Chimarra of Sabah and Sarawak, Northern Borneo (Trichoptera: Philopotamidae)
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Chimarra of Sabah and Sarawak, northern Borneo (Trichoptera: Philopotamidae) Roger J. Blahnik, Ralph W. Holzenthal & Jolanda Huisman Thirty new species of Chimarra Stephens (Trichoptera: Philopotamidae) from the Malaysian part of Borneo (Sabah and Sarawak) are described, and eight previously described species are redescribed (Chimarra dulitensis Kimmins, 1955 and C. demeter Malicky, 2000, originally described from the Malaysian part of Borneo; and also C. phlegyas Malicky, 2008; C. polyneikes Malicky, 2008; C. prokrustes Malicky, 2008; C. terramater Malicky, 2008; C. thaumas Malicky, 2008; and C. tityos Malicky, 2008, originally described from the Indonesian part of Borneo, but now also recorded from the Malaysian part of Borneo). New species described include: Chimarra antheae, C. caduca, C. chanchuluni, C. cuspidata, C. cygnus, C. danumensis, C. dejongi, C. denticula, C. devogeli, C. drepane, C. fuilianae, C. gyrospina, C. jannekae, C. karlijnae, C. kinabaluensis, C. lambi, C. liwaguensis, C. noloyan, C. noohi, C. phillipsae, C. physanoton, C. preapicalis, C. scolops, C. silausilau, C. sinitorum, C. stenodactylus, C. vantoli, C. vanwelzeni, C. ventritropis, and C. xiphosella. Two major groups, the Chimarra tsudai group and the Chimarra digitata group are recognized, based on characters of the male genitalia and wing venation, to include most of these species. Roger J. Blahnik, Department of Entomology, 1980 Folwell Ave., 219 Hodson Hall, University of Minnesota, St. Paul, Minnesota, 55108, U.S.A. [email protected] Ralph W. Holzenthal*, same address, [email protected] Jolanda Huisman, same address, [email protected] Introduction C. tawitawi also occur in the northern, Malaysian Currently 229 species of Chimarra are known from part of Borneo. This paper thus redescribes and reil- the Oriental region, of the approximately 630 spe- lustrates all of the species currently known from cies known for the world. The vast majority of Borneo, excluding only C. atripennis, which is based these have been described in just the last 20 years. on an unassociated female holotype, and C. thyestes Until the recent paper by Malicky (2008), only and C. tawitawi which have not yet been collected three species were previously described from Bor- from the Malaysian part of Borneo. It includes an neo: Chimarra atripennis Banks, 1931, C. duliten- additional 30 new species from the Malaysian states sis Kimmins, 1955 and C. demeter Malicky, 2000. of Sabah and Sarawak, bringing to 40 the number The seven additional species recently described by of species now recorded from Borneo, all of them Malicky from the Indonesian part of Borneo (East- endemic, except C. dulitensis, which has also been ern Kalimantan) include C. phlegyas, C. polyneikes, reported from Sumatra by Malicky (1989), and C. prokrustes, C. terramater, C. thaumas, C. thyestes, C. tawitawi, otherwise only known from the Phil- and C. tityos. Chimarra tawitawi Malicky, originally ippines. It should be noted, however, that one of described from the Philippines (Malicky 1994), was the new species described here (C. devogeli, sp. n.) also reported from Kalimantan by Malicky (2008). is very similar to C. tawitawi, and it is possible that All of the described species, except C. thyestes and the record of C. tawitawi from Kalimantