronotum 1.6 times wider than long, widest medially. Margins with relatively wide border, lateral margins with 4 long setae. Pronotum evenly convex, with minute punctures. Elytra convex, with distinct humeral and apical tubercules. Maximum width approximately at basal 1/3. First stria distinct and reaching the apex of elytron, other striae indistinct. Epipleura with long, sparse, brown setae. Base of elytra with border from scutellum to humeral humps. Elytra sparsely punctate with relatively large punctures and minute punctures between them. Protibiae with 3 outer teeth, lateral margin basad of outer teeth not crenulate. Apex with robust, spur-like seta

REVIEW OF THE MADAGASCAN ORPHNINAE Zootaxa 4207 (1) © 2016 Magnolia Press · 43 and a few smaller setae basally. Mesothoracic and metathoracic legs similar in shape. Longer tibial spur as long as 2 basal tarsomeres in mesothoracic legs and shorter than 3 basal tarsomeres in metathoracic legs. Metafemora with a few pectinate setae on the inner side apically.

FIGURE 21. Madecorphnus falcatus, holotype. Habitus in dorsal view (A), aedeagus in lateral view and parameres in dorsal and apical view (B), labels (C), internal sac of aedeagus (D—photograph, E—schematic representation of armature), distributional records (F).

44 · Zootaxa 4207 (1) © 2016 Magnolia Press FROLOV ET AL. Parameres relatively long, their apices curved downwards (in lateral view), with small teeth laterally visible in apical view (Fig. 21B). Internal sac with 1 long, spur-shaped sclerite, 2 smaller, double sclerites, and a field small adjacent spinules (Figs. 21D–E). Female. Unknown. Distribution and habitat. Madecorphnus falcatus is known from two localities in eastern Madagascar (Fig. 21F).

Madecorphnus falciger (Lansberge, 1886) (Figs. 22A–E)

Drepanognatus falciger Lansberge, 1886: 93. Sissantobius falciger (Lansberge): Paulian 1937: 140, 1977: 1203. Madecorphnus falciger (Lansberge): Frolov 2010c: 1107; Paulian 1992: 171.

FIGURE 22. Madecorphnus falciger, holotype. Habitus in dorsal view (A), aedeagus in lateral view and parameres in dorsal view (B), labels (C), internal sac of aedeagus (D—photograph, E—schematic representation of armature).

REVIEW OF THE MADAGASCAN ORPHNINAE Zootaxa 4207 (1) © 2016 Magnolia Press · 45 Type material examined. Holotype (Figs. 22A–E), male, “Sissantobius Rits. ? falciger Lansb. Madag.”, Ex- Musaeo VAN LANSBERGE / G.J.Arrow vidit 1908 / MUSEUM PARIS 1952 COLL. R. OBERTHÜR / Sissantobius falciger Lansb. HOLOTYPE” (MNHN). Diagnosis. This species differs from other Madecorphnus species in having a characteristic shape of the parameres (Fig. 22B) and armature of the internal sac of the aedeagus consisting of 2 large, semicircular sclerites; 1 short, spur-like sclerite; and numerous small spinules (Figs. 22D–E). Description. Male. Body length 6.0 mm. Color brown (Fig. 22A). Right mandible 2.5 longer than left, without tooth behind apex. Labrum trapezoidal, length about 1/5–1/6 width (in dorsal view). Frontoclypeus asymmetrical, slightly convex anteriorly, obtuse, with 2 long and a number of smaller setae. Canthus and frontal suture indistinct. Frontoclypeus slightly depressed in the middle anteriorly, punctate. Pronotum 2 times wider than long, widest medially. Margins with relatively wide border, lateral margins with 4 long setae. Pronotum evenly convex, sparsely punctate. Elytra convex, with distinct humeral and apical tubercules. Maximum width approximately at basal 1/3. First stria distinct and reaching the apex of elytron, other striae indistinct. Epipleura with long, sparse, brown setae. Base of elytra with border from scutellum to humeral humps. Elytra sparsely punctate with relatively large punctures. All legs of the holotype are broken with only the right profemur, left mesofemur, right mesofemur and mesotibia, and half of right metafemur remaining. Parameres angulate apically (in lateral view), with small but distinct teeth laterally (Fig. 22B). Internal sac with 2 large semicircular sclerites, 1 short spur-like sclerite and numerous small spinules (Figs. 22D–E). Female. Unknown. Distribution and habitat. The only known specimen of this species lacks any precise record of the locality where it was collected.

Madecorphnus hanskii Frolov, 2012 (Figs. 23A–F)

Madecorphnus hanskii Frolov, 2012b: 160.

Type material examined. Holotype (Figs. 23A–E), male, “Madagascar Ambohitantely [18°10' S, 47°17' E] 24.3.2005, wet forest, I. Hanski-group leg. / Holotypus Madec. hanskii Frolov 2009” (ZIN). Diagnosis. Madecorphnus hanskii is similar to M. falciger and M. punctatus in having internal sac of the aedeagus with 3 sclerites: 1 relatively long, spur-like, and 2 comma-shaped or somewhat bifurcate (Figs. 16E–F, 22D–E, 23D–E). It can be separated from these species by the shape and relative size of these sclerites and, from the latter, by the shape of parameres and sparse punctation of head and pronotum. Description. Male. Body length 5.3 mm. Color uniformly brown (Fig. 23A). Right mandible as long as left, without tooth behind apex. Labrum trapezoidal, with broadly rounded sides, length about 1/6 width (in dorsal view). Frontoclypeus slightly asymmetrical, apically obtuse, with 2 long and 6 shorter setae on the apical margin. Canthus and frontoclypeus almost flat, minutely depressed apicomedially. Head without traces of frontoclypeal suture, finely punctate with minute punctures separated by greater than 5 times their diameter. Pronotum approximately 1.6 times wider than long, widest medially. Disc of pronotum convex, without any depressions, tubercles, or ridges. Punctation on pronotum similar to that on head. Margins with relatively wide border, lateral margins with 4 long setae: 1 seta on basal angle, 1 seta approximately in the middle of lateral margin, and 2 setae on the apical angle. Scutellum triangular, right angled apically, about 1/13 length of elytra. Elytra convex, with distinct humeral humps, widest at basal third, glabrous. First stria distinct and reaching the apex of elytron, other striae indistinct. Disc of elytra with a few relatively large punctures. Epipleura with long, sparse, brown setae. Base of elytron with border. Wings fully developed. Protibiae with 3 outer teeth, lateral margin basad of outer teeth not crenulate. Protibial apex with robust, spur- like seta and 2 smaller setae basally. Mesothoracic and metathoracic legs similar in shape to each other,

46 · Zootaxa 4207 (1) © 2016 Magnolia Press FROLOV ET AL. metathoracic legs somewhat longer. Longer tibial spur shorter than 2 basal tarsomeres in mesothoracic legs and longer than 2 basal tarsomeres in metathoracic legs.

FIGURE 23. Madecorphnus hanskii, holotype. Habitus in dorsal view (A), aedeagus in lateral view and parameres in dorsal view (B), labels (C), internal sac of aedeagus (D—photograph, E—schematic representation of armature), distributional records (F).

Parameres with teeth laterally, narrowly rounded apically in dorsal view and curved downwards in lateral view (Fig. 23B). Internal sac with a long, spur-like sclerite and 2 smaller, somewhat comma-shaped sclerites (Figs. 23D– E). Female. Unknown. Distribution and habitat. The type specimen of M. hanskii was collected in wet forest in Ambohitantely

REVIEW OF THE MADAGASCAN ORPHNINAE Zootaxa 4207 (1) © 2016 Magnolia Press · 47 special reserve (Fig. 23F). The reserve is situated at altitudes of about 1500 m and houses the last remnants of indigenous forest on the central plateau of Madagascar. Madecorphnus hanskii is apparently a forest litter dweller and, due to severe deforestation of the central plateau, its current range may be limited to this small forest remnant.

Madecorphnus cuccodoroi Frolov, 2010 (Figs. 24A–F)

Madecorphnus cuccodoroi Frolov, 2010a: 64.

Type material examined. Holotype (Figs. 24A, C–E), male, “MADAGASCAR, Mt Ambre, 3 km NE N.P. camp site, 12°33'09''S, 49°09'40''E, 1300 m. 28.II.2003 #3a G. Cuccodoro / Holotypus Madec. cuccodoroi Frolov 2010” (MHNG). Paratypes: one male, “Madagascar, Mt Ambre, 2,5 km SE N.P. camp site, 12°31'12''S, 49°11'42''N, 1050 m. 11.III.2003, G. Cuccodoro” (MHNG); one male, “Madagascar, Mt Ambre, 1 km NW N.P. camp site, 12°31'55''S, 49°10'32''N, 1000 m. 2.III.2003, G. Cuccodoro” (MHNG); two males, “Madagascar, Mt Ambre, 3 km NE N.P. camp site, 12°32'30''S, 49°10'02''N, 1250 m. 15.III.2003, G. Cuccodoro” (MHNG, ZIN); one female, “Madagascar, Mt Ambre, 2,5 km SE N.P. camp site, 12°31'15''S, 49°11'40''N, 950 m. 03.III.2003, G. Cuccodoro” (MHNG); two females, “Madagascar, Mt Ambre, 0,5 km NE N.P. camp site, 12°31'50''S, 49°10'25''N, 1000 m. 01.III.2003, G. Cuccodoro” (MHNG, ZIN). Additional material examined. MADAGASCAR: Antsiranana: six males, “MADAGASCAR: Province d'Antsiranana, Parc National Montagne d'Ambre 3.6 km 235° SW Joffreville Elev 925 m 20–26 Jan 2001 / 12°32′4″S, 49°10′46″E coll. Fisher, Griswold et al. California Acad. of Sciences, sifted litter, montane rainforest, code: BLF2564” (CASC). Diagnosis. This species is externally similar to M. peyrierasi Frolov, 2010 and M. perinetensis Frolov, 2010 and can reliably be separated from them only by the shape of the parameres and, especially, the internal sac armature. In M. peyrierasi and M. perinetensis, the internal sac armature consists of 2 larger sclerites basally; 2 smaller, comma-shaped sclerites apically; and an area of numerous minute spinules situated between the sclerites (Figs. 26D–E, 27 D–E). In M. cuccodoroi, the 2 larger sclerites are probably homologous to those in the previous species, but they differ slightly in shape. The area composed of numerous minute spinules is larger in M. cuccodoroi, the spinules are somewhat coarser, and a part of the apical spinules is more robust and they are almost tooth shaped (Fig. 24C). Madecorphnus cuccodoroi also differs in the shape of the parameres, which are somewhat wider and more curved at apices (Figs. 24D, 26B, 27B). Description. Male. Body length 5.0–5.5 mm. Color uniformly dark brown, almost black, legs somewhat paler (Fig. 24A). Right mandible slightly longer than left, without tooth behind apex. Labrum trapezoidal, with slightly rounded sides, length about 1/6 width (in dorsal view). Frontoclypeus slightly asymmetrical, apically obtuse, with 2 long and 2 shorter setae on the apical margin. Frontoclypeus slightly depressed apicomedially. Head finely punctate with minute punctures separated by greater than 4 times their diameter. Pronotum approximately 1.5 times wider than long, widest medially. Disc of pronotum convex, without any depressions, tubercles, or ridges. Punctation on pronotum finer than on head, almost indistinct. Margins with relatively wide border, lateral margins with 4 long setae: 1 seta on basal angle, 1 seta approximately in the middle of lateral margin, and 2 setae on the apical angle. Scutellum triangular, angulate apically, about 1/12 length of elytra. Elytra convex, with distinct humeral and apical, widest at basal third. First stria distinct and reaching the apex of elytron, other striae indistinct. Disc of elytra sparsely punctate with relatively large punctures. Epipleura with long, sparse, brown setae. Base of elytron with border from scutellum to humeral callus. Protibiae with 3 outer teeth, lateral margin basad of outer teeth not crenulate. Apex of protibia with robust, spur-like seta and a few smaller setae basally. Mesothoracic and metathoracic legs similar in shape to each other. Longer tibial spur shorter than 2 basal tarsomeres in mesothoracic legs and as long as 2 tarsomeres in metathoracic legs. Parameres with small teeth laterally, narrowly rounded apically in dorsal view and curved downwards in lateral

48 · Zootaxa 4207 (1) © 2016 Magnolia Press FROLOV ET AL. view (Fig. 24D). Internal sac with 2 long sclerites and a large number of small spinules; some spinules more robust and form 2 indistinctly separated clusters in apical part (Fig. 24C). Female. Females (Fig. 24B) differ from males in having the apical spur of the protibiae that is slightly longer and more robust than apical seta in males. Variation. Body length of the examined specimens varies from 5.0–5.5 mm. Left and right mandibles of some males are almost equal in length and similar to those in females.

FIGURE 24. Madecorphnus cuccodoroi, holotype male (A, C, D, E), paratype female (B). Habitus in dorsal view, internal sac of aedeagus (C), aedeagus in lateral view and parameres in dorsal view (D), labels (E), distributional records (F).

REVIEW OF THE MADAGASCAN ORPHNINAE Zootaxa 4207 (1) © 2016 Magnolia Press · 49 Distribution and habitat. All specimens were collected in the Amber Mountain National Park in Antsiranana Province (northern Madagascar, Fig. 24F). The park occupies the Madagascan northernmost remnant of indigenous humid forest covering about 200 km2. All specimens were collected by sifting forest litter. The number of the collected specimens suggests that the population density of M. cuccodoroi is low at least during the season when the collecting was done.

Madecorphnus brunneus Frolov, 2010 (Figs. 25A–F)

Madecorphnus brunneus Frolov, 2010c: 1108.

Type material examined. Holotype (Figs. 25A–E), male, “ANDRINGITRA Est Anjavidilava 1850–1950 m 18- XII/15-I-1971 / Holotypus Madecorphnus brunneus Frolov 2009” (MNHN). Diagnosis. Madecorphnus brunneus is similar to M. perinetensis and M. peyrierasi but differs from them in having the parameres less acute in lateral view and in the different internal sac armature. Description. Male. Body length 6.2 mm. Color brown, pronotum and head dark brown (Fig. 25A). Right mandible about 2 times longer than left, without tooth basad of apex. Labrum trapezoidal, its length about 1/5 width (in dorsal view). Frontoclypeus asymmetrical, slightly convex anteriorly, obtuse, with 2 long setae. Frontoclypeus slightly depressed in the middle anteriorly, finely punctate. Pronotum 1.9 times wider than long, widest medially. Margins with relatively wide border, lateral margins with 4 long setae. Pronotum evenly convex, minutely punctate. Elytra convex, with distinct humeral and apical tubercules. Maximum width approximately at basal 1/3. First stria distinct and reaching the apex of elytron, other striae indistinct. Disc of elytra with sparse punctation. Epipleura with long, sparse, brown setae. Base of elytra with border from scutellum to humeral callus. Protibiae with 3 outer teeth, lateral margin basad of outer teeth not crenulate. Apex with robust, spur-like seta and a few smaller setae basally. Mesothoracic and metathoracic legs similar in shape. Longer tibial spur shorter than 2 basal tarsomeres in mesothoracic legs and as long as 2 tarsomeres in metathoracic legs. Parameres acutely rounded in lateral view (Fig. 25B). Internal sac of the aedeagus with 1 longer sclerite and 1 smaller, somewhat bifurcated sclerite (Figs. 25D–E). Female. Unknown. Distribution and habitat. Madecorphnus brunneus is known from one locality in the Adringitra Massif, southeastern Madagascar (Fig. 25F).

