Moss-Inhabiting Flea Beetles (Coleoptera: Chrysomelidae: Galerucinae: Alticini) with Description of a New Genus from Cangshan, C

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Journal of Natural History Publication details, including instructions for authors and subscription information: http://www.tandfonline.com/loi/tnah20 Moss-inhabiting flea beetles (Coleoptera: Chrysomelidae: Galerucinae: Alticini) with description of a new genus from Cangshan, China Alexander Konstantinova, Maria Lourdes Chamorroa, K. D. Prathapanb, Si-Qin Gec & Xing-Ke Yangc a Systematic Entomology Laboratory, USDA, ARS, c/o Smithsonian Institution, National Museum of Natural History, Washington, DC, USA b Department of Entomology, Kerala Agricultural University, Trivandrum, Kerala, India c Key Laboratory of Zoological Systematics and Evolution, Institute of Zoology, Chinese Academy of Sciences, Beijing, China Published online: 03 May 2013.

To cite this article: Alexander Konstantinov, Maria Lourdes Chamorro, K. D. Prathapan, Si-Qin Ge & Xing-Ke Yang (2013) Moss-inhabiting flea beetles (Coleoptera: Chrysomelidae: Galerucinae: Alticini) with description of a new genus from Cangshan, China, Journal of Natural History, 47:37-38, 2459-2477, DOI: 10.1080/00222933.2012.763068 To link to this article: http://dx.doi.org/10.1080/00222933.2012.763068

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Moss-inhabiting ﬂea beetles (Coleoptera: Chrysomelidae: Galerucinae: Alticini) with description of a new genus from Cangshan, China Alexander Konstantinova*, Maria Lourdes Chamorroa , K.D. Prathapanb , Si-Qin Gec and Xing-Ke Yangc

aSystematic Entomology Laboratory, USDA, ARS, c/o Smithsonian Institution, National Museum of Natural History, Washington, DC, USA; bDepartment of Entomology, Kerala Agricultural University, Trivandrum, Kerala, India; cKey Laboratory of Zoological Systematics and Evolution, Institute of Zoology, Chinese Academy of Sciences, Beijing, China

(Received 14 September 2012; ﬁnal version received 23 November 2012; ﬁrst published online 3 May 2013)

The world diversity of moss cushion-inhabiting and moss-feeding flea beetles is documented and discussed. A new genus (Cangshanaltica) with a single new species (Cangshanaltica nigra sp. nov.) from Yunnan Province in China is described and illustrated. It is similar to Benedictus Scherer, Ivalia Jacoby, Minota Weise, Paraminota Scherer, and Phaelota Jacoby. An identification key for all flea beetle genera known to occur in mosses in the eastern hemisphere is provided. http://www.zoobank.org/urn:lsid:zoobank.org:pub:6341724E-BA3F-4979-B226- 108D0CA64B92 Keywords: moss; bryobionts; new genus; species; flea beetles; Asia

Introduction Moss (bryophyte) cushions constitute a special environment characterized by several important features: the perennial nature of most bryophytes, rendering them present as a food source and a habitat for overwintering when many tracheophytes are absent; buffered temperature and moisture; and the small size of spaces among the stems and leaves within cushions (Glime 2006). Because of these features moss cushions are populated by a particular fauna of phytophagous, saprophagous and predatory invertebrates. Various terms have been suggested for animals associated with moss cushions. Chernov (1985) called invertebrates living in mosses semi-edophores (which

Downloaded by [Institute of Zoology] at 18:00 19 December 2013 means partly living in soil). Gerson (1982) suggested dividing moss faunae into: bryobionts (animals living exclusively within mosses); bryophiles (animals that are usually associated with mosses but can be found elsewhere); bryoxenes (animals spending part of their life cycle within mosses); and occasionals (animals that at times are found in mosses but do not depend on them for survival) (Glime 2006). Leaf beetles (Chrysomelidae) constitute a relatively small portion of moss-living invertebrates. Adults of the vast majority of the more than 50 000 known species of leaf beetles live on the leaf surface of various ﬂowering plants on which they feed. To date, only 27 leaf beetle species from 14 genera (Table 1) are documented to live within moss cushions (bryobionts). All moss-inhabiting leaf beetles belong to the tribe Alticini, a group of about 12 000 species in the world. Nineteen species from nine

*Corresponding author. Email: [email protected]

© 2013 Taylor & Francis 2460 A. Konstantinov et al. ) (2005) Continued (2009) (2009) (2009) (2009) (2009) (2009) (2009) ( Konstantinova (2011) et al. et al. et al. et al. et al. et al. et al. (2006) Chamorro- Lacayo (2006) Duckett Konstantinov and Sprecher-Uebersax Sprecher-Uebersax Sprecher-Uebersax Sprecher-Uebersax Sprecher-Uebersax Sprecher-Uebersax Konstantinov and Konstantinov and Duckett et al. sp. Mitt. sp. (Hypnaceae) Host plants Citation (Hypnaceae) Hypnum Isopterygium (m) 1066 3478 3443 3667 2863 3850 3892 3000 3300 Sprecher-Uebersax 1270 3300 2300 forest forest forest forest forest forest forest forest forest forest forest plantation premontane wet mixed montane Rhododendron Rhododendron Rhododendron Rhododendron Rhododendron Rhododendron Rhododendron premontane wet Rhododendron pine trees in tea Rico, El Yunque Cangshan mountains, Yatong La pass, pass Nobding National Park. Chilal La pass, Thimphu, Dochu La pass, Cangshan mountains, Dominican Republic, Sierra de Baoruco Cangshan mountains, Ghats, Tamil Nadu eastern China, Yunnan, eastern Bhutan, Tongsa dist, eastern Bhutan, Thumsi La eastern Bhutan, env. of eastern Nepal, Lantang eastern Bhutan, env. of Ha, eastern Bhutan, env. of eastern China, Yunnan, eastern China, Yunnan, eastern South India, Western western West Indies, Puerto western West Indies, Downloaded by [Institute of Zoology] at 18:00 19 December 2013 Konstantinova, (2011) Chamorro, Prathapan, Ge, Yang, sp. nov. et al., (2009) et al., (2009) et al., (2009) et al., (2009) et al., (2009) et al., (2009) et al., (2009) Chamorro-Lacayo, (2006) Duckett, (2005) (2006) onstantinov and K Konstantinov and Konstantinov, Sprecher-Uebersax Sprecher-Uebersax Sprecher-Uebersax Sprecher-Uebersax Sprecher-Uebersax Sprecher-Uebersax Sprecher-Uebersax Konstantinov and Prathapan et al., dali elyunque nigra yatongla thumsila nobding lauribina dochula cangshanicus baorucae korakundah Clavicornaltica Borinken Cangshanaltica Benedictus Benedictus Benedictus Benedictus Benedictus ha Benedictus Taxon nameBenedictus Author(s) and date Hemisphere Geographic range Habitat Altitude Kiskeya Ivalia Table 1. Diversity of moss inhabiting ﬂea beetles of the world. Journal of Natural History 2461 ) (2009) Continued ( (1997) Konstantinova (2011) Chamorro- Lacayo (2006) Cox Konstantinov and Unpublished data Nadein sp. sp. Hypnum A. rugelii sp. Hedw. (Hedw.) (Hedw.) B. S. sp. Neckera crispa Eurhynchium Brachythecium (Lindb. ex Limpr.) Warnst. (Hypnaceae) and Schimp. (Brachythecieae) (Hedw.) Mitt., cupressiforme (Hypnaceae); Thamnobyum alopecurum Nieuwl., Hedw. (Neckeraceae); Metzgeria conjugata Lindb. (Metzgeriaceae); Anomodon attenuatus (Hedw.) Hub., (Mull.) Keisl. (Anomodontaceae); Brachythecium populeum G., (Brachytheciaceae); Fissidens (Fissidentaceae) Rhytidiadelphus Ctenidium molluscum 10661597 1010 Konstantinov and 200–1900 400–1500 including pine and junipers forest forest leaved forest Various forests, premontane wet premontane wet Europe, from Great Britain to Ukraine Rico, El Yunque Dominican Republic, Sierra de Baoruco Caucasus eastern Russia, Western eastern Southern and Central eastern Japan, Kyushu Isl. montane broad western West Indies, Puerto western West Indies, Downloaded by [Institute of Zoology] at 18:00 19 December 2013 Konstantinova, (2011) Chamorro-Lacayo, (2006) Koch, (1803) Konstantinov and Konstantinov and Nadein, (2009) sp. muscorum elyunque caucasica Mniophila Table 1. (Continued). Kiskeya Kiskeya neibae Minota Mniophila 2462 A. Konstantinov et al. ) (2009) 2009a Konstantinov (2009) Konstantinov (2009) Konstantinova (2011) ( (2002) Konstantinov (2003) Konstantinov (2009) Prathapan and Konstantinov and Konstantinov et al. Konstantinov Döberl and Prathapan and Prathapan and Unpublished data Nadein Unpublished data (Spruce) Host plants Citation (Mitt.) W.R. Buck (Leucodontaceae) (Beuch ex Spruce) Schimp., Homalothecium philippeanum Schimp. (Brachytheciaceae), and Plagiomnium rostratum (Schrad.) T. Kop. (Mniaceae) Forsstroemia thomsonii Brachythecium glareosum 793 571 (m) 1993 1066 1450 3850 3500 2060 2600 100–800 temperate forest forest forest forest forest temperate forest temperate forest montane wet premontane wet montane wet Rhododendron Rhododendron montane wet montane wet Ghats, Tamil Nadu Rico, El Yunque Nicaragua, Serrania Dariense National Park National Park. Ghats, Tamil Nadu Ghats, Tamil Nadu eastern Nepal, Lantang eastern South India, Western eastern Ukraine, Crimea eastern South India, Western eastern South India, Western western West Indies, Puerto westernwestern West Indies, St Lucia Central America, moss western West Indies, St Lucia tree moss Downloaded by [Institute of Zoology] at 18:00 19 December 2013 ) 2009a Konstantinov, (2009) Konstantinova, (2011) ( Konstantinov, (2009) Konstantinov, (2009) Prathapan and Konstantinov and Konstantinov et al., Konstantinov, (2002)Döberl, (1991) eastern Nepal, Lantang Prathapan and Prathapan and Nadein, (2009) sp. 2 sp. 1 viridipennis selvanegra lauribina nepalensis maculipennis taurica Phaelota Ulrica eltoro Monotalla Nicaltica Paraminota Paraminotella Phaelota Phaelota saluki Table 1. (Continued). Taxon nameMniophila Author(s) and date Hemisphere Geographic range HabitatMonotalla Altitude Journal of Natural History 2463