Madecorphnus perinetensis Frolov, 2010 (Figs. 26A–F)

Madecorphnus perinetensis Frolov, 2010c: 1108.

Type material examined. Holotype (Figs. 26A–E), male, “Madagascar East Perinet A.Peyrieras XI-1972 / Holotypus Madec. perinetensis Frolov 2009” (MNHN). Paratypes: one male and two females with the same data as the holotype (MNHN); one male “Madagascar East Perinet A.Peyrieras X-1972” (MNHN). Additional material examined. MADAGASCAR: Toamasina: male, “Madagascar Est: Analamazoatra 6- 11.I.2011 Ballerio leg.” (ABCB). Diagnosis. This species is similar to M. peyrierasi externally and in the shape of the internal sac armature, but can be separated from it by the different shape of the parameres and somewhat larger eyes (Figs. 26A–B). Description. Male. Body length 5.5–5.8 mm. Color uniformly brown (Fig. 26A). Right mandible about 2 times longer than left, without tooth behind apex. Labrum trapezoidal, length about 1/ 6 width (in dorsal view).

50 · Zootaxa 4207 (1) © 2016 Magnolia Press FROLOV ET AL. FIGURE 25. Madecorphnus brunneus, holotype. Habitus in dorsal view (A), aedeagus in lateral view and parameres in dorsal view (B), labels (C), internal sac of aedeagus (D—photograph, E—schematic representation of armature), distributional records (F).

Frontoclypeus asymmetrical, slightly convex anteriorly, obtuse, with 2 long and 2 shorter setae on the anterior margin. Frontoclypeus slightly depressed in the middle anteriorly, finely punctate. Pronotum 1.8 times wider than long, widest medially. Margins with relatively wide border, lateral margins with 4 long setae. Pronotum evenly convex, minutely punctate. Elytra convex, with distinct humeral and apical tubercules. Maximum width approximately at basal 1/3. First stria distinct and reaching the apex of elytron, other striae indistinct. Disc of elytra sparsely punctate with relatively

FIGURE 26. Madecorphnus perinetensis, holotype. Habitus in dorsal view (A), aedeagus in lateral view and parameres in dorsal view (B), labels (C), internal sac of aedeagus (D—photograph, E—schematic representation of armature), distributional records (F).

52 · Zootaxa 4207 (1) © 2016 Magnolia Press FROLOV ET AL. Protibiae with 3 outer teeth, lateral margin basad of outer teeth not crenulate. Apex with robust, spur-like seta and a few smaller setae basally. Mesothoracic and metathoracic legs similar in shape. Longer tibial spur shorter than 2 basal tarsomeres in mesothoracic legs and as long as 2 basal tarsomeres in metathoracic legs. Parameres less tapering apically than in the previous species, wider in dorsal view, and acute in lateral view. Internal sac with 2 longer and 2 smaller sclerites (Figs. 26D–E). Female. Females differ from males in having short subsymmetrical mandibles, long protibial spur, and wider elytra. Distribution and habitat. Madecorphnus perinetensis is known from a small area near Andasibe, eastern Madagascar, in primary rain forest habitat (Fig. 26F).

FIGURE 27. Madecorphnus peyrierasi, holotype. Habitus in dorsal view (A), aedeagus in lateral view and parameres in dorsal view (B), labels (C), internal sac of aedeagus (D—photograph, E—schematic representation of armature), distributional records (F).

Madecorphnus peyrierasi Frolov, 2010c: 1109.

Type material examined. Holotype (Figs. 27A–E), male, “MADAGASCAR EST PERINET XI.1972 A.PEYRIERAS / Holotypus Madec. peyrierasi Frolov 2009” (MNHN). Paratypes: one female with the same data as the holotype (MNHN); one female, “Madagascar East Perinet A.Peyrieras X-1972” (MNHN); one male, “La Mandraka A. Peyrieras 25-III-1975” (MNHN); one male, “Madagascar Est route d'Anosibe A.Peyrieras 6.IV.1975” (MNHN). Diagnosis. This species is similar to M. perinetensis externally and in the shape of the internal sac armature, but can be separated from it by the different shape of the parameres and somewhat smaller eyes (Figs. 27A–B). Description. Male. Body length 4.6–6.2 mm. Color uniformly brown, head slightly darker (Fig. 27A). Right mandible about 1.7–1.2 times longer than left, without tooth behind apex. Labrum trapezoidal, length about 1/6 width (in dorsal view). Frontoclypeus asymmetrical, slightly convex anteriorly, obtuse, with 2 long and 2 shorter setae on the anterior margin. Frontoclypeus slightly depressed in the middle anteriorly, finely punctate. Pronotum 1.8 times wider than long, widest medially. Margins with relatively wide border, lateral margins with 4 long setae. Pronotum evenly convex, minutely punctate. Elytra convex, with distinct humeral and apical tubercules. Maximum width approximately at basal 1/3. First stria distinct and reaching the apex of elytron, other striae indistinct. Disc of elytra with sparsely punctate with relatively large punctures. Epipleura with long, sparse, brown setae. Base of elytra with border from scutellum to humeral callus. Protibiae with 3 outer teeth, lateral margin basad of outer teeth not crenulate. Apex with robust, spur-like seta and a few smaller setae basally. Mesothoracic and metathoracic legs similar in shape. Longer tibial spur shorter than 2 basal tarsomeres in mesothoracic legs and as long as 2 tarsomeres in metathoracic legs. Parameres tapering apically, acute in lateral view (Fig. 27B). Internal sac with 2 longer and 2 smaller sclerites (Figs. 27D, E). Female. Females differ from males in having short subsymmetrical mandibles, long protibial spur, and wider elytra. Distribution and habitat. Madecorphnus peyrierasi is known from one locality in Adringitra Massif, southeastern Madagascar (Fig. 27F).

Genus Renorphnus Frolov & Montreuil, 2009

Renorphnus Frolov & Montreuil, 2009: 65. Type species. Orphnus clementi Petrovitz, by original designation.

Diagnosis. Renorphnus are small beetles (body length 7.0–9.5 mm), with uniform brown coloration. Mandibles are subsymmetrical. Frontoclypeus of both sexes is not tuberculate but with more or less raised anterior margin in the middle. Pronotum is wider than long, smooth, similar in both sexes of most examined specimens, with shallow, sometimes almost indistinct depression anteriorly and 2 small tubercles at each side of the depression. Propleurae are smooth, convex, without carinae separating anterolateral areas from basal area. Scutellum is triangular, narrowly rounded apically, about 1/12 length of elytra. Elytra convex with humeral humps, only first stria distinct. Wings fully developed. Metepisternon triangular, tapering caudally, with slightly widened anterodorsal angle slightly overlapping epypleuron. Protibiae of males with apical outer tooth directed almost parallel to the inner margin of tibia, while in female it is directed somewhat laterad. Phallobase strongly sclerotized dorsally and with a thin membrane ventrally. Parameres tapering apically in lateral view and with a minute tooth at apex, without setae. Internal sac of the aedeagus with an area composed of small spinules and with 2 characteristic semicircular sclerites. From the other Madagascar genera, Renorphnus can be separated by having larger eyes (width of eye in dorsal view about 1.8 times smaller than distance between eyes), disc of pronotum with 2 feebly developed tubercles in

Renorphnus clementi (Petrovitz, 1971) (Figs. 28A–E)

Orphnus clementi Petrovitz, 1971: 21; Paulian 1977: 1201. Renorphnus clementi (Petrovitz): Frolov & Montreuil 2009: 66.

Type material examined. Holotype, male, “Ivoloina (Tamatave) Madagask.lg.Clement” (MHNG). Additional material examined. MADAGASCAR: Antsiranana: two males, one female, “Madagascar, Province Diego-Suarez, Montaigne Francaise, elev 150 m, 6–20 March 2001 / 12 18′8″S, 49 38′51″E R.Harin'Hala coll. malaise along forested limestone ridge MA-01-06-07 / CASENT 8014178” (CASC); Toamasina: two females, “Ivoloina (Tamatave) Madagask.lg.Clement [15°34′00″S, 49°37′00″E, Clement leg.]” (MHNG); one female, “Andasibe protected area, Mar2004, wet forest, alt. 800 m, fish bated trap [18°55′48″S, 48°25′12″E], Ilkka Hanski leg” (ZIN); one female, “Ambila Mar2004 littoral forest, alt. 50 m, fish baited trap [18°49′48″S, 49°09′00″E], Ilkka Hanski leg.” (ZIN); one female, “MADAGASCAR: Toamasina Ambatovy, 12.4 km NE Moramanga elev 1080 m pitfall trap, 4–7 March 2007 18˚50'22"S 048˚18'30"E California Acad. of Sciences coll. B.L.Fisher et al. montane rainforest BLF16917” (CASC); Fianarantsoa: one female, “MADAGASCAR: Province Fianarantsoa, Manombo Special Reserve camp site 32 km SSE of Farafangana 5–15 December 2004 23°01.31'S, 47°43.20'E California Acad of Sciences coll: M. Irwin, R. Harin'Hala malaise trap, lowland rainforest, elev 36m MA-28-06”; one female, same data but collected 23.XI–5.XI.2004 (CASC); one female, same data but collected 27.II–13.III.2004 (CASC); one female, same data but collected 9–16.X.2005 (CASC); one female, same data but collected 15–27.II.2005 (CASC); one male and two females, same data but collected 26.III– 10.IV.2005 (CASC); one unsexed specimen, same data but collected 9–16 October 2005 (CASC); Mahajanga: one male, one female, “MADAGASCAR TAM, Morarano-Chrome, Foret 25 km W [Forest 25 km W of Morarano- Chrome, 17°44′54″S, 47°55′30″E], 13–25.IV.1991, A. Pauly col.” (MRAC); Toliara: one male, “Andahahelo 23.3.2004, wet forest, fish bated trap [24°45′36″S, 46°51′36″E], Ilkka Hanski leg.” (ZIN); two males, two females, “Andahahelo 26.4.2006, wet forest, fish bated trap [24°45′36″S, 46°51′36″E], Ilkka Hanski leg.” (ZIN); one male, “Madagascar, Beza-Mahafaly, Betioky 10.4.2006, alt. 140 m., semu-dry Eucal[ipt]. Forest fish bait.-pitfall trap. [23°09′00″S, 44°37′48″E] I. Hanski-group leg.” (ZIN); one male (UHHF), one male (MNHN), and one male (ZIN), “MADAGASCAR Ambatotsirongorongo [25°00′00″S, 46°45′00″E] Feb 2005 fish baited trap Ilkka Hanski leg.”; two males, two females (ZIN) and two males, two females (MNHN), “Madagascar, St. Luce, Manafiati 16.4.2006 alt. 0 m, hum. lit. for. fish bait.-pitfall trap. [humid littoral forest, fish baited pitfall trap, 24°46′12″S, 47°10′48″E] I. Hanski-group leg.”; 6 unsexed specimens, “MADAGASCAR: Toliara Prov., Parc National d'Andohahela, Col du Sedro, 3.8 km 113° ESE Mahamavo, 37.6 km NNW Tolagnaro, 21–25 I 2002 // sifted litter, montain rainforest, 900 m a.s.l., 24°45′50″S, 46°45′06″E, 21–25.I.2002, Fisher, Griswold et al. leg., (CASC); Madagascar (no precise locality): one female, “Madagascar Mus. Pragense” (NMPC). Description. Male. Small-sized beetle with elongate, strongly shiny body (Fig. 28A). Color brown, legs and apices of elytra somewhat paler. Frontoclypeus slightly convex anteriorly, obtusely rounded, anterior margin setose and crenulate in dorsal view. Eyes relatively large (diameter larger than the distance between eye and gula in ventral view), incompletely divided by canthus into smaller dorsal and larger ventral parts. Frontoclypeus not tuberculate, with more or less raised anterior margin in the middle; in some specimens it forms a flattened triangular tubercle (although it is never as robust and horn-like as in Triodontus and Pseudorphnus males). Dorsal surface of head impunctate. Labrum somewhat bilobate, slightly sinuate in the middle and relatively feebly protruding past frontoclypeus. Length in the middle about 1/5 width (in dorsal view). Pronotum 1.6 times wider than long, widest medially. Anterior margin with wide border, base with fine border. Lateral margins densely punctate, appearing crenulate in dorsal view. Disc of pronotum with shallow, sometimes almost indistinct depression anteriorly, with 2 small tubercles aside the depression. Surface of pronotum smooth, without punctures. Lateral margins with long, brown setae. Scutellum triangular, narrowly rounded apically, about 1/10 length of elytra.

REVIEW OF THE MADAGASCAN ORPHNINAE Zootaxa 4207 (1) © 2016 Magnolia Press · 55 FIGURE 28. Renorphnus clementi. Habitus in dorsal view (A—male, B—female), aedeagus in lateral view and parameres in dorsal view (C), internal sac of aedeagus (D), distributional records (E), protibia (F, male left, female right).

Elytra convex, with feebly marked humeral humps. Maximum width approximately at the middle. First stria distinct and reaching the apex of elytron, other striae feebly marked to indistinct. Disc of elytra with sparse punctures. Epipleura with long, sparse, brown setae. Base of elytra bordered. Protibiae with 3 outer teeth. Apical tooth directed almost parallel to inner margin of tibia. Lateral margin basad of outer teeth not crenulate. Apex and internal margin of tibia with robust, spur-like setae becoming slender towards base of tibia. Protarsi well developed, about 4/5 length of protibiae. Claws 1/2 length of apical tarsomere.

56 · Zootaxa 4207 (1) © 2016 Magnolia Press FROLOV ET AL. Apical protarsomere as long as tarsomeres 3 and 4 combined, as thick as other tarsomeres. Ventral surface of protibiae smooth, with 2 rows of setae along sides and sparse, longer setae in the middle. Ventral surface of femora smooth. Mesothoracic and metathoracic legs similar in shape; metafemora and metatibiae about 1/8 longer than mesofemora and mesotibiae. Tibiae somewhat triangular, with 2 apical spurs. Longer tibial spur as long as 2 basal tarsomeres. Claws 1/3 length of apical tarsomere. Femora almost impunctate, with 2 rows of long setae. Abdominal sternites with somewhat granular sculpture. Sternite 8 medially as long as sternites 4–5 combined. Pygidium transverse, irregularly punctate, hidden under elytra. Parameres symmetrical, tapering apically in lateral view and with a minute tooth at apex (Fig. 28C). Internal sac with an area composed of small spinules and with 2 characteristic semicircular sclerites (Fig. 28D). Female. In comparison to other Orphninae genera, the sexual dimorphism in R. clementi is weak. Females (Fig. 28B) are similar to males in shape of pronotum, although they generally have smaller medial tubercles. The female of R. clementi can be readily separated from the male by a character shared by all orphnines — the presence of a long, robust apical spur on protibia. Variation. The length of the examined males varies from 8.0–8.7 mm, females from 7.0–9.5 mm. Most of specimens of both sexes have distinct tubercles on disc of pronotum, although some specimens have feebler tubercles or only traces of them; two females from St. Luce have no tubercles, although the males from the same series have distinct tubercles. Distribution and habitat. Renorphnus clementi is widely distributed in Madagascar. The beetles were collected in wet forests in the eastern part of the island including littoral forests on the eastern coast and in relatively dry areas with secondary forest in the western part of Madagascar (Fig. 28E). In all cases where the data are available, the beetles were collected with pitfall traps baited with fish. Trapping setup and timing is described in more detail by Frolov and Montreuil (2006). Short-time exposures of the traps suggest that specimens of R. clementi were attracted to the carrion.