genera are known in the eastern hemisphere and eight species from five genera in the western hemisphere (Table 1). All moss-inhabiting flea beetles share a similar habitus. They are among the smallest leaf beetles, with a round body, robust appendages and antennae with more or less enlarged apical antennomeres, reduced hind wings, greatly simplified and shortened mesothorax and metathorax, and the elytra lacking humeral calli. Little is known about the biology of moss-inhabiting flea beetles. It is particularly difficult to identify what these beetles are feeding on because usually they are not observed feeding, their larvae are mostly unknown [larvae are described for only Ivalia korakundah Prathapan, Konstantinov and Duckett (Duckett et al. 2006)and Mniophila muscorum (Koch) (Cox 1997)], and DNA investigation of their gut contents has not been attempted. Host plants are documented for only six bryobiont species (Table 1) but it is unclear how the host plant relationships were established for Mniophila caucasica Nadein and Mniophila taurica Nadein. Most likely these species were living within cushions consisting of the mosses in question (Table 1) (Nadein 2009). The fact of feeding still needs to be confirmed either by observation of adult or larval feeding, or by identification of the gut contents. It should be noted that actual feeding was observed only for M. muscorum (Cox 1997) and an undescribed species of Ivalia Jacoby (Prathapan unpublished data). The gut contents containing cell samples of a moss from the genus Hypnum Hedw. (Hypnaceae) provided evidence for moss feeding in Cangshanaltica nigra sp. nov. (Figure 6G). However, the fact that M. muscorum associated with mosses from differ- ent families and the range of potential hosts for M. caucasica and M. taurica suggests that moss-feeding flea beetles are most likely polyphagous. It is also possible that some of them are detritophagous as many congeners of these beetles live in leaf litter where detritus is the likely food source. Habitats of moss-inhabiting flea beetles are also poorly known. Species of Benedictus Scherer, Paraminota Scherer and Paraminotella Döberl and Konstantinov were collected in alpine Himalayan forests (2800–4400 m) by moss sifting (Konstantinov 2002; Döberl and Konstantinov 2003; Sprecher-Uebersax et al. 2009). Mosses are abundant in high-altitude forests, especially where species of Rhododendron L. (Ericaceae) are dominant. Moss cushions covering tree trunks, rocks, soil surface and even hanging from the branches of trees comprise the habitat of these flea beetles in the Himalayas. In southern India, where the highest peak attains an altitude of only 2965 m, mosses are comparatively less abundant. Moss dwelling apterous

Downloaded by [Institute of Zoology] at 18:00 19 December 2013 species of Phaelota Jacoby and Ivalia were collected in the humid environments of the tropical evergreen forests of the Western Ghats chain of mountains at altitudes ranging from 1000 to 2600 m (Duckett et al. 2006; Prathapan and Konstantinov 2009; Prathapan unpublished data). However, an undescribed species of alate Ivalia occurs from sea level to 2600 m (Duckett et al. 2006; Prathapan unpublished data). This species was collected from moss in a variety of habitats ranging from urban environments to montane rain forests. Being endowed with the capacity of ﬂight, this beetle has adapted well to a variety of habitats. A single specimen of this species was also picked up in artiﬁcial light. Borinken, Konstantinov and Konstantinova, Kiskeya Konstantinov and Chamorro-Lacayo, Nicaltica Konstantinov, Chamorro-Lacayo and Savini, and Ulrica Scherer occur in lower mountains (1200–1500 m) in the West Indies and Central America (Konstantinov and Chamorro-Lacayo 2006; Konstantinov et al. 2009a; Konstantinov and Konstantinova 2011). Clavicornaltica dali Konstantinov 2464 A. Konstantinov et al.

and Duckett was collected in moss under Rhododendron trees at 3300 m in China (Konstantinov and Duckett 2005). Mniophila species in the Crimea and the Caucasus occur between 400 and 1500 m (Nadein 2009). The diversity of moss-inhabiting flea beetles is vastly understudied; every success- ful moss-sifting event in a previously unexplored locality uncovers 100% of new species and about 30% of new genera. Moss sifting, surprisingly, is not a popular method of beetle collecting compared with leaf litter sifting. The latter method has revealed a much more diverse flea beetle fauna around the world with many genera that occur both in leaf litter and moss cushions. Among the 14 flea beetle genera occurring in mosses, six are bryobionts (Kiskeya, Borinken, Cangshanaltica, Mniophila, Nicaltica and Ulrica) and eight are bryophiles (Benedictus, Clavicornaltica, Ivalia, Monotalla, Minota, Paraminota, Paraminotella and Phaelota). Here we provide a list of flea beetles documented to occur in moss cushions in the western and eastern hemispheres (Table 1) and describe Cangshanaltica nigra sp. nov.

Materials and methods Beetles were retrieved from moss samples by Berlese extraction. Initially, moss samples collected in the ﬁeld were placed directly in Berlese funnels. However, by the end of the collecting trip it became clear that there was not enough time to process all the collected moss through Berlese funnels. Therefore, moss samples were sifted, which dramatically reduced the size of the remaining substrate. It is still unclear if sifting causes loss of beetle specimens. The food source of Cangshanaltica nigra sp. nov. was identiﬁed based on the gut contents, which included undigested cells that were recognized as belonging to mosses from the genus Hypnum. The gut contents included the tip and base of at least two leaves. Dissecting techniques, measurements and terminology follow Konstantinov (1998). Scanning electron microscopy images were taken with a Hitachi TM 3000 Tabletop Microscope. Observations were made with a Zeiss Discovery V20 microscope and digital images were taken with an AxioCam HRC digital camera attached to it. Specimens are deposited in the National Museum of Natural History, Smithsonian Institution, Washington DC, USA (USNM) and the Institute of Zoology, Chinese Academy of Sciences, Beijing, China (IZAC). Downloaded by [Institute of Zoology] at 18:00 19 December 2013 Cangshanaltica Konstantinov, Chamorro, Prathapan, Ge, and Yang, gen. nov. (Figures 1–6, 9)