Genus Triodontus Westwood, 1845

Triodontus Westwood, 1845: 172; Paulian 1977: 1208. Triodentus (lapsus calami): Paulian 1937: 13. Orphnidius Benderitter, 1914b: 83; objective junior synonym. Type species: Orphnus nitidulus Guérin-Méneville, 1844, by monotypy.

Diagnosis. Triodontus are small to mid-sized (length 6.5–17.0 mm) beetles, with uniform brown to almost black coloration. Labrum bilobate, slightly sinuate in the middle and relatively feebly protruding past frontoclypeus; length in the middle is 1/6 width (in dorsal view). Mandibles subsymmetrical. Frontoclypeus with rounded anterior margin, with tubercle or horn in males; females have frontoclypeus without tubercle or with minute tubercle. Pronotum of larger males of most species with a bulge medially, large excavations aside the bulge, and with 2 ridge-shaped tubercles laterally of each excavation. Females with convex pronotum without excavations or tubercles. Propleurae smoothly convex, without carinae separating anterolateral areas from basal area. Prosternum with a distinct longitudinal ridge in the middle, slightly tapering caudally. Scutellum rounded apically, about 1/15 length of elytra. Elytra convex, smooth except for base, with marked striae in some species, with feebly marked humeral humps. Wings fully developed. Metepisternon triangular, tapering caudally, with slightly widened anterodorsal angle slightly overlapping epipleuron. Protibiae with 3 outer teeth, not crenulated basad of the teeth. Apices with 3 robust, spur-like setae and a few small setae. Protarsi about 3/4 length of protibiae. Claws 1/3 length of apical tarsomere. Apical protarsomere as long as tarsomeres 3 and 4 combined, as thick as other tarsomeres. Ventral surface of protibiae with a longitudinal keel. Mesothoracic and metathoracic legs similar in shape; metafemora and metatibiae slightly longer than mesothoracic legs. Tibiae somewhat triangular, with 2 apical spurs, with inner margin only slightly concave. Longer tibial spur as long as or slightly longer than 2 basal tarsomeres. Claws about 1/3 length of apical tarsomere. Ventral sides of femora almost impunctate, with sparse, long setae. Phallobase strongly sclerotized dorsally and with a thin membrane ventrally. Parameres relatively short, complex, consisting of outer and inner lobes, without setae. Internal sac of aedeagus with 1 characteristic symmetrical sclerite or a group of fused sclerites.

REVIEW OF THE MADAGASCAN ORPHNINAE Zootaxa 4207 (1) © 2016 Magnolia Press · 57 Diagnostic characters. The most reliable character to differentiate Triodontus species is the shape of the parameres. The shape of the prothoracic armature, especially when fully developed, is specific to species groups. Species composition, distribution, and habitat. Triodontus is endemic to Madagascar. It is the second most speciose orphnine genus of the island with 15 species. The species are distributed throughout the island but mostly in the eastern and northeastern parts. The majority of the localities agree well with the current distribution of the remnants of indigenous rain forests.

Key to Triodontus species (males)

1 Central pronotal bulge with 3 more-or-less similar tubercles (Figs. 31A–B); outer lobes of parameres rather large, mostly covering inner lobes (in lateral view: Fig. 31E) ...... 2 - Central pronotal bulge with 2 tubercles (Fig. 35A); outer lobes of parameres smaller, not covering inner lobes (in lateral view: Fig. 35 E) ...... 4 2 Body length less than 9.0 mm ...... 3 - Body length 9.5–12.0 mm ...... T. hova (Fairmaire) 3 Abdominal sternite 8 narrower and more sinuate medially, without oval transverse concavity (Fig. 31F); lateral lobes of parameres with small, keel-shaped processes on proximal parts (Fig. 31E)...... T. nitidulus (Guérin-Méneville) - Abdominal sternite 8 wider and less sinuate medially, with oval transverse concavity (Fig. 33G); lateral lobes of parameres without small, keel-shaped processes on proximal parts (Fig. 33F) ...... T. bicavatus (Fairmaire) 4 Abdominal sternite 8 with more-or-less developed tubercule medially (Fig. 34F) ...... 5 - Abdominal sternite 8 without tubercule medially (Fig. 36G) ...... ………. 6 5 Tubercules of abdominal sternite 8 rounded apically (Fig. 34F); lateral lobes of parameres with keel-shaped processes on proximal parts (in dorsal view, Fig. 34D)...... T. ankarafantsikae Frolov, Montreuil & Akhmetova - Tubercule of abdominal sternite 8 with concavity apically (Fig. 35G); lateral lobes of parameres with semicircular notches on proximal parts (in dorsal view, Fig. 35E) ...... T. lemoulti Frolov, Montreuil & Akhmetova 6 Body length less than 6.5 mm ...... T. modestus (Benderitter) - Body length 9.0–12.0 mm ...... 7 7 Outer lobes of parameres strongly sclerotized, with distinct lateral notches and teeth situated more-or-less apically (Fig. 37D) ...... 8 - Outer lobes of parameres less sclerotized, somewhat rolled up in most species; lateral notches, if present, situated more basally (Fig. 41E) ...... 10 8 Outer lobes of parameres slender in lateral and dorsal view (Fig. 37D)...... T. owas Westwood - Outer lobes of parameres wider, at lease in dorsal view (Figs. 39E, 40E) ...... 9 9 Outer lobes of parameres with slit-shaped lateral notches (Fig. 39E) ...... T. viettei Frolov, Montreuil & Akhmetova - Outer lobes of parameres with wide lateral notches (Fig. 40E) ...... T. hildebrandtii (Fairmaire) 10 Outer lobes of parameres long, curved downward and inward, with more-or-less separated strongly sclerotized lateral parts and membranous, overlapping medial parts; lateral notches absent (Fig. 41E) ...... T. alticola Paulian - Outer lobes of parameres of different shape, without distinctly separated lateral and medial, membranous parts; lateral notches absent or present ...... 11 11 Outer lobes of parameres without lateral notches (Fig. 42F) ...... T. fairmairei Frolov, Montreuil & Akhmetova - Outer lobes of parameres with more-or-less distinct lateral notches and teeth (Figs. 43F, 44D, 45E, 46D) ...... 12 12 Outer lobes of parameres with distinct, slit-shaped lateral notches (in lateral view, Figs. 43F, 44D) ...... 13 - Outer lobes of parameres less distinct, wider lateral notches (in lateral view, Figs. 45E, 46D)...... 14 13 Abdominal sternite 8 without fossa (Fig. 43E); slit-shaped lateral notches of outer lobes of parameres longer (Fig. 43F) ...... T. itremoi - Abdominal sternite 8 with shallow but distinct, somewhat transversal fossa (Fig. 44E); slit-shaped lateral notches of outer lobes of parameres shorter (Fig. 44D)...... T. copridoides Paulian 14 Outer lobes of parameres 1.3 times longer than inner lobes; inner lobes smaller (in lateral view) (Fig. 45E). . T. hanskii Frolov - Outer lobes of parameres as long as inner lobes; inner lobes larger (in lateral view, Fig. 46D) T. inexpectatus Frolov, Montreuil & Akhmetova

Triodontus hova (Fairmaire, 1868) (Figs. 29A–H; 30A–C)

Orphnus hova Fairmaire, 1868: 783; Fairmaire 1897: 100. Triodontus hova (Fairmaire): Paulian 1977: 1217. Orphnus distinctus Fairmaire, 1897: 100: junior objective synonym. Orphnus nigrita Brancsik, 1893: 224, new synonym Triodontus nigritus (Brancsik): Paulian 1937: 13, 1977: 1217.

58 · Zootaxa 4207 (1) © 2016 Magnolia Press FROLOV ET AL. FIGURE 29. Triodontus hova. Habitus in dorsal view (A, C—male [A—holotype], B—female), labels of holotype (D), habitus of male in dorso-lateral view (E), aedeagus in lateral view and parameres in dorsal and ventral view (F), internal sac of aedeagus (G), distributional records (H).

Type material examined. Orphnus hova: holotype (Figs. 29A, D), male, “MUSEUM PARIS MADAGASCAR Collection Leon Fairmaire 1906 / Orphnus distinctus hova [Fair] Madag. / TYPE” (MNHN).

REVIEW OF THE MADAGASCAN ORPHNINAE Zootaxa 4207 (1) © 2016 Magnolia Press · 59 Orphnus nigrita: syntype (Fig. 30B), female, “Fauna Ins. Nossibe / Coll. Dr. Ed. Knirsch. Orphnus nigrita Brancsik Determ. Dr. Knirsch / Co-Typus / CNHM 1955 Karl Brancsik Colln. ex Eduard Knirsch” (FMNH); syntype, female, “Coll. Dr. Ed. Knirsch. Orphnus nigrita Brancsik Determ. Dr. Knirsch / Typus / Typ/ CNHM 1955 Karl Brancsik Colln. ex Eduard Knirsch” (FMNH). Additional material examined. MADAGASCAR: Antsiranana: one male, “Sambirano N.O. Madagasc. [Sambirano, 13°43′00″S, 48°22′00″E]” (MNHN); one female, “Museum Paris Madagascar Nossi-Be H. Pierron 1885 [Nosy Be Island, 1885, H. Pierron leg.]” (MNHN); six males, six females, “Nossi-Be Madagascar” (MNHN); one male, “Madagascar 5415 / Nosy-Be VII 1983” (MNHN); Nosy Be Island, two females (NMPC); two females “Nossi-Be 1937 Cie. Sucriere” (IRSNB); four females, “Nossi-Be Hellville Tschitscherine [Nosy Be Island, Hellville, 13°24′00″S, 48°16′00″E, Tschitscherine leg.]” (MNHN); one male, “Madagascar Sambirano Nosy-Be foret de Lokobe XII-58 Andria Robinson [Nosy Be, Lokobe Forest, 13°22′59″S, 48°19′59″E, XII.1958, A. Robinson leg.]” (MNHN); one female, “MADAGASCAR: Province d'Antsiranana, Ampasindava, Foret d'Ambilanivy, 3.9 km 181° S Ambaliha [13°47′54″S, 49°09′42″E] elev. 600 m. 4–9 March 2001” (CASC); Mahajanga: one male, two females, “Madagascar Ouest: Antsalova, Antsingy. R.N. 9 I.1975 A. Peyrieras [Antsalova, Antsingy, "Reserve Naturelle 9", 19°06′00″S, 44°52′59″E]” (MNHN); one male, “Madagascar Siberbielle H.Perrier [Suberbieville = Maevatanana, 16°57′00″S, 46°49′59″E]” (MNHN); one male, “Madagascar Ile de Beraphia Cote Ouest 1934 [Beraphia Island, 14°01′59″S, 47°48′00″E, 1934]” (MNHN); one male, “Ankarafantsika Forest [16°09′00″S, 47°01′59″E] // MADAGASCAR I.1956 C.Koch leg.” (MHNG); Madagascar (no exact locality): one male (IRSNB). Diagnosis. Triodontus hova is similar to T. nitidulus and T. bicavatus in having 3 tubercles on the central pronotal bulge and relatively large outer lobes of parameres but differs from them in the larger overall size of the body, almost flat elytral intervals (as opposed to being somewhat convex in T. nitidulus and T. bicavatus), and in having the abdominal sternite 8 wider and less sinuate medially. Description. Male. Color of head, pronotum and elytra dark brown to brown; legs, antennae and underside of the body brown (Figs. 29A, C, E). Frontoclypeus convex anteriorly, rounded laterally, anterior margin crenulate and setose in dorsal view. Eyes relatively small (their diameter slightly smaller than the distance between eye and gula in ventral view), incompletely divided by canthus into small dorsal and large ventral parts. Frontoclypeus with small tubercles mediad of each eye and in some specimens with a horn in center of frontoclypeus; horn, when the most developed, is about as long as the width of the head, acutely rounded apically, straight to somewhat curved caudally at the apex, slightly rugose on posterior side. Pronotum with a bulge medially, with excavations aside the bulge, and with 2 ridge-shaped tubercles laterally of each excavation. Central bulge has 3 small but distinct tubercles. Lateral margins with wide border appearing somewhat crenulate in dorsal view, with a row of brown setae. Anterior margin with wide, smooth border. Posterior margin with fine border, not crenulate, punctate with narrow longitudinal punctures. Surface of pronotum smooth. Scutellum rounded apically, its visible part is about 1/12 length of elytra. Elytra convex, with distinct humeral humps, their maximum width is approximately at middle. Elytra with 10 fine striae on disc and with oblique line from base of 6th stria to approximately middle of 8th stria. Stria with small punctures. Intervals with minute punctures, almost smooth and flat. Base of elytra with irregular row of coarse punctures. Sternite 8 relatively wide, feebly sinuate medially (in ventral view), and somewhat excavated (Fig. 30A). Parameres (Fig. 29F) with large outer lobes covering inner lobes (in lateral view). Proximal parts of outer lobes with small keel-shaped processes. Internal sac of the aedeagus with 1 strongly sclerotized symmetrical, somewhat pentagonal sclerite, and with 2 similar protuberances of the membrane (Fig. 29G). Female. Females (Figs. 29B; 30B) differ from males in having long apical spur on protibiae, absence of head and pronotum armature including tubercles mediad of each eye, densely punctate dorsal side of head, and longer abdominal sternites including 6th sternite, which is evenly rounded apically. Variation. Pronotal and head armature of males is subject to allometric variability from fully developed (as described above) to almost undeveloped (with the males having small frontoclypeal tubercles in the middle and small fossa on the pronotum anteriorly. Body length of examined specimens varied from 9.5–12.0 mm (males) and from 9.0–14.0 mm (females).

Distribution. Triodontus hova occurs in western and northwestern Madagascar (Fig. 29H). Previously it was recorded mostly from Nosy Be Island (Paulian 1977), but it is also known from Nosy Berafia Island, Sambirano River basin, and from three distant localities in the western Madagascar up to Tsingy de Bemaraha Nature Reserve in the south. Remarks. Synonymy of the names Orphnus distinctus Fairmaire and O. hova Fairmaire was discussed by Paulian (1977). Brancsik (1893) described Orphnus nigrita based on two females from Nosy Be Island. He noted that they differed from T. hova in larger size (up to 14 mm) and darker, almost black coloration. Although most of T. hova specimens we examined are smaller (up to 12 mm) and more or less brown rather than black, we think that the type specimens of O. nigrita might be aberrant forms of T. hova or lie within the interspecific variability of this species. Other characters, but the size and color, agree well with the females of T. hova as well as the occurrence on Nosy Be. Therefore the new synonymy is here proposed.

Triodontus nitidulus (Guérin-Méneville, 1844) (Figs. 31A–G; 32A–C)

Orphnus nitidulus Guérin-Méneville, 1844: 86. Triodontus nitidulus (Guérin-Méneville): Westwood 1845: 173; Paulian 1937: 13, 1977: 1220. Triodontus occidentalis Paulian, 1977: 1220, new synonym Orphnus obsoletus Brancsik, 1893: 224, new synonym

Type material examined. Orphnus nitidulus: lectotype (here designated, Fig. 32A), male, “Type Guerin Men / Type / Coll. R. I. Sc. N. B. Madagascar Orphnus nitidulus Guer. [...] ♂ Madag / Coll. R. I. Sc. N. B. Madagascar ex coll. Guerin ex coll. Monchicourt ex coll. de Bonneuil Le Moult vendit / TYPE / Orphnus M.Leay Madagascarensis Dup. Madag” (IRSNB). Orphnus obsoletus: one syntype (Fig. 32C), male, “Nosy Be” (FMNH). Triodontus occidentalis: holotype (Fig. 32B), male, “Berati D. Ananalava P. Griveaud XII.1960 / MUSEUM PARIS Madagascar / Triodontus occidentalis n. sp. R. Paulian det. / TYPE”; paratypes, one male and four females with the same locality data as the holotype.

REVIEW OF THE MADAGASCAN ORPHNINAE Zootaxa 4207 (1) © 2016 Magnolia Press · 61 FIGURE 31. Triodontus nitidulus. Habitus in dorsal view (A—male, C—female), head and pronotum of male in dorsolateral view (B), internal sac of aedeagus (D), aedeagus in lateral view and parameres in dorsal and ventral view (E), abdomen (F), distributional records (G).

62 · Zootaxa 4207 (1) © 2016 Magnolia Press FROLOV ET AL. FIGURE 32. Triodontus nitidulus. Habitus of in dorsal view. Lectotype of T. nitidulus (A), holotype of T. occidentalis (B), syntype of T. obsoletus (C).

Additional material examined. MADAGASCAR: Antsiranana: three males, 15 females, “Madagascar-Est dct. Sambava Marojejy Ambinanitelo 500m XII-58 Raharizonina [Sambava District, Ambinanitelo]” (MNHN); six males, four females, “Madagascar Est 17.XIII.72 Marojejy 300 m. [Marojejy Range, 300 m, 17.XIII.1972]” (MNHN); two males, two females, “Andranovolo / Madagascar R-on Antalaha XII.38 Vadon [R-on Antalaha, Andranovolo, 15°01′59″S, 50°07′00″E]” (MNHN); one male, “Madagascar Est m-if du Marojejy env-ons Manantenina 150 m., II.1974 A.Peyrieras [Marojejy Range, environs of Manantenina, 14°28′59″S, 49°49′00″E]” (MNHN); 21 males, 35 females, “Madagascar Ron Antalaha [Antalaha District] XI.35 Vadon” (MNHN); one male, “Marojejy 400m f[or]et Ambatomitatao A.Peyrieras I.1973 [Ambatomitatao Forest, Marojejy Nature Reserve, 14°25′59″S, 49°43′59″E]” (MNHN); three males, “Madagascar Ron Antalaha XII.35 Vadon Andapa [Antalaha District, Andapa, 14°39′00″S, 49°39′00″E]” (MNHN); eight unsexed specimens, “Sambirano [13°43′00″S, 48°22′00″E] N.O. Madagasc.” (SMTFD); three females, “Madagascar Andapa 1.69 [Andapa, 14°39′00″S, 49°39′00″E]” (IRSNB); Toamasina: one male, “Reserve nat III Ambatovositra Andranomalaza I-57 P Soga [Ambatovositra, Andranomalaza, "Reserve naturelle III", 17°39′00″S, 48°37′00″E]” (MNHN); seven males, four females, “Madagascar Est Moramanga -57 Gruvel [Moramanga, 18°55′59″S, 48°12′00″E]” (MNHN); 42 males, 16 females, “Nandihizana / Madagascar R-ne Maroantsetra XII.1938 Vadon! [Maroantsetra District, Nandihizana, 15°45′00″S, 49°19′00″E]” (MNHN); one male, “Madagascar Beanana XII.45 Vadon! [Beanana, 15°43′59″S, 49°28′00″E]” (MNHN); six males, six females, “Museum Paris Madagascar Andevorante A. Mathiaux 1900 [Andevoranto, 18°57′00″S, 49°06′00″E, 1900, A. Mathiaux leg.]” (MNHN); 52 males, 16 females, “Madacascar Antsianaka et lac Alaotra 2e Trimestre 1889 Perrot Freres [Antsianaka and Alaotra Lake, 17°30′00″S, 48°30′00″E]” (MNHN); three males, five females, “Madagascar Fenerive E.Perrot [Fenerive, 17°22′00″S, 49°25′00″E]” (MNHN); two males, three females, “Madagascar Analamazoatra / Museum Paris 1931 Lasere” (MNHN); four males, two females, “Madacascar Antsianaka Perrot Freres 2 Semestre 1890 [Antsianaka, 17°30′00″S, 48°30′00″E]” (MNHN); one male, two females, “Madagascar-Est dct. Moramanga Fanovana X I-58 R.Vieu [Moramanga District, Fanovana, 18°55′00″S, 48°34′00″E]” (MNHN); one male, “Madagascar Est distr. Mananara - N. Antanambe Vadon et Peyrieras [Mananara District, Antanambe, 16°25′59″S, 49°51′00″E]” (MNHN); two females, “Museum Paris Madagascar Prov. de Fenerive Reg. de Soanierana A. Mathiaux 1905

REVIEW OF THE MADAGASCAN ORPHNINAE Zootaxa 4207 (1) © 2016 Magnolia Press · 63 [District of Soanierana (=Soanierana Ivongo)]” (MNHN); one female, “Madagascar Brickaville [18°49′00″S, 49°04′00″E] Cap. Refroigney[?].” (MNHN); one female, “Perinet [18°55′59″S, 48°25′00″E]” (MNHN); one male, “Museum Paris Madagascar Env. de Tamatave A. Raffrey 1884 [Tamatave (=Toamasina), 18°10′00″S, 49°22′59″E]” (MNHN); one male, “Madagascar Tamatave prov. Moramanga env. [Moramanga environs, 18°55′59″S, 48°12′00″E] 27–30.12.1996 Ivo. Jenis leg.” (MHNG); one male, two females, “Madagascar Tamatave prov. Andasibe (Maromizaha) [Forest Maromizaha, Anevoka, 18°57′00″S, 48°28′00″E] 19–20.12.1996 Ivo. Jenis leg.”; one male, “Madagascar Tamatave prov. Andasibe (Maromizaha) [Forest Maromizaha, Anevoka, 18°57′00″S, 48°28′00″E] 21–24.11.1995 Ivo. Jenis leg.” (MHNG); 14 males, five females, “Rogez Madagascar Mus Praha 800–990 m [Rogez, 800–990 m, 18°47′53″S, 48°36′05″E]” (NMPC); 71 unsexed specimens, “Madagascar C. Moramanga env. [Moramanga, 18°55′59″S, 48°12′00″E] 10–18.XII.1997 P.Pacholatko leg.” (NHMB); 41 unsexed specimens, “Tamatave [Toamasina, 18°10′00″S, 49°22′59″E], 1884, A. Raffrey leg., (SMTFD); one male, “Maroantsetra [15°25′59″S, 49°43′59″E] Madagasc.” (SMTFD); one male, “Moramanga [18°55′59″S, 48°12′00″E] Madagasc. or.” (SMTFD); one male, “mittl. O.-Madag. Ambalarond[r]a Dr.Denso 1931.11 [Ambalarondra, 18°28′06″S, 49°00′04″E, XI.1931, Denso leg.]” (SMTFD); eight males, two females, “Tamatavo [Tamatave (=Toamasina), 18°10′00″S, 49°22′59″E] Madagascar” (IRSNB); Antananarivo: eight males, four females, “Museum Paris Madagascar Antsirabe Waterlot 1916 [Antsirabe, 19°51′00″S, 47°01′59″E, 1916, Waterlot leg.]” (MNHN); one male, “Andrangoloaka 1600 m O.S.O. de Tananarive [Andrangoloaka, 1600 m, 19°01′00″S, 47°52′00″E]” (MNHN); three males, 10 females, “Madagascar Tanarive III.33 [Antananarivo, 18°55′00″S, 47°31′00″E] / Museum Paris 1933 J.Vadon et E.Lebis” (MNHN); two females, “Madagascar Est Ankadimanga Manjakandriana XII-57 Jean-Elie [Ankadimanga, Manjakandriana, 18°58′00″S, 47°45′00″E]” (MNHN); one male, one female, “Madagascar Ambatolampy [19°22′59″S, 47°25′00″E]” (MNHN); one female, “Madagascar Tananarive Vadon ! [Antananarive, 18°55′00″S, 47°31′00″E]” (MNHN); eight males, five females, “Madagascar La Mandraka Vadon! XII-36 [La Mandraka, 18°55′00″S, 47°55′59″E]” (MNHN); one male, “Tananarive III.33 [Antananarive, 18°55′00″S, 47°31′00″E]” (MNHN); one female, “Madagascar Antananarivo prov. Manankazo env. [18°09′00″S, 47°12′00″E] 15–17.12.1996 Ivo. Jenis leg.” (MHNG); 22 unsexed specimens, “La Mandraka [18°55′00″S, 47°55′59″E] II.1953 Madagascar” (NHMB); Fianarantsoa: eight males, 14 females, “Madagascar Region de Mananjary [21°13′00″S, 48°19′59″E] A. Mathiaux” (MNHN); two males, “Madagascar: Prov. Fianarantsoa, Ranomafana, 600 m. [Ranomafana, 600 m, 21°18′00″S, 47°30′00″E] 21 October 1988 W.E.Steiner & R. Van Epps collrs.” (MNHN); five females, “Madagascar Centre dct. Ambohimahasoa foret Tsarafidy 1450 m XII-59 & I.60 P.Griveaud [Ambohimahasoa Disctrict, Ankafina Tsarafidy, forest Tsarafidy, 1450 m, 21°12′00″S, 47°15′00″E]” (MNHN); two females, “Madagascar Vohilava 60 m. Faraony [Vohilava Faraony, 60 m, 21°46′00″S, 47°55′00″E]” (MNHN); four males, two females, “Ambositra [20°31′00″S, 47°15′00″E] Madagascar” (MNHN); six males, six females, “Madagascar Fianarantsoa prov. Ranomafana env. [21°18′00″S, 47°30′00″E] 29.11.– 2.12.1995 Ivo. Jenis leg.” (MHNG); one female, “MADAGASCAR - CE 2010 Ranomafana NP ~ 1120 m Vohiparara env., at light on river bank [Ranomafana National Park, Vohiparara environs, ca. 1120 m, 21°13′35″S, 47°22′11″E]; 17.XI. local coll. lgt., BMNH(E) 2010-29” (BMNH); two unsexed specimens, “Ambositra Madagascar 14.11.55 G.Minet [Ambositra , 20°31′00″S, 47°15′00″E]” (NHMB); one male, “MADAGASCAR, Fianarantsoa Province, Ranomafana National Park, Vohiparara area, 1170 m, mixed tropical forest. 2–22 January 2001, 21.22644°S / 47.36979°E. Stop# DHK-01-004 D.H. & K.M. Kavanaugh. R.L. Brett, E. Elsom, F. Vargas, R. Ranaivosolo, E.F. Randrianirina, N. Rasoamananana, T.J. Ravelomanana, and H.C. Raveloson leg.” (CASC); one female, “MADAGASCAR, Fianarantsoa Province, Ranomafana National Park, Talatakely area, 900 m, mixed tropical forest 2–22 January 2001, 21.25041°S / 47.41945°E, COL-DHK-2001-001 D.H. & K.M. Kavanaugh. R.L. Brett, E. Elsom, F. Vargas, R. Ranaivosolo, E.F. Randrianirina, N. Rasoamananana, T.J. Ravelomanana, and H.C. Raveloson leg.“ (CASC); one male, “MADAGASCAR, 2007, Ranomafana Nat. Park near Ranomafana vill. [21°18′00″S, 47°30′00″E], M. Tryzna leg., 26–31.i., BMNH(E) 2010-29 [26–31.I.2007, M. Tryzna leg.]” (BMNH); three males, three females, “MADAGASCAR CE 2011, Ranomafana N. P. [Ranomafana National Park] 17–21.xi. S21°15'22.6'' E47°25'17.8'' 958 m; at light; M. Tryzna leg. BMNH(E) 2010-29” (BMNH); one male, “Deans Cowan Betsileo Madagascar / 1881 3000 ft 4000 ft [3000–4000 ft, 1881]” (SMTFD); one male, “Ifanadiana [21°18′12″S, 47°38′19″E] Madagascar Mus. Praha” (NMPC); one male, three females, “Masagascar Midongy [Midongy-du-Sud, 23°35′13″S, 47°01′00″E]” (NMPC); Mahajanga: 57 males, 14 females, “Ste Marie de Madagascar Perrot Freres X–XII 1896 [Sainte Marie de Madagascar, near Marovoay, 16°06′00″S, 46°37′59″E]” (MNHN); one male, one female, “Berati D. Ananalava P. Griveaud XII.1960 [Ananalava District, Beraty,

64 · Zootaxa 4207 (1) © 2016 Magnolia Press FROLOV ET AL. 14°01′00″S, 48°15′00″E]” (MNHN); one female, “Randiantsileo / Madagascar Ron Mandritsara XII-37 Vadon ! [Randiantsileo, Mandritsara District]” (MNHN); three females, “Morafenobe [unclear locality] Madag. occ.” (SMTFD); one male, “Maromandia [14°13′00″S, 48°04′59″E] Masagascar Mus. Praha” (NMPC); 79 males, 20 females, “Coll. IRScNB MADAGASCAR Sainte Marie de Madagascar [16°06′00″S, 46°37′59″E] 1897 Ex. Coll. Oberthur” (IRSNB); Toliara: one male, one female, “Madagascar Ouest Bongolava 1300 m. [Bongolava, 18°30′00″S, 45°30′00″E] / A.Peyrieras XII.1974” (MNHN); one female, “Betroka XI.31 [Betroka, 23°16′00″S, 46°04′59″E, XI.1931]” (MNHN); Madagascar (no precise locality): two females, “Madagascar E.Perrot” (MNHN); six unsexed specimens, “Madagascar” (SMTFD). Diagnosis. Triodontus nitidulus is most similar to T. bicavatus. Males of the former species can be separated by having abdominal sternite 8 narrower and more sinuate medially, without oval transverse concavity (Fig. 31F), and by having the lateral lobes of the parameres without small keel-shaped processes on proximal parts (Fig. 31E). Females of these species are difficult to separate. Description. Male. Color of head, pronotum and elytra dark brown; legs, antennae, and underside of the body brown (Fig. 31A). Frontoclypeus convex anteriorly, rounded laterally, anterior margin crenulate and setose in dorsal view. Eyes relatively small (diameter slightly smaller than the distance between eye and gula in ventral view), incompletely divided by canthus into small, dorsal and large, ventral parts. Frontoclypeus with small tubercles mediad of each eye and in some specimens with a horn in the center of the frontoclypeus. The horn, when the most developed, is about as long as the width of the head, acutely rounded apically, straight to somewhat curved caudally at the apex, slightly rugose on posterior side. Pronotum with a bulge medially, with excavations aside the bulge, and with 2 ridge-shaped tubercles laterally of each excavation. The median bulge has 3 small but distinct tubercles. Lateral margins with wide border appearing somewhat crenulate in dorsal view, with a row of brown setae. Anterior margin with wide, smooth border. Posterior margin with fine border, not crenulate, punctate with narrow longitudinal punctures. Surface of most pronotum smooth, anterior and posterior angles with sparse and coarse punctation. Scutellum rounded apically, visible part about 1/12 length of elytra. Elytra convex, with distinct humeral humps, their maximum width is approximately at the middle. Elytra with 10 distinct striae on disc and with oblique line from base of 6th stria to approximately middle of 8th stria. Stria with small punctures. Intervals with minute punctures, almost smooth. Base of elytra with an irregular row of coarse punctures. Sternite 8 medially deeply sinuate (in ventral view) and somewhat excavated (Fig. 31F). Parameres (Fig. 31E) with large outer lobes covering inner lobes (in lateral view). Proximal parts of outer lobes without keel-shaped processes. Internal sac of the aedeagus with 1 strongly sclerotized symmetrical sclerite, somewhat horseshoe shaped, and with 2 similar protuberances of the membrane (Fig. 31D). Female. Females (Fig. 31C) differ from males in having a long, apical spur on the protibiae; absence of the head and pronotum armature including tubercles mediad of each eye; densely punctate dorsal side of the head; and longer abdominal sternites including 6th sternite, which is evenly rounded apically. Variation. Pronotal and head armature of males is subject to allometric variability from fully developed (as described above) to almost undeveloped (with the males having small frontoclypeal tubercles in the middle and small fossa on the pronotum anteriorly. Body length of examined specimens varied from 6.5–9.0 mm (males) and from 6.0–7.5 mm (females). Remarks. The lectotype of T. nitidulus was unknown to Paulian (1977). We found one male specimen originated from the collection of Guérin-Méneville and deposited now in IRSNB. This specimen agrees with the original description of the species. Since Guérin-Méneville mentioned both sexes, evidently he examined more than one specimen although he provided no exact data on this account. To ensure the stability of the nomenclature, we designate this specimen (Fig. 32A) the lectotype of Orphnus nitidulus Guérin-Méneville, 1844. Brancsik (1893), in the original description of O. obsoletus, noted that it was similar to O. nitidulus (= T. nitidulus). From the short diagnosis of the former, it can be inferred that it differs from the later mostly in the less developed head and pronotum armature. Paulian (1977) considered O. obsoletus a junior synonym of T. bicavatus, however he did not discuss this synonymy. Examination of the male syntype of O. obsoletus showed that it has all the diagnostic characters of T. nitidulus, including the characteristic shape of the abdominal sternite 8, therefore the new synonymy is here proposed.