Description Body length 1.62–1.77 mm, width 1.25–1.37 mm, rounded (Figure 1), convex in lateral view (1.66 times as long as thick). Colour black without metallic lustre, legs and antennae light chestnut brown. Head (Figure 2A–D) nearly hypognathous, only slightly convex in lateral view. Frons and vertex forming slightly convex line (Figure 2D) in lateral view. Supraorbital pore well developed, placed close to supraorbital sulcus, with no other pores nearby. Antennal calli poorly developed, slightly shorter than wide, straight, connected to Journal of Natural History 2465

Figure 1. Cangshanaltica nigra sp. nov., habitus.

each other. Supracallinal, midfrontal and suprafrontal sulci shallow, barely visible. Supraorbital sulcus deep, straight. Orbit narrow, 0.50 times as wide as transverse diam-

Downloaded by [Institute of Zoology] at 18:00 19 December 2013 eter of eye. Interantennal space 2.10 times wider than transverse diameter of eye and 3.0 times as wide as transverse diameter of antennal socket. Frontal ridge wide, as wide as anterofrontal ridge, occupying nearly all frontal side of head, with parallel sides. Anterofrontal ridge not separated from frontal ridge. Both ridges form swollen, loosely defined rectangle sparsely covered with shallow, poorly defined, small punctures. Eyes slightly protruding laterally. Vertex covered with shallow, poorly defined, small punctures. Labrum with six setiferous pores, apically deeply incised. Labium (Figure 2C) with three small palpomeres per palpus, two distal palpomeres longer than wide. Maxillary palpus with four palpomeres (Figure 2F), distal palpomere conical, much shorter and narrower than preapical. Antenna with 11 antennomeres. First antennomere slightly wider and much longer than second. Third and fourth antennomeres much thinner than second. Distal antennomeres wider than third and fourth. Antennomere seven with distal protrusion directed anteriorly (Figure 2G). 2466 A. Konstantinov et al. Downloaded by [Institute of Zoology] at 18:00 19 December 2013

Figure 2. Scanning electron micrographs of Cangshanaltica nigra sp. nov. (A) Head, ventrolat- eral view; (B) head and prothorax, ventral view; (C) mouthparts, ventral view; (D) head, lateral view; (E) last maxillary palpomere; (F) maxillary palpi; (G) antennomeres 6–9. Journal of Natural History 2467

Figure 3. Scanning electron micrographs of Cangshanaltica nigra sp. nov. (A) Thorax and abdomen, ventral view; (B) thorax, lateral view; (C) pronotum, lateral view; (D) metasternum. Downloaded by [Institute of Zoology] at 18:00 19 December 2013 Pronotum (Figure 1) 1.74 times wider than long (measured in middle), without impressions basally, covered with sharp, small, very sparsely placed punctures. Sides weakly rounded and relatively narrowly explanate, converging gradually from base to apex. Marginal anterolateral callosity poorly developed. Its setiferous pore placed nearly in middle of lateral margin of pronotum (Figure 3C). Posterolateral callosity slightly protruding. Basal margin extends posteriorly, with thin border. Pronotal side near anterior corner with shallow S-shaped impression separating low and wide callosity. Procoxal cavity open. Intercoxal prosternal process relatively narrow, with sharp longitudinal ridge, slightly widening apically, slightly convex at apex (Figure 2B). Mesoscutellum ﬂat, broadly triangular (Figure 1). Mesocoxae separated by both mesosterna and metasterna. Mesosternum not covered by metasternum, partly horizontal. Intercoxal mesosternal process much wider than long (Figure 3A, B). 2468 A. Konstantinov et al.

Downloaded by [Institute of Zoology] at 18:00 19 December 2013 Figure 4. Scanning electron micrographs of Cangshanaltica nigra sp.nov.(A)Foreleg;(B) middle leg; (C) metatarsus; (D) hind leg.