REVIEW OF THE MADAGASCAN ORPHNINAE Zootaxa 4207 (1) © 2016 Magnolia Press · 65 Paulian (1977) distinguished T. occidentalis Paulian from T. nitidulus by the absence of the fossette on the abdominal sternite 8 in the males of the former (as opposed to the presence of a “vague fossette” in the later). However T. nitidulus has no distinct fossette or fovea on the abdominal sternite 8 (as in T. bicavatus) but rather its abdominal sternite 8 is apically sinuate and somewhat excavated, that may appear “foveate” depending on the angle of view. We found no noticeable differences between the type of T. occidentalis and available material of T. nitidulus and hence we propose the new synonymy. Distribution and habitat. This is the most widespread and the only relatively abundant species of the Orphninae in Madagascar. It is currently known from 50 localities throughout the island (Fig. 31G). Most of the localities are in the eastern Madagascar, in the area occupied by primary and secondary forests. The relatively wide distribution and high abundance of this species may be, at least partly, a result of the secondary adaptation to feeding on cultivated crops. Larvae of T. nitidulus were collected in upland rice fields in the Central Plateau (Randriamanantsoa et al. 2010).

Triodontus bicavatus (Fairmaire, 1905) (Figs. 33A–H)

Orphnus bicavatus Fairmaire, 1905: 118. Triodontus bicavatus (Fairmaire): Paulian 1977: 1221. Triodontus obsoletus Brancsick: synonymy by Paulian 1977.

Type material examined. Holotype (Figs. 33A, D), male, “MUSÉUM PARIS 1906 Coll. Leon Fairmaire / Orphnus bicavatus opm n. sp. XXX / TYPE” (MNHN) Additional material examined. MADAGASCAR. Antsiranana: one male, “Mt. d'Ambre [Amber Mountains, 12°31′50″S, 49°10′24″E] / Museum Paris 1934 Brouhard” (MNHN); one male, “Mad. Vohemar, Ampanefena XII.36 Vadon [Vohemar, Ampanefena, 13°52′00″S, 49°58′00″E]” (MNHN); one male, “Diego Suarez [12°16′00″S, 49°16′59″E] / Museum Paris 1936 Coll. A. Boucomont” (MNHN); four males, five females, “Andapa / Madagascar Ron Antalaha XII.37 Vadon [Antalaha District, Andapa, 14°39′00″S, 49°39′00″E” (MNHN); four males, nine females, “Madagascar-Est dct. Sambava R.N.XII-Marojejy Ambatosoratra 1700m XI-60 P.Soga [Ambatosoratra, Natural Reserve of Marojejy (RNI XII), 1700 m, 14°25′59″S, 49°43′59″E]” (MNHN); seven males, 10 females, “Madagascar Ron Antalaha XII.35 Vadon Andapa [Antalaha District, Andapa, 14°39′00″S, 49°39′00″E]” (MNHN); 10 males, 32 females, “Mt. d'Ambre [Amber Mountains] Madagascar” (MNHN); one male, “Madagascar Nord Montagne D'Ambre Foret de Sakaramy 28.XII.2005 A.Ballerio” (ABCB); Mahajanga: three males, “Anginjanbary [dubious locality] / Madagascar Ron Mandritsara XII-37 Vadon !” (MNHN). Diagnosis. Triodontus bicavatus is most similar to T. nitidulus. Males of the former species can be separated by having abdominal sternite 8 rather wide, not sinuate medially, and with oval transverse concavity, and also by having the lateral lobes of the parameres with small keel-shaped processes on proximal parts. Females of these species are difficult to separate. Description. Male. Color of head, pronotum, and elytra dark brown; legs, antennae, and underside of the body brown (Figs. 33A–B). Frontoclypeus convex anteriorly, rounded laterally, anterior margin crenulate and setose in dorsal view. Eyes relatively small (diameter slightly smaller than the distance between eye and gula in ventral view), incompletely divided by canthus into small, dorsal and large, ventral parts. Frontoclypeus with small tubercles mediad of each eye and in some specimens with a horn in the center of the frontoclypeus. The horn, when the most developed, is about as long as the width of the head, acutely rounded apically, straight to somewhat curved caudally at the apex, slightly rugose on posterior side. Pronotum with a bulge medially, with excavations aside the bulge, and with 2 ridge-shaped tubercles laterally of each excavation. The median bulge has 3 small but distinct tubercles. Lateral margins with wide border appearing somewhat crenulate in dorsal view, with a row of brown setae. Anterior margin with wide, smooth border. Posterior margin with fine border, not crenulate, punctate with narrow longitudinal punctures. Surface of most pronotum smooth, anterior and posterior angles with sparse and coarse punctation. Scutellum rounded apically, its visible part is about 1/12 length of elytra.

66 · Zootaxa 4207 (1) © 2016 Magnolia Press FROLOV ET AL. FIGURE 33. Triodontus bicavatus. Habitus in dorsal view (A, B—male [A—holotype], C—female), labels of holotype (D), internal sac of aedeagus (E), aedeagus in lateral view and parameres in dorsal and ventral view (F), abdomen of holotype (G), distributional records (H).

REVIEW OF THE MADAGASCAN ORPHNINAE Zootaxa 4207 (1) © 2016 Magnolia Press · 67 Elytra convex, with distinct humeral humps, their maximum width is approximately at the middle. Elytra with 10 distinct striae on disc and with oblique line from base of 6th stria to approximately middle of 8th stria. Stria with small punctures. Intervals with minute punctures, almost smooth. Base of elytra with an irregular row of coarse punctures. Sternite 8 medially feebly sinuate (in ventral view), with elongate transversal fovea (Fig. 33G). Parameres (Fig. 33F) with large outer lobes covering inner lobes (in lateral view). Proximal parts of outer lobes with small keel-shaped processes. Internal sac of the aedeagus with 1 strongly-sclerotized, somewhat horseshoe- shaped, symmetrical sclerite and with 2 similar protuberances of the membrane (Fig. 33E). Female. Females (Fig. 33C) differ from males in having long apical spur on protibiae, absence of head and pronotum armature including tubercles mediad of each eye, densely punctate dorsal side of head, and longer abdominal sternites including 6th sternite, which is evenly rounded apically. Variation. The variation is similar to that of T. nitidulus: pronotal and head armature of males is subject to allometric variability from fully developed (as described above) to almost undeveloped (with the males having small frontoclypeal tubercles in the middle and small fossa on the pronotum anteriorly). Body length of examined specimens varied from 6.5–8.2 mm (males) and from 6.5–7.5 mm (females). Remarks. Paulian (1977) considered O. obsoletus a junior synonym of T. bicavatus, however the type specimen of the former species has all the diagnostic characters of T. nitidulus (see above). Distribution. T. bicavatus is known from a few localities in northern Madagascar including the northernmost part of the island (Fig. 33H).

Triodontus ankarafantsikae Frolov, Montreuil & Akhmetova, new species (Figs. 34A–G)

Type material. Holotype (Figs. 34A–F), male, “ANKARAFANTSIKA AMPIJOROA 1.10-11-1973/ MUSEUM PARIS MADAGASCAR OUEST / Triodontus ankarafantsikae HOLOTYPUS Frolov et. al. 2010” (MNHN). Diagnosis. Triodontus ankarafantsikae is similar to T. lemoulti new species in having the abdominal sternite 8 with a more or less developed tubercule medially, but differs from it in having the tubercule rounded apically (as opposed to being concave in T. lemoulti) and lateral lobes of parameres with keel-shaped processes on proximal parts (in dorsal view, Fig. 34D). Description. Holotype, male. Body length 10.6 mm. Body strongly shiny (Figs. 34A–B). Color of head, pronotum and elytra dark brown; legs, antennae, and underside of the body light brown. Frontoclypeus convex anteriorly, rounded laterally, anterior margin slightly crenulate and setose in dorsal view. Eyes relatively small (diameter slightly smaller than the distance between eye and gula in ventral view), incompletely divided by canthus into small dorsal and large ventral parts. Frontoclypeus with small tubercles mediad of each eye and with a long horn in center of frontoclypeus; horn is longer than width of head, acutely rounded apically, somewhat curved caudally, slightly rugose on posterior side. Pronotum convex, almost smooth, feebly excavated anteriorly, with a medial bulge divided into 2 separated tubercles and 2 tubercles laterad of the bulge. Anterior margin with wide, smooth border, feebly sinuate medially. Posterior margin with fine border, not crenulate, punctate with small, narrow punctures. Scutellum rounded apically, its visible part about 1/15 length of elytra. Elytra convex, with distinct humeral humps. Maximum width approximately at the middle. Elytra with 10 feebly visible striae and with oblique line from base of 6th stria to approximately middle of 8th stria. Stria with small, indistinct punctures. Intervals with minute punctures, almost smooth. Base of elytra with an irregular row of coarse punctures each bearing small, yellow seta. Abdominal sternite 8 with a developed tubercule medially, tubercule is rounded apically (Fig. 34F). Parameres have outer lobes with keel-shaped processes on proximal parts (in dorsal view, Fig. 34D) and long feebly sclerotized apices. Internal sac of the aedeagus with 1 strongly sclerotized somewhat pentagonal sclerite, and 2 additional similar sclerites (Fig. 34E). Female. Unknown. Distribution. The species is known from one locality in western Madagascar, from dry tropical forest area (Fig. 34G).

68 · Zootaxa 4207 (1) © 2016 Magnolia Press FROLOV ET AL. Etymology. The species name is derived from the name of the type locality, Ankarafantsika. This name should be treated as a noun in apposition.

FIGURE 34. Triodontus ankarafantsikae, holotype male. Habitus in dorsal (A) and dorsolateral view (B), labels (C), aedeagus in lateral view and parameres in dorsal and ventral view (D), internal sac of aedeagus (E), abdomen (F), distributional records (G).

Triodontus nigritus: Paulian, 1977: 1217, incorrect identification.

Type material. Holotype (Figs. 35A–B, D—F), male, “MADAGASCAR Plantations du Sambirano Collection Le Moult / MUSÉUM PARIS 1938 COLL. A. BOUCOMONT / Triodontus nigritus Brancs. R. Paulian det. / HOLOTYPUS Triodontus lemoulti Frolov et al. 2010” (MNHN). Paratypes: three specimens with the same locality data as the holotype, one male (MHNG), one male (IRSNB), one male (MNHN); one male and one female, “Sambirano [Sambirano River] N.O. Madagasc.” (MHNG); one male and one female, “Mandritsara [Mahajanga: Randiantsileo, Mandritsara District] XII-37 Vadon!” (MNHN); one male and two females, “C. Katsepy (Majunga) [15°46′00″S, 46°13′59″E] 24–31.XII.1997 P.Pacholatko leg.” (NHMB); four males, “MADAGASCAR: Mahajanga Prov., Parc National Tsingy de Bamaraha, 3.4 km 93° E Bekopaka, Tombeau Vazimba elev. 50m, 6–10 Nov 2001 / Parc National Tsingy de Bamaraha, 3.4 km 93° E Bekopaka, Tombeau Vazimba, 50 m a.s.l., pitfall trap in tropical dry forest [19°08′31″S, 44°49′41″E] / 19°8′31″S, 44°49′41″E coll. Fisher, Griswold et al. California Acad. of Sciences, pitfall trap in tropical dry forest, code:BLF4230” (CASC); one male, “Madagascar Sahapetraka, 10.12.2006, cow dung pasture, Hanski group leg. / (GPS 855) Elev. 1288m, S19 03'05.1'' E 046 44'29.8''” (ZIN); two males, “MADAGASCAR Baie de Baly Jan 2004 dry forest fish baited trap [16°02′31″S, 45°14′45″E] Ilkka Hanski leg.” (ZIN). Diagnosis. Triodontus lemoulti is similar to T. ankarafantsikae in having the abdominal sternite 8 with a more or less developed tubercule medially, but differs from it in having the tubercule concave apically (as opposed to being convex in T. ankarafantsikae) and lateral lobes of parameres without keel-shaped processes on proximal parts but with semicircular notches (in dorsal view, Fig. 35E). Description. Holotype, male. Body length 10.2 mm. Body strongly shiny (Figs. 35A–B, D–F). Color of head, pronotum, and elytra dark brown; legs, antennae, and underside of the body brown. Frontoclypeus convex anteriorly, rounded laterally, anterior margin slightly crenulate and setose in dorsal view. Eyes relatively small (diameter slightly smaller than the distance between eye and gula in ventral view), incompletely divided by canthus into small dorsal and large ventral parts. Frontoclypeus with small tubercles mediad of each eye and with a short, conical horn in the center of the frontoclypeus. The horn is somewhat curved caudally. Pronotum convex, almost smooth, feebly excavated anteriorly, with 2 feebly distinct tubercles medially. Anterior margin with wide, smooth border, feebly sinuate medially. Posterior margin with fine border, not crenulate, punctate with small, narrow punctures. Scutellum rounded apically, visible part is about 1/15 length of elytra. Elytra convex, with distinct humeral humps. Maximum width approximately at the middle. Elytra with 10 feebly visible striae and with oblique line from base of 6th stria to approximately middle of 8th stria. Stria with small, indistinct punctures. Intervals with minute punctures, almost smooth. Base of elytra with an irregular row of coarse punctures each bearing small, yellow seta. Abdominal sternite 8 with a developed tubercule medially, the tubercule is concave apically (Fig. 35G). Parameres have outer lobes with semicircular notches on proximal parts (in dorsal view, Fig. 35E) and long feebly sclerotized apices. Internal sac of the aedeagus with 1 strongly sclerotized sclerite, and 2 additional similar sclerites (Fig. 35F). Female. Females differ from males in having long apical spur on protibiae, rugose punctation of head, and small frontoclypeal tubercle (Fig. 35C). Paratypes. Body length of the paratypes varies from 8.5–11.0 mm in males and from 10.0–10.5 mm in females. Larger male paratype has slightly larger pronotal excavations and smaller males have almost no excavations and very short frontoclypeal tubercle. Distribution. This species is known from a few localities in western and central Madagascar (Fig. 35H). Apparently its distribution range is limited to the dry tropical forests. Remarks. Paulian (1977) considered the specimen designated here the holotype of T. lemoulti as T. nigritus. Both syntypes of T. nigritus are females but they are similar to females of T. hova in head sculpture, whereas the female of T. lemoulti differs in having a small but distinct frontal tubercle. Collecting localities of the specimens also suggest that T. nigritus is conspecific with T. hova. The former species was described from Nosy Be Island.

70 · Zootaxa 4207 (1) © 2016 Magnolia Press FROLOV ET AL. FIGURE 35. Triodontus lemoulti¸ holotype male (A, B, D, E), paratype male (F, G), paratype female (C). Habitus in dorsal (A , C) and dorsolateral view (B), labels (D), aedeagus in lateral view and parameres in dorsal and ventral view (E), internal sac of aedeagus (F), abdomen (G), distributional records (H).

REVIEW OF THE MADAGASCAN ORPHNINAE Zootaxa 4207 (1) © 2016 Magnolia Press · 71 Etymology. This species is dedicated to the memory of E. Lemoult, French entomologist and collector of the first half of 19th century.

Triodontus modestus (Benderitter, 1914) (Figs. 36A–H)

Orphnidius modestus Benderitter, 1914a: 291. Triodontus modestus Paulian, 1977: 1222.

Type material examined. Lectotype (here designated) (Figs. 36A, B, D–G), male, “Mianinarévo [Miarinarivo] / MADAGASCAR Collection Le Moult / Orphnus ♂ modestus type E. Benderitter det. / Orphnus modestus Bend. LECTOTYPUS Frolov des. 2010” (MNHN). Paralectotypes: one male and four females with the same locality data as the lectotype (MNHN). Additional material examined. MADAGASCAR. Antananarivo: one female, “Madagascar Tananarive Collection Le Moult [Antananarivo]” (MNHN); Madagascar (no precise locality): one female, “Museum Paris Madagascar Collection Leon Fairmaire 1906” (MNHN). Diagnosis. Triodontus modestus can easily be separated from other Triodontus species with 2 tubercles on the pronotal bulge by being smaller (body length up to 6.5 mm whereas the other species are longer than 9.0 mm), and having outer lobes of the parameres with large circular situations lateroapically. All examined specimens are light brown but additional material is needed to clarify if such a pale coloration is species specific. Description. Male. Body convex, strongly shiny, with uniform light brown coloration (Figs. 36A–B). Frontoclypeus slightly convex anteriorly, obtusely rounded laterally, anterior margin setose in dorsal view. Eyes small (diameter a few times smaller than distance between eye and gula in ventral view), incompletely divided by canthus into small dorsal and large ventral parts. Frontoclypeus with small tubercles mediad of each eye and with short, tubercle-like horn in the center of the frontoclypeus. Frontoclypeus coarsely punctate apical of the horn. Pronotum without a distinct medial tubercule but with 4 relatively small tubercles in proximal 1/3, and with a shallow excavation anteriad of the tubercles. Lateral margins with wide, densely setose border. Anterior margin with wide, smooth border. Posterior margin with fine border, not crenulate; punctate with small, narrow, longitudinal punctures. Surface of pronotum sparsely punctate with small punctures. Scutellum rounded apically, its visible part is about 1/15 length of elytra. Elytra convex, with distinct humeral humps. Maximum width approximately at the middle. Elytra with 10 striae on disc and with oblique line from base of 6th stria to approximately middle of 8th stria. Stria with coarse punctures. Intervals with small but distinct punctures. Base of elytra with an irregular row of coarse punctures each bearing a small, yellow seta. Abdominal sternite 8 medially about as wide as sternites 3–5 combined, with slightly sinuate apical margin and with slight excavation in the middle. Parameres with outer lobes having large circular situations lateroapically (Fig. 36E). Internal sac of aedeagus with somewhat fork-shaped sclerite (Fig. 36F). Female. Females (Fig. 36C) can be separated from males by having apical protibial spur; relatively smaller pronotum; minute tubercle in the middle of frontoclypeus; wider abdominal sternite 8, which is not sinuated apically; and scutellum more angulate apically. Variation. Body length of examined specimens varied from 6.2–6.5 mm (males) and from 5.9–6.5 mm (females). Remarks. Benderitter (1914a) described T. modestus from two males and four females from “Mianinarévo” (misspelling of Miarinarivo). Paulian (1977) was unaware of the depository of these specimens. However, we found the complete type series at the MNHN. For the purpose of the stability of the nomenclature, one of the male specimens (Figs. 36A–B) is here designated the lectotype. Distribution. This species is known from two localities in the Central Plateau (Fig. 36H).

72 · Zootaxa 4207 (1) © 2016 Magnolia Press FROLOV ET AL. FIGURE 36. Triodontus modestus, lectotype male (A, B, D), paralectotype male (E, F), paralectotype female (C). Habitus in dorsal (A, C) and dorsolateral view (B), labels of lectotype (D), aedeagus in lateral view and parameres in dorsal and ventral view (E), internal sac of aedeagus (F), abdomen (G), distributional records (H).

Triodontus owas Westwood, 1852: 74; Paulian, 1937: 14, 1977: 1212. Triodontus vadoni Paulian, 1977: 1215, new synonym Triodontus perrotorum Paulian, 1977: 1219, new synonym

Type material examined. Orphnus owas: holotype (Figs. 37A–B), male, “Madag / Owas Westw. (Reiche) Madagasc. / Orphnus nov. sp. / Type / 67.45/ II” (BMNH). Triodontus vadoni: holotype (Fig. 38B), male, “Ambohitsitondrona Madagascar XII-52 Michel” (MNHN). Triodontus perrotorum: holotype (Fig. 38C), male, “Ste Marie de Madagascar Perrot Freres X-XII 1896 / Triodontus perrotorum n. sp. R. Paulian det. / MUSEUM PARIS 1952 COLL R OBERTUR / TYPE” (MNHN). Paratypes: two males and two females with the same locality data as the holotype (MNHN). Additional material examined. MADAGASCAR: Antsiranana: two males, one female, “Amber geb. [Amber Mountains, 12°37′00″S, 49°09′00″E]” (SMTFD); Toamasina: one male, “Ambohitsitondrona Madagascar XII-52 Michel [15°34′59″S, 50°01′00″E]” (MNHN); three males, “Madagascar Ambodivoangy I.1945 Vadon! [15°17′49″S, 49°36′47″E,]” (MNHN); one male, “Madagascar Est Dist. Mananara - N Mont Antampona Vadon Peyrieras [Mananara District, North of Antampona Mountain, 15°58′00″S, 49°13′00″E]” (MNHN); one male, “Madagascar Fenerive E.Perrot [17°22′00″S, 49°25′00″E]” (MNHN); one male, “Madagascar Ambohitsitondrona I-1946 Vadon! [15°34′59″S, 50°01′00″E]” (MNHN); one male, one female, “Madagascar Ambohitsitondrona II- 1948 Michel [15°34′59″S, 50°01′00″E]” (MNHN); one male, “Madagascar Fampanambo [15°22′15″S, 49°37′59″E] XII” (MNHN); Fianarantsoa: one male, “Madagascar Foret Tanala Alluaud 1901 [Tanala Forest,]” (MNHN); one male, “Madagascar Fianarantsoa Perrot Freres 2 Semestre 1892 [21°25′59″S, 47°04′59″E]” (MNHN); one male, one female, “Madagascar Est Ranomafana XI.2004 [Ranomafana National Park, 21°18′00″S, 47°30′00″E, XI.2004, O. Montreuil]” (OMCP); Mahajanga: three males, two females, “Ste Marie de Madagascar Perrot Freres X–XII 1896 [Sainte Marie de Madagascar, near Marovoay, 16°06′00″S, 46°37′59″E]” (MNHN); Madagascar (no precise locality): three males (SMTFD); three males (IRSNB); one male (NMPC). Differential diagnosis. From the most similar species, T. viettei new species and T. hildebrandtii, T. owas differs in having the outer lobes of the parameres relatively slender in lateral and dorsal view (Fig. 37D). Description. Male. Color brown to dark brown, legs, antennae, and underside of the body slightly paler (Figs. 37A, 38A, C). Frontoclypeus slightly convex anteriorly, rounded laterally, anterior margin crenulate and setose in dorsal view. Eyes relatively small (diameter slightly smaller than distance between eye and gula in ventral view), incompletely divided by canthus into small dorsal and large ventral parts. Frontoclypeus with tubercles mediad of each eye and with long horn in the center of the frontoclypeus. The horn is longer than width of the head, acutely rounded apically, somewhat curved caudally, slightly rugose on posterior side. Pronotum with a bulge medially, with large excavations adjacent to bulge, and with 2 ridge-shaped tubercles laterally of each excavation. The median bulge is rounded and somewhat depressed on top, with 2 distinct tubercles and a depression, receiving frontoclypeal horn. Lateral margins with wide border appearing somewhat crenulate (in dorsal view), with a row of brown setae. Anterior margin with wide, smooth border. Posterior margin with fine border; not crenulate; punctate with small, narrow, longitudinal punctures. Surface of most pronotum almost smooth, with minute, feebly visible punctation. Scutellum rounded apically, visible part is about 1/13 length of elytra. Elytra convex, with distinct humeral humps. Maximum width approximately at basal 1/3. Elytra with 10 feebly visible striae on disc and with oblique line from base of 6th stria to approximately middle of 8th stria. Stria with small punctures. Intervals with sparse, minute punctures, almost smooth. Base of elytra with an irregular row of coarse punctures, each bearing a small, yellow seta. Sternite 8 medially as wide as sternites 2–5 combined, without tubercle or concavity in middle, with apical border deeply sinuate medially (Fig. 37F). Parameres (Fig. 37D) with outer lobes relatively slender in lateral and dorsal view, strongly sclerotized, with distinct lateral notches and teeth situated more or less apically. Internal sac of the aedeagus with 1 strongly sclerotized sclerite and 2 additional similar sclerites (Fig. 37E).

74 · Zootaxa 4207 (1) © 2016 Magnolia Press FROLOV ET AL. FIGURE 37. Triodontus owas, holotype male (A, B, D), male (E, F), female (C). Habitus in dorsal view (A, C), labels of holotype (B), aedeagus in lateral view and parameres in dorsal and ventral view (D), internal sac of aedeagus (E), abdomen (F), distributional records (G).

REVIEW OF THE MADAGASCAN ORPHNINAE Zootaxa 4207 (1) © 2016 Magnolia Press · 75 FIGURE 38. Triodontus owas, habitus in dorsal view. Male specimen from Fampanambo (A), holotype of T. vadoni (B), holotype of T. perrotorum (C).

Female. Females (Fig. 37C) differ from males in having an apical spur on protibiae; absence of head and pronotum armature including tubercles mediad of each eye; and longer abdominal sternites including 6th sternite, which is evenly rounded apically. Variation. Pronotal and head armature of males is subject to allometric variability from fully developed (as described above) to less developed (with the males having short frontoclypeal tubercles and small excavation on the pronotum anteriorly). Body length of examined specimens varied from 11.0–14.0 mm (males) and from 11.0– 12.0 mm (females). Distribution. Triodontus owas is known from a few localities mostly from the eastern Madagascar: from Amber Mountain in the north to Ranomafana National Park in the south (Fig. 37G). The localities are situated in the eastern rainfall forest ecoregion except for Marovoay, which is situated in the dry deciduous forest ecoregion in western areas of the island. Remarks. It can be inferred from the Paulian’s (1977) work, that he did not examine the holotype of T. owas. The characters and illustrations provided suggest that the specimens he considered T. owas belonged to T. owas, T. itremoi, and T. viettei, as treated here. Figures of the parameres (Paulian 1977, Figs. 22–23) belonged to T. itremoi, as treated here, while the description of the pronotum shape corresponds rather to T. owas, T. copridoides, or T. viettei. Examination of the types of T. vadoni and T. perrotorum showed that they both have diagnostic characters of T. owas. The differences between T. vadoni and T. perrotorum are due to the allometric variability of T. owas and, therefore, two new synonymies are here proposed.

Triodontus viettei Frolov, Montreuil & Akhmetova, new species (Figs. 39A–H)

Type material. Holotype (Figs. 39A–C, E–G), male, “Madagascar Antsianaka et lac Alaotra 2e Trimestre 1889 Perrot Freres / HOLOTYPUS Triodontus viettei Frolov et al. 2010” (MNHN). Paratypes: one male, “Tananarive Lamberton” (MNHN); one male and two females, “Madagascar Nord, contreforts du Tsaratanana, Haut Sambirano, 1200 m, vallee de la Besanetribe [Upper Sambirano River, 1200 m

FIGURE 39. Triodontus viettei, holotype male (A, B, C, E, F, G), paratype female (D). Habitus in dorsal (A, D) and dorsolateral view (B), labels of holotype (C), aedeagus in lateral view and parameres in dorsal and ventral view (E), internal sac of aedeagus (F), abdomen (G), distributional records (H).

REVIEW OF THE MADAGASCAN ORPHNINAE Zootaxa 4207 (1) © 2016 Magnolia Press · 77 Diagnosis. Triodontus viettei is most similar to T. hildebrandtii but differs from it in having outer lobes of the parameres with slit-shaped, lateral notches, as opposed to having wide lateral notches in T. hildebrandtii. Description. Holotype, male. Body length 14.5 mm. Color uniformly dark brown, legs, antennae, and underside of the body slightly paler (Fig. 39A–B). Frontoclypeus slightly convex anteriorly, rounded laterally, anterior margin crenulate and setose in dorsal view. Eyes relatively small (diameter slightly smaller than the distance between eye and gula in ventral view), incompletely divided by canthus into small dorsal and large ventral parts. Frontoclypeus with tubercles mediad of each eye and with long horn in the center of frontoclypeus. The horn is longer than width of the head, acutely rounded apically, somewhat curved caudally, slightly rugose on posterior side. Pronotum with a bulge medially, with large excavations at each side of bulge, and with 2 ridge-shaped tubercles laterally of each excavation. The median bulge is rounded and somewhat depressed on the top, with 2 distinct tubercles and a depression, receiving frontoclypeal horn. Lateral margins with wide border appearing somewhat crenulate (in dorsal view), with a row of brown setae. Anterior margin with wide, smooth border. Posterior margin with fine border; not crenulate; punctate with small, narrow, longitudinal punctures. Surface of most pronotum almost smooth, with minute, feebly visible punctation. Scutellum rounded apically, visible part is about 1/13 length of elytra. Elytra convex, with distinct humeral humps. Maximum width approximately at basal 1/3. Elytra with 10 feebly visible striae on disc and with oblique line from base of 6th stria to approximately middle of 8th stria. Stria with small punctures. Intervals with sparse, minute punctures, almost smooth. Base of elytra with an irregular row of coarse punctures each bearing a small, yellow seta. Sternite 8 medially as wide as sternites 2–5 combined, without tubercle or concavity in the middle, with apical border deeply sinuate medially (Fig. 39G). Parameres with outer lobes being relatively wide (in dorsal view) and having slit-shaped, lateral notches (Fig. 39E). Internal sac of the aedeagus with 1 strongly sclerotized sclerite and 2 additional similar sclerites (Fig. 39F). Female. Females differ from males in having apical spur on protibiae; absence of the head and pronotum armature; densely punctate dorsal side of the head; and in longer abdominal sternites including 6th sternite, which is evenly rounded apically (Fig. 39D). Variation. Body length of the examined paratypes varied from 13.5–14.0 mm (males) and from 11.7–11.9 mm (females). Distribution. Triodontus viettei is known from four localities in eastern and northern Madagascar (Fig. 39H). Etymology. The new species is dedicated to the memory of Pierre Viette, French entomologist, specialist of Noctuidae (Lepidoptera) of Madagascar.