Metasternum (Figure 3D) very short between mesocoxae and metacoxae, protruding anteriorly between mesocoxae, broadly rounded at apex anteriorly, with narrow rounded emargination posteriorly. Mid part of metasternum with ﬂat area separated anteriorly and laterally by distinct ridge. Elytron (Figure 1) widest near mid-length. Humeral callus lacking. Elytral punctures small, widely separated (diameter of punctures many times smaller than distance between them) and not arranged in regular rows. Elytral apex narrowly rounded, surrounded by distinct border. Epipleura broad, horizontal, gradually narrowing posteriorly, nearly attaining sutural margin of elytron. Elytral lock at base of elytra Journal of Natural History 2469

Figure 5. Scanning electron micrographs of Cangshanaltica nigra sp. nov. (A) Protarsus, ventral view; (B) mesotarsus, ventral view; (C) apex of right elytron; (D) apex of left elytron; (E) base Downloaded by [Institute of Zoology] at 18:00 19 December 2013 of right elytron with elytral lock; (F) sensillar patch on ventral side of right elytron; (G) base of left elytron with elytral lock.

consists of three ridges including one in middle on right elytron (Figure 5E). In left elytron middle ridge narrow, groove above it relatively wide and deep (Figure 5G). Ventral surface of each elytron with one sensilla patch (Figure 5F). Hind wings absent. Profemur and mesofemur slightly flattened dorsoventrally (Figure 4A, B). Metafemur robust, flat dorsoventrally, fairly symmetrical (Figure 3A), 1.84 times as long as wide. Protibia and mesotibia generally cylindrical, slightly wider in distal third, with sharp longitudinal ridge on outer surface, with spurs apically. Metatibia (Figure 4D) slightly curved in dorsal and lateral views, generally cylindrical, abruptly widening distally (in dorsal view), flat only at apex, with longitudinal ridge dorsally, without serration on outer and inner edges and without preapical excavations on either 2470 A. Konstantinov et al. Downloaded by [Institute of Zoology] at 18:00 19 December 2013

Figure 6. Abdomen, male and female genitalia of Cangshanaltica nigra sp. nov. (A) Aedeagus, ventral and lateral view; (B) spermatheca; (C) vaginal palpus; (D) abdominal tergites; (E) abdominal ventrites; (F) tignum; (G) gut with its content, including cells of moss of the genus Hypnum, indicated by arrows.

inner or outer edges; apical spur straight and long (Figure 4C), longer than tarsal claw. Claw appendiculate (Figure 5A). Third tarsomere narrow, deeply incised (Figure 4D). First metatarsomere as long as two following tarsomeres together. Abdomen with ﬁve distinctly visible ventrites (Figure 3A). First ventrite as long as two preceding sternites combined, without appendages basally, with longitudinal Journal of Natural History 2471 Downloaded by [Institute of Zoology] at 18:00 19 December 2013

Figure 7. Habiti of ﬂea beetles. (A) Benedictus shivalayanicus Sprecher-Uebersax, Konstantinov, Prathapan and Döberl; (B) Clavicornaltica dali Konstantinov and Duckett; (C) Ivalia korakundah Prathapan, Konstantinov and Duckett; (D) Minota obesa (Waltl). 2472 A. Konstantinov et al. Downloaded by [Institute of Zoology] at 18:00 19 December 2013

Figure 8. Habiti of ﬂea beetles. (A) Mniophila muscorum (Koch); (B) Paraminota lauribina Konstantinov; (C) Paraminotella nigrita Döberl and Konstantinov; (D) Phaelota mauliki Prathapan and Konstantinov.

ridge in middle. Apical tergite of female broadly triangular, unevenly covered with long setae, sclerotized along its posterior margin and in middle, without groove in middle (Figure 6D). Spermatheca (Figure 6B) with receptacle gradually transitioning into pump without distinct border. Receptacle much longer than wide and wider and longer than Journal of Natural History 2473 Downloaded by [Institute of Zoology] at 18:00 19 December 2013

Figure 9. Habitats of Cangshanaltica nigra sp. nov. (A) Cangshan mountains; (B) moss cushion.

pump. Duct straight distally, basally directed away from receptacle, with gland situ- ated at its distal end. Tignum (Figure 6F) anteriorly slightly narrower than posteriorly. Vaginal palpi (Figure 6C) robust, connected at base, making relatively wide loop, distal sclerotizations narrow and straight. 2474 A. Konstantinov et al.

Median lobe of aedeagus (Figure 6A) simple, convex in lateral view, ﬂat dorsoventrally, without any sculpture ventrally.

Type species Cangshanaltica nigra Konstantinov, Chamorro, Prathapan, Ge, and Yang.