Triodontus hildebrandtii (Fairmaire, 1883) (Figs. 40A–E)

Orphnus hildebrandtii Fairmaire, 1883: 365, 1884: 131. Orphnus hildebrandti: Arrow 1912: 29. Triodontus hildebrandti: Paulian 1937: 142, 1977: 1213.

Type material examined. Holotype (Figs. 40A–E), male, “MUSÉUM PARIS 1906 Coll. Leon FAIRMAIRE / Orphnus Hildebrandti Waterh / 434” (MNHN). Additional material examined. Two males, “Madagascar” (IRSNB). Diagnosis. Triodontus hildebrandtii is most similar to T. viettei but differs from it in having outer lobes of parameres with wide lateral notches, as opposed to having slit-shaped lateral notches in T. viettei. Description. Male. Body length 15.0 mm. Color uniformly dark brown, legs, antennae, and underside of the body slightly paler (Figs. 40A–B). Frontoclypeus slightly convex anteriorly, rounded laterally, anterior margin crenulate and setose in dorsal view. Eyes relatively small (diameter slightly smaller than the distance between eye and gula in ventral view), incompletely divided by canthus into small, dorsal and large, ventral parts. Frontoclypeus with tubercles mediad of each eye and with long horn in the center of the frontoclypeus. The horn is longer than width of the head, acutely rounded apically, somewhat curved caudally, slightly rugose on posterior side.

78 · Zootaxa 4207 (1) © 2016 Magnolia Press FROLOV ET AL. FIGURE 40. Triodontus hildebrandtii, holotype male. Habitus in dorsal (A) and lateral view (B), internal sac of aedeagus (C), labels of holotype (D), aedeagus in lateral view and parameres in dorsal and ventral view (E).

Pronotum with a bulge medially, with large excavations each side of bulge, and with 2 ridge-shaped tubercles laterally of each excavation. Median bulge is rounded and somewhat depressed on top, with 2 distinct tubercles and a depression, receiving frontoclypeal horn. Lateral margins with wide border appearing somewhat crenulate (in dorsal view), with a row of brown setae. Anterior margin with wide, smooth border. Posterior margin with fine border; not crenulate; punctate with small, narrow, longitudinal punctures. Surface of most pronotum almost smooth, with minute, feebly-visible punctation. Scutellum rounded apically, visible part is about 1/13 length of elytra. Elytra convex, with distinct humeral humps. Maximum width approximately at basal 1/3. Elytra with 10 feebly visible striae on disc and with oblique line from base of 6th stria to approximately middle of 8th stria. Stria with small punctures. Intervals with sparse, minute punctures, almost smooth. Base of elytra with an irregular row of coarse punctures each bearing a small, yellow seta. Sternite 8 medially as wide as sternites 2–5 combined, without tubercle or concavity in the middle, with apical border deeply sinuate medially. Parameres with outer lobes being relatively wide (in dorsal view) and having slit-shaped lateral notches (Fig. 40E). Internal sac of aedeagus with 1 strongly sclerotized sclerite and 2 additional smaller sclerites (Fig. 40C).

REVIEW OF THE MADAGASCAN ORPHNINAE Zootaxa 4207 (1) © 2016 Magnolia Press · 79 Female. Unknown. Variation. The non-type specimens differ from the holotype in slightly less-developed frontoclypeal and pronotal armature. Body length varies from 14.0–14.5 mm. Distribution. The distribution of this species is yet unknown since all three known specimens lack precise locality data. Remarks. Paulian wrote that the type of T. hildebrandtii was unknown to him, but he treated it a valid species and mentioned some material from Tanala forest (collection of Alluaud) (Paulian 1937, 1977). There is one Triodontus specimen in the collection of MNHN, which bears the label “Forêt Tanala (coll. Alluaud)”. This specimen, in our opinion, belongs to T. owas. We think that the application of the name T. hildebrandtii to this or other specimens was unjustified. In the collection of MNHN we found a specimen with a label “Orphnus Hildebrandti Waterh”. Although the label is not handwritten by Fairmaire, the specimen originated from Fairmaire’s collection and agrees with the original description of Orphnus hildebrandtii. Therefore we consider it the holotype.

Triodontus alticola Paulian, 1977 (Figs. 41A–H)

Triodontus alticola Paulian, 1977: 1215. Triodontus maroantsetrae Paulian, 1977: 1219, new synonym

Type material examined. Triodontus alticola. Holotype (Figs. 41C, D), male, “Madagascar Ambohitsitondrona [Toamasina Province] XI-1949 Michel / Triodontus alticola n. sp. R. Paulian det. / TYPE” (MNHN). Paratypes: two females with the same locality and date label (MNHN); two males and two females with the same locality label but the date “XII-52” (MNHN); one male, “Madagascar Sambava [Antsiranana Province] XI [...] Vadon!” (MNHN); one male, “Madagascar Ambodivoangy [Toamasina Province] I.1950 Vadon!” (MNHN). Triodontus maroantsetrae. Holotype, male, “Madagascar Ron Maroantsetra XII.35 Vadon Antakotako / MUSEUM PARIS 1938 J. Vadon et E. Lebis / TYPE / Triodontus maroantsetrae n. sp. R. Paulian det.” (MNHN). Additional material examined. MADAGASCAR: Toamasina: two males, “Madagascar Ambohitsitondrona II-1948 Michel [15°34′59″S, 50°01′00″E]” (MNHN); one male, “Madagascar Ambodivoangy [15°17′49″S, 49°36′47″E] I.1946 Vadon!” (MNHN); one male, same label but XII.1947 (MNHN); one female, “Madagascar Antakotako [15°19′00″S, 49°48′00″E] II.1946 Vadon!” (MNHN); one male, “Madagascar Est distr. Mananara - N. Antanambe Vadon et Peyrieras [Mananara District, Antanambe, 16°25′59″S, 49°51′00″E]” (MNHN); four males, one female, “Madagascar Ambohitsitondrona I-1946 Vadon! [15°34′59″S, 50°01′00″E]” (MNHN); one male, “Madagascar Ambohitsitondrona 25-I-1948 Michel [15°34′59″S, 50°01′00″E, 25.I.1948]” (MNHN); one male, one female, “Madagascar Ambohitsitondrona X-1947 Vadon! [15°34′59″S, 50°01′00″E]” (MNHN). Diagnosis. Triodontus alticola can be separated from other species of the genus by the characteristic shape of the parameres having the outer lobes long, curved downward and inward; with more-or-less separated, strongly sclerotized lateral parts and membranous, overlapping medial parts; and without lateral notches. Description. Male. Color of head, pronotum, and elytra dark brown, legs, antennae, and underside of the body slightly paler (Figs. 41A, C). Frontoclypeus slightly convex anteriorly, obtusely rounded laterally, anterior margin setose in dorsal view. Eyes relatively small (diameter slightly smaller than the distance between eye and gula in ventral view), incompletely divided by canthus into small, dorsal and large, ventral parts. Frontoclypeus with tubercles mediad of each eye and with a horn in center. The horn is longer than width of the head, acutely rounded apically, somewhat curved caudally, with slightly flattened anterior side and slightly rugose posterior side. Pronotum with a bulge medially in center, with excavations each side of bulge, and with 2 ridge-shaped tubercles laterally of each excavation. The median bulge with 2 distinct tubercles and somewhat excavated between the tubercles. Lateral margins with wide border bearing a row of brown setae. Anterior margin with wide, smooth border, almost not sinuate medially. Posterior margin with fine border, not crenulate, punctate with a few narrow, longitudinal punctures. Surface of most pronotum punctate with minute, feebly visible puctures. Scutellum rounded apically, visible part is about 1/15 length of elytra.

80 · Zootaxa 4207 (1) © 2016 Magnolia Press FROLOV ET AL. FIGURE 41. Triodontus alticola, holotype male (C, D, E, F), male (A), female (B). Habitus in dorsal view (A, B, C), aedeagus in lateral view and parameres in dorsal and ventral view (E), labels of holotype (D), internal sac of aedeagus (F), abdomen (G), distributional records (H).

REVIEW OF THE MADAGASCAN ORPHNINAE Zootaxa 4207 (1) © 2016 Magnolia Press · 81 Elytra convex, with distinct humeral humps. Maximum width approximately at middle. Elytra with 10 striae; stria 1 (sutural) distinct, other feebly marked with small punctures. Intervals with minute punctures, almost smooth. Base of elytra with an irregular row of coarse punctures. Abdominal sternite 8 medially deeply sinuate, without tubercles or concavities in middle (Fig. 41G). Parameres (Fig. 41E) with outer lobes long, curved downward and inward; with more-or-less separated, strongly sclerotized lateral parts and membranous, overlapping medial parts; and without lateral notches. Internal sac of aedeagus with a hoof-shaped sclerite (Fig. 41F). Female. Females (Fig. 41B) differ from males in having a protibial apical spur; absence of head and pronotum armature including tubercles mediad of each eye; and longer abdominal sternites including 6th sternite, which is evenly rounded apically. Variation. The pronotal and head armature of males is less developed in some specimens. Body length of examined specimens varied from 8.8–12.0 mm (males) and from 9.0–11.0 mm (females). Distribution. This species is known from a few localities in the northeastern part of the island, mostly north of Antongil Bay (Fig. 41H). Remarks. Triodontus maroantsetrae was described from one male specimen from Antakotako. Comparison of this specimen with the type series of T. alticola has shown that they have similar shape of the parameres. The only difference is the feebly developed frontoclypeal horn and almost absent ridges and tubercles on the pronotum in the holotype of T. maroantsetrae. We believe all these specimens fall within the limits of allometric variability of one species, therefore the new synonymy is here proposed.

Triodontus fairmairei Frolov, Montreuil & Akhmetova, new species (Figs. 42A–F)

Type material. Holotype (Figs. 42A–F), male, “Madag […] / MUSEUM PARIS MADAGASCAR Collection Leon Fairmaire 1906 / HOLOTYPUS Triodontus fairmairei Frolov et al. 2010 ” (MNHN). Diagnosis. From other Triodontus species, T. fairmairei can be separated by the outer lobes of the parameres without lateral notches, relatively large inner lobes of the parameres, and somewhat Y-shaped sclerite of the internal sac of the aedeagus. Description. Holotype, male. Body length 8.9 mm. Color of head, pronotum, and elytra blackish brown; legs, antennae, and underside of the body brown (Figs. 42A–B). Frontoclypeus slightly convex anteriorly, obtusely rounded laterally, anterior margin setose in dorsal view. Eyes relatively small (diameter slightly smaller than the distance between eye and gula in ventral view), incompletely divided by canthus into small, dorsal and large, ventral parts. Frontoclypeus with tubercles mediad of each eye and with a horn in the center. The horn is shorter than width of the head, acutely rounded apically, somewhat curved caudally, with slightly flattened anterior side and slightly rugose posterior side. Pronotum with a bulge medially in center, with excavations aside the bulge, and with 2 ridge-shaped tubercles laterally of each excavation. The median bulge with 2 distinct tubercles and somewhat excavated between the tubercles. Lateral margins with wide border bearing a row of brown setae. Anterior margin with wide, smooth border, almost not sinuate medially. Posterior margin with fine border; not crenulate; punctate with a few narrow, longitudinal punctures. Surface of most pronotum punctate with minute, feebly visible puctures. Scutellum rounded apically, visible part about 1/14 length of elytra. Elytra convex, with humeral humps. Maximum width approximately at the middle. Elytral striae indistinct (except for stria 1). Intervals with minute punctures, almost smooth. Base of elytra with an irregular row of coarse punctures. Sternite 8 medially slightly sinuate, without tubercles or concavities in the middle (Fig. 42D). Parameres (Fig. 42F) have outer lobes without lateral notches, inner lobes relatively large. Internal sac of aedeagus with Y-shaped sclerite (Fig. 42C). Female. Unknown. Distribution. The only known specimen of this species lacks information about the collection locality except for the reference to Madagascar. Remarks. The holotype lacks left prothoracic leg and left metatibia.

82 · Zootaxa 4207 (1) © 2016 Magnolia Press FROLOV ET AL. FIGURE 42. Triodontus fairmairei, holotype male. Habitus in dorsal (A) and dorsolateral view (B), internal sac of aedeagus (C), abdomen (D), labels of holotype (E), aedeagus in lateral view and parameres in dorsal and ventral view (F).

Etymology. The new species is dedicated to the memory of Leon Fairmaire, French entomologist of the 19th century, who described a significant part of the Madagascan Coleoptera fauna.

Triodontus itremoi Paulian, 1977 (Figs. 43A–I)

Triodontus itremoi Paulian, 1977: 1210.

Type material examined. Holotype (Figs. 43A–C), male, “MADAGASCAR CENTRE massif de l'Itremo / A. Peyrieras II.1974 / Triodontus itremoi n. sp. R. Paulian det. / TYPE” (MNHN).

REVIEW OF THE MADAGASCAN ORPHNINAE Zootaxa 4207 (1) © 2016 Magnolia Press · 83 FIGURE 43. Triodontus itremoi, holotype male (A, B, C), paratype male (E right, F, G, H), paratype female (E left, D). Habitus in dorsal (A, D) and dorsolateral view (B), labels (C), abdomen (E, H), aedeagus in lateral view and parameres in dorsal and ventral view (F), internal sac of aedeagus (G), distributional records (I).