Etymology This genus name is derived from Cangshan, the mountain region in western Yunnan (China), where these beetles occur, and the name of the type genus of the tribe (Altica) to which the genus belongs. The name is feminine.

Host plant Hypnum sp. (Hypnaceae) (Figures 9A, B).

Discussion A few species of Benedictus, Ivalia, Paraminota, Phaelota and Paraminotella are known to occur inside moss cushions or leaf litter in various regions of Asia. These species are characterized by a generally small and round body, and the absence of wings, so they resemble Cangshanaltica, but all of them can be easily separated with the key at the end of this paper. Among Oriental ﬂea beetle genera, Cangshanaltica is particularly similar to Phaelota and Ivalia. They share about the same body size and shape, major head features with wide frontal ridge, lacking well-developed antennal calli, pronotum without antebasal transverse and longitudinal furrows, and spermatheca without distinct border between receptacle and pump. Cangshanaltica can be separated from Phaelota based on the following features: head without supracallinal sulci; pronotal side near anterior corner with shallow S-shaped impression separating low and wide callosity; procoxal cavity posteriorly open; epipleura nearly horizontal; metatarsal spur as long as metatarsal claw; vaginal palpi widely connected to each other. Cangshanaltica can be separated from Ivalia with the help of the following features: head without suprafrontal sulci; antennomere 7 with distal protrusion directed anteriorly, prono- Downloaded by [Institute of Zoology] at 18:00 19 December 2013 tal side near anterior corner with shallow S-shaped impression separating low and wide callosity; pronotal anterolateral setiferous pore placed almost in middle of lateral margin; metatibia only slightly curved in dorsal view, with dorsal surface convex.

Cangshanaltica nigra Konstantinov, Chamorro, Prathapan, Ge, and Yang, sp. nov. (Figures 1–6, 9)

Description Body length 1.62–1.77 mm, width 1.25–1.37 mm. Colour black without metallic lustre, legs and antennae light chestnut brown, mandibles, apical antennomeres, and coxae and trochanters (especially metatrochanters) dark yellow. Journal of Natural History 2475

Vertex smooth, with tiny punctures and without wrinkles, vertex punctures with relatively long setae. Proportions of antennomere lengths in female: 7 : 4 : 4 : 3 : 3 : 3 : 4 : 3.5 : 4 : 4 : 7; in male: 7 : 4 : 4 : 4 : 4.5 : 3 : 4.5 : 3.5 : 4 : 4 : 7. Pronotum sparsely covered with shallow, small punctures. Distance between punctures many times more than diameter of puncture. Elytron with punctures slightly smaller than those of pronotum. Interspaces uneven, slightly shagreened. Proportions of tarsomere lengths of female (starting with ﬁrst): protarsomeres 3.5 : 3 : 4.5 : 7; mesotarsomeres 4 : 3 : 4 : 7; metatarsomeres 9 : 4 : 5 : 7. In male, proportions as follows: protarsomeres 4:3:4:7;mesotarsomeres4:3:5:7;metatarsomeres9:4:5:7. Receptacle of spermatheca with external side concave, internal side convex (Figure 6B). Spermathecal pump much shorter than receptacle, slightly curved, apex with denticle. Posterior part of tignum (Figure 6F) short, weakly sclerotized, slightly wider than anterior part. Vaginal palpi (Figure 6C) twice as long as wide, with its entire length sclerotized and posterior sclerotizations evenly conical. Median lobe of aedeagus (Figure 6A) in lateral view with apex bent dorsally and in ventral view gradually narrowing without well-deﬁned denticle.

Comments Males and females of this species are nearly indistinguishable. Characters that usually help to separate males and females in many ﬂea beetles (e.g. width of the ﬁrst protar- somere and mesotarsomere, shape of the antennae and the apex of the last abdominal ventrite) are not helpful in C. nigra.

Etymology The speciﬁc name is derived from the Latin nigrum, referring to the black colour of the beetle.