84 · Zootaxa 4207 (1) © 2016 Magnolia Press FROLOV ET AL. Paratypes: three males and six females with the same locality data (MNHN). Additional material examined. MADAGASCAR: Toamasina: one male, “Madagascar route Tamatave PK181 Beforona [road Tamatave PK181, Beforona, 18°58′00″S, 48°34′59″E] 8-X-1973 / R.Viossat” (MNHN); four males, “Perinet [18°55′59″S, 48°25′00″E]” (MNHN); one male, one female, “Madagascar C. Moramanga env. [Moramanga environs, 18°55′59″S, 48°12′00″E] 10–18.XII.1997 P.Pacholatko leg.” (NHMB); four males, two females, “MADAGASCAR: Toamasina Ambatovy, 12.4 km NE Moramanga elev 1040 m pitfall trap, 5–8 March 2007 18˚ 51' 29" S 048˚ 17' 06" E California Acad. of Sciences coll. B.L.Fisher et al. grassland BLF16919” (CASC); one male, “MADAGASCAR: Toamasina Ambatovy, 12.4 km NE Moramanga elev 1080 m pitfall trap, 4–7 March 2007 18˚ 50' 22" S 048˚ 18' 30" E California Acad. of Sciences coll. B.L.Fisher et al. montane rainforest BLF16917” (CASC); one male, “Madagascar Andasibe [18°55′48″S, 48°25′12″E] XI 2005 O.Montreuil leg.” (MNHN); one male, “Madagascar Centr. Plat., Andasibe, Perinet 950 m [18°55′59″S, 48°25′00″E], 1– 5.01.2002 V.Dolin & R.Andreeva” (NMPC); one male, two females, “Madagascar 135 km E Antananarivo Andasibe env. h=900 m [Andasibe environs, 900 m, 18°55′48″S, 48°25′12″E] 20–30.XII.2000 S.Murzin leg” (ABCB); Antananarivo: three males, “Tananarive Lamberton [Antananarive, 18°55′00″S, 47°31′00″E, Lamberton leg.]” (MNHN); Fianarantsoa: three males, two females, “Madagascar Ranomafana NP, Vatoharana transect trail S 600 m [21°18′00″S, 47°30′00″E], 15.1.2001, Johana Rainio leg.” (UHHF); one male, “Deans Cowan Betsileo Madagascar [Betsileo, D. Cowan leg.] / 1881 3000 ft 4000 ft [3000–4000 ft, 1881]” (SMTFD); two males, “[Ranomafana NP H.Viljanen] Nov 21-[20]02 PFT Primary forest S725” Ranomafana National Park, primary forest, 21°18′00″S, 47°30′00″E, 21.XI.2002, H.Viljanen leg., (ZIN); six males, 10 females, “Madagascar, Fianarantsoa Prov., Foret d'Atsirakambiaty, 7.6 km 285° WNW Itremo, elev 1550 m., 22–26 January 2003 / 20°35′36″S, 046°33′48″E colls. Fisher Griswold et al. Calif. Acad. of Sci. pitfall trap - montane rainforest collection code BLF7150” (CASC); Toliara: one male, “S.-P. Tsiroanomandidy [...] du Bongolava 1300 m. A.Peyrieras 6/13.XII.1974 [Tsiroanomandidy District, Bongolava, 1300 m, 18°30′00″S, 45°30′00″E, 6– 13.XII.1974, A. Peyrieras]” (MNHN); Madagascar (no precise locality): one male (MNHN); one male (IRSNB). Diagnosis. Males of T. itremoi are most similar to T. copridoides especially in the shape of the parameres and internal sac armature. Triodontus itremoi can be separated from the later by having the abdominal sternite 8 without trace of concavity (as opposed to being slightly but distinctly concave in T. copridoides), reasonably larger tubercles laterad of each eye, and somewhat longer and more oblique fossae on the lateral sides of outer lobes of the parameres. Description. Male. Body length varied from 12.5–14.5 mm. Color of head, pronotum, and elytra dark brown; legs, antennae, and underside of the body paler, castaneous (Figs. 43A–B). Frontoclypeus slightly convex anteriorly, rounded laterally, anterior margin crenulate and setose in dorsal view. Eyes relatively small (diameter slightly smaller than the distance between eye and gula in ventral view), incompletely divided by canthus into small dorsal and large ventral parts. Frontoclypeus with tubercles mediad of each eye and with long horn in center of frontoclypeus. The horn is longer than width of head, acutely rounded apically, somewhat curved caudally, slightly rugose on posterior side. Pronotum with a bulge medially in distal 1/3, with large excavations aside the bulge, and with 2 ridge-shaped tubercles laterally of each excavation. The median bulge is rounded and somewhat depressed on the top, without distinct tubercles but with a longitudinal depression receiving frontoclypeal horn. Pronotal and head armature is less developed in some specimens. Lateral margins with wide border appearing somewhat crenulate on the hind angles (in dorsal view), with a row of brown setae. Anterior margin with wide, smooth border. Posterior margin with fine border; not crenulate; punctate with small, narrow, longitudinal punctures. Surface of most pronotum smooth, anterior and posterior angles with sparse and coarse punctation. Scutellum rounded apically, visible part is about 1/15 length of elytra. Elytra convex, with distinct humeral humps. Maximum width approximately at the middle. Elytra with 10 feebly visible striae on disc and with oblique line from base of 6th stria to approximately middle of 8th stria. Stria with small punctures. Intervals with minute punctures, almost smooth. Base of elytra with an irregular row of coarse punctures each bearing small, yellow seta. Abdominal sternite 8 medially about 1.5 as short as other sternites combined, with slight excavation in ventral view, without any tubercles or concavities in the middle (Figs. 43E right, H). Parameres (Fig. 43F) with outer lobes somewhat depressed, with feebly sclerotized wide apices, with deep oblique fossae on lateral sides (Fig. 43F, arrow). Internal sac of aedeagus with long, sclerotized sclerites (Fig.

REVIEW OF THE MADAGASCAN ORPHNINAE Zootaxa 4207 (1) © 2016 Magnolia Press · 85 43G). Female. Females (Figs. 43D–E left) differ from males in having long apical spur on protibiae; absence of head and pronotum armature including tubercles mediad of each eye; coarsely punctate dorsal side of head; and longer abdominal sternites including 6th sternite, which is evenly rounded apically. Body length of examined females varied from 12.0–13.0 mm. Distribution and habitat. Triodontus itremoi is known from a few localities in central and eastern Madagascar (Fig. 43I). Most of the localities are situated within primary rain forests.

Triodontus copridoides Paulian, 1977 (Figs. 44A–F)

Triodontus copridoides Paulian, 1977: 1211.

Type material examined. Holotype (Figs. 44A–B, D–E), male , “Madagascar Cowans [= Ankafina Tsarafidy, according to Paulian (1977)] / Ex Musæo D.Sharp 1890 / MUSÉUM PARIS 1952 COLL R OBERTHUR / Triodontus copridoides n. sp. R. Paulian det. / TYPE” (MNHN). Additional material examined. MADAGASCAR: Antsiranana: two males, “Amber geb. [Amber Mountains]” (SMTFD). Diagnosis. Triodontus copridoides is most similar to T. itremoi in general appearance and in the shape of the parameres but it can be separated from it by having the abdominal sternite 8 with a shallow but distinct, somewhat transversal fossa, larger tubercles laterad of each eye, and somewhat shorter and less oblique notches on the lateral sides of the outer lobes of the parameres. Description. Male. Color of head, pronotum, and elytra dark brown; legs, antennae, and underside of the body paler, castaneous (Figs. 44A, C). Frontoclypeus slightly convex anteriorly, rounded laterally, anterior margin crenulate and setose in dorsal view. Eyes relatively small (diameter slightly smaller than the distance between eye and gula in ventral view), incompletely divided by canthus into small, dorsal and large, ventral parts. Frontoclypeus with tubercles mediad of each eye and with long horn in center of frontoclypeus. The horn is longer than width of the head, acutely rounded apically, somewhat curved caudally, slightly rugose on posterior side. Pronotum with a bulge medially in distal 1/3, with large excavations aside the bulge, and with 2 ridge-shaped tubercles laterally of each excavation. The median bulge is rounded and somewhat depressed on the top, without distinct tubercles but with a longitudinal depression receiving frontoclypeal horn. Lateral margins with wide border appearing somewhat crenulate on the hind angles (in dorsal view), with a row of brown setae. Anterior margin with wide, smooth border. Posterior margin with fine border; not crenulate; punctate with small, narrow, longitudinal punctures. Surface of pronotum mainly smooth, anterior and posterior angles with sparse and coarse punctation. Scutellum rounded apically, visible part is about 1/15 length of elytra. Elytra convex, with distinct humeral humps. Maximum width approximately at the middle. Elytra with 10 feebly visible striae on disc and with oblique line from base of 6th stria to approximately middle of 8th stria. Stria with small punctures. Intervals with minute punctures, almost smooth. Base of elytra with an irregular row of coarse punctures each bearing small, yellow seta. Protibiae with 3 outer teeth, not crenulate basad of the teeth. Apices with 3 robust, spur-like setae and a few small setae. Anterior tarsi about 3/4 length of protibiae. Claws 1/3 length of apical tarsomere. Apical protarsomere as long as tarsomere 3 and 4 combined, as thick as other tarsomeres. Ventral surface of protibiae with a longitudinal keel. Mesothoracic and metathoracic legs similar in shape; metafemora and metatibiae slightly longer than mesofemora and mesotibiae, respectively. Tibiae somewhat triangular, with 2 apical spurs, with inner margin only slightly concave. Longer tibial spur as long as or slightly longer than 2 basal tarsomeres. Claws about 1/3 length of apical tarsomere. Ventral sides of femora almost impunctate; with sparse, long setae. Sternite 8 medially about 1.5 as short as other sternites combined; with distinct, elongate, transverse fossa in the middle. Parameres with outer lobes somewhat depressed, feebly sclerotized wide apices, and deep oblique fossae on

FIGURE 44. Triodontus copridoides, holotype male (A, B, D, E), male (C). Habitus in dorsal view (A, C), labels (D), aedeagus in lateral view and parameres in dorsal and ventral view (D), abdomen (E), distributional records (F).

Female. Unknown.

REVIEW OF THE MADAGASCAN ORPHNINAE Zootaxa 4207 (1) © 2016 Magnolia Press · 87 Variation. Body length of examined specimens varied from 15.5–16.0 mm, otherwise the three specimens are rather similar. Distribution. This species is known from two rather distant localities in central Madagascar, Ankafina Tsarafidy (as suggested by Paulian (1977)) and northern Madagascar (Amber Mountains) (Fig. 44F).

Triodontus hanskii Frolov, 2010 (Figs. 45A–G)

Triodontus hanskii Frolov, 2010b: 1031.

Type material examined. Holotype, male, “MADAGASCAR Ambatotsirongorongo Feb 2005 fish baited trap Ilkka Hanski leg. / Triodontus hanskii Frolov 2010 HOLOTYPUS” (ZIN). Diagnosis. From other species with 2 medial tubercles on the pronotum and simple shape of the 6th abdominal sternite in males (T. owas Westwood, T. itremoi Paulian, T. copridoides Paulian), T. hanskii can be separated by having the parameres without teeth and notches laterally (Fig. 45E). Description. Male. Body length 10.1 mm. Body strongly shiny, uniformly colored (Figs. 45A–B). Color of head, pronotum, and elytra dark brown; legs, antennae, and underside of the body brown. Frontoclypeus convex anteriorly, rounded laterally, anterior margin crenulate and setose in dorsal view. Eyes relatively small (diameter slightly smaller than the distance between eye and gula in ventral view), incompletely divided by canthus into small, dorsal and large, ventral parts. Frontoclypeus with small but distinct tubercles mediad of each eye and with short horn in center of frontoclypeus. Pronotum with 2 obtuse tubercles in the middle and with excavations aside the tubercles. Lateral margins with wide border appearing somewhat crenulate in dorsal view. Anterior margin with wide, smooth border. Posterior margin with fine border; not crenulate; punctate with small, narrow, longitudinal punctures laterally. Surface of most pronotum smooth, anterior angles with a few coarse punctures. Scutellum rounded apically, visible part is about 1/15 length of elytra. Elytra convex, with distinct humeral humps. Maximum width approximately at middle. Elytra with 10 feebly visible striae and with oblique line from base of 6th stria to approximately middle of 8th stria. Stria with small, indistinct punctures. Intervals with minute punctures, almost smooth. Base of elytra with an irregular row of coarse punctures each bearing small, yellow seta. Sternite 8 medially about 3 times as short as other sternites combined, slightly sinuate apically, without tubercles or distinct concavities in the middle (Fig. 45F). Parameres with upper lobes somewhat tapering apically and sinuate on lateral margin basally (Fig. 45E). Internal sac of aedeagus with a somewhat pentagonal main sclerite and smaller auxiliary sclerites (Fig. 45D). Female. Unknown. Distribution and habitat. The species is known from a single specimen collected in Ambatotsirongorongo forest in Tolagnaro region of South Madagascar. This is the southernmost locality of Triodontus species known to date and it is also the southernmost remnant of indigenous rain forest on the island (Fig. 45G). The only known specimen of T. hanskii was collected in a pitfall trap baited with fish carrion. Short trap exposure (1 day and night) suggests that the specimen was attracted to carrion rather than captured accidently.

Triodontus inexpectatus Frolov, Montreuil & Akhmetova, new species (Figs. 46A–F)

Type material. Holotype (46 A–E), male, “MADAGASCAR E. Masoala, 50–470 m R. Antsamanarana 3–7/xii/ 1993, Flight Intercept Trap BMNH(E) 1994-138 / Holotypus Triodontus inexpectatus Frolov et al. 2010” (BMNH). Diagnosis. Triodontus inexpectatus is most similar to T. hanskii but can be separated from it in having inner lobes of the parameres relatively larger (in lateral view) and outer lobes as long as the inner lobes. Description. Male. Body length 12.5 mm, width 6.5 mm. Body strongly shiny, uniformly colored (Figs. 46A– B). Color of head, pronotum, and elytra dark brown; legs, antennae, and underside of body brown. Frontoclypeus convex anteriorly, rounded laterally, anterior margin crenulate and setose in dorsal view. Eyes

88 · Zootaxa 4207 (1) © 2016 Magnolia Press FROLOV ET AL. relatively small (diameter slightly smaller than the distance between eye and gula in ventral view), incompletely divided by canthus into small, dorsal and large, ventral parts. Frontoclypeus with tubercles mediad of each eye and with short obtuse horn in center of frontoclypeus.

FIGURE 45. Triodontus hanskii, holotype male. Habitus in dorsal (A) and dorsolateral view (B), labels (C), internal sac of aedeagus (D), aedeagus in lateral view and parameres in dorsal and ventral view (E), abdomen (F), distributional records (G).

REVIEW OF THE MADAGASCAN ORPHNINAE Zootaxa 4207 (1) © 2016 Magnolia Press · 89 FIGURE 46. Triodontus inexpectatus, holotype male. Habitus in dorsal (A) and dorsolateral view (B), labels (C), aedeagus in lateral view and parameres in dorsal and ventral view (D), abdomen (E), distributional records (F).

Pronotum convex, without tubercles or ridges, with shallow concavity in the middle. Lateral margins with wide border appearing somewhat crenulate in dorsal view. Anterior margin with wide, smooth border. Posterior

90 · Zootaxa 4207 (1) © 2016 Magnolia Press FROLOV ET AL. margin with fine border; not crenulate; punctate with narrow, longitudinal punctures laterally. Surface of pronotum mainly smooth, anterior angles with a few coarse punctures. Scutellum rounded apically, visible part is about 1/15 length of elytra. Elytra convex, with distinct humeral humps. Maximum width approximately at middle. Elytra with 10 feebly visible striae and with oblique line from base of 6th stria to approximately middle of 8th stria. Stria marked with small, indistinct punctures. Intervals with minute punctures, almost smooth. Base of elytra with an irregular row of coarse, semicircular punctures each bearing small, yellow seta. Sternite 8 medially about 2 times as short as other sternites combined, slightly sinuate apically, without tubercles or concavities in the middle (Fig. 46E). Parameres with apices of upper lobes being obtusely rounded, strongly sclerotized; upper lobes deeply sinuate laterally, the sinuation forms a tooth in dorsal view (Fig. 46D). Internal sac of aedeagus with a somewhat pentagonal main sclerite. Female. Unknown. Distribution. The species in known from a single specimen collected in the Masoala Peninsula (Fig. 46F). Etymology. From Latin, “inexpectatus” for unexpected. This name is an adjective in the nominative singular.

Acknowledgments

We are thankful to the curators of the collections who provided access to specimens. We especially thankful to Thierry Deuve, Antoine Mantilleri, Stéphane Boucher, and Azadeh Taghavian for assistance to A.V.F. and L.A.A. while working at the MNHN. We are thankful to the late Ilkka Hanski and Heidi Viljanen (University of Helsinki, Finland) for sharing the Orphninae material and information gathered by them in Madagascar. We thank the two anonymous reviewers and Andrew Smith for their valuable comments. This project was partly supported by the Russian state research project 01201351189 and Russian Foundation for Basic Research (grants 13-04-01002-a, 16-04-00412-a).

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