Host plant Hypnum sp. (Hypnaceae) (Figures 9A, B). Type material: Holotype male. (1) China: Yunnan, env. Dali, Cangshan Mountains, 3478 m, 19 July 2011, 25◦41.100 N, 100◦06.228 E, moss, Konstantinov, Chamorro and Volkovitsh; (2) Holotype Cangshanaltica nigra Konstantinov, Downloaded by [Institute of Zoology] at 18:00 19 December 2013 Chamorro, Prathapan, Ge, Yang, sp. nov., 2011 (IZAC). Paratypes with same labels as holotype (2 females IZAC; 1 male, 2 females USNM).

Key for identiﬁcation of moss-inhabiting ﬂea beetle genera from the eastern hemisphere 1. Antennaeclavate...... Clavicornaltica Scherer (Figure 7B) Antennaenotclavate...... 2 2. Anteriorcoxalcavityclosed...... 3 Anteriorcoxalcavityopen...... 4 3. Fontalridgebetweenantennalsocketsmuchwiderthannearclypeus...... Minota Weise (Figure 7D) 2476 A. Konstantinov et al.

Fontalridgebetweenantennalsocketsmuchnarrowerthannearclypeus...... Phaelota Jacoby (Figure 8D) 4. Base of pronotum with transverse impression ...... Benedictus Scherer (Figure 7A) Base of pronotum without transverse impression ...... 5 5. First abdominal ventrite with one or two longitudinal ridges between metacoxae...... 6 First abdominal ventrite without longitudinal ridges between metacoxae . . . 7 6. Pronotal anterolateral setiferous pore placed nearly in middle of lateral margin. Antennomere seven with distal protrusion directed anteriorly. Head without suprafrontal sulci. Metatibia only slightly curved in dorsal view, with dorsalsurfaceconvex...... Cangshanaltica gen. nov. (Figure 1) Pronotal anterolateral setiferous pore placed close to anterior end of lateral margin. Antennomere seven without distal protrusion. Head with suprafrontal sulci. Metatibia strongly curved in dorsal view, with dorsal surfaceﬂattoconcave...... Ivalia Jacoby (Figure 7C) 7. Frontal ridge narrowing gradually from antennal sockets towards anterofrontal ridge. Supracallinal sulcus absent. First metatarsomere only slightly less than half-length of metatibia. Third metatarsomere entire ...... Paraminotella Döberl and Konstantinov (Figure 8C) Frontal ridge narrowing abruptly from antennal sockets towards anterofrontal ridge. Supracallinal sulcus present. First metatarsomere only much less than half length of metatibia. Third metatarsomere divided into two abruptly narrowing lobes ...... 8 8. Body rounded (Figure 8A). Eighth antennomere substantially smaller than seventh...... Mniophila Stephens (Figure 8A) Body more elongate (Figure 8B). Eighth antennomere about as large as seventh...... Paraminota Scherer (Figure 8B)

Acknowledgements We are grateful to D. Nelson (USDA, Forest Service, International Research Programs,

Downloaded by [Institute of Zoology] at 18:00 19 December 2013 Washington DC, USA) for partial funding for exploration in China. M. Volkovitsh (Zoological Institute, St Petersburg, Russia), Hongxia Xu (Institute of Zoology, Chinese Academy of Sciences, Beijing, China) and T. Petrice (USDA, Forest Service, Northern Research Station, MI, USA) are thanked for camaraderie and companionship during the 2011 explorations in China. M. Gates and A. Norrbom (Systematic Entomology Laboratory, Washington DC, USA), and C. Staines (Department of Entomology, Smithsonian Institution, Washington DC, USA) thoroughly reviewed earlier versions of this manuscript and provided valuable sugges- tions. We thank M. Bufﬁngton and M. Gates (Systematic Entomology Laboratory, Washington, DC) for their help with the digital cameras and image capture and editing. We are grateful to M. Nair (Department of Botany, Zamorin’s Guruvayoorappan College, Kozhikode and the Malabar Botanic Garden, Calicut, Kerala, India) for identiﬁcation of the moss samples from the gut of Cangshanaltica nigra. KDP is funded by the Indian Council of Agricultural Research Network Project on Insect Systematics and the Kerala State Council for Science, Technology and Environment, Trivandrum. Habitus illustration are the work of K. Arakawa (Ibaraki, Journal of Natural History 2477

Japan), A. N. Brown (USA), S. V. Edgerton (San Francisco CA), and E. Roberts (Systematic Entomology Laboratory, Washington DC, USA). USDA is an equal opportunity provider and employer.

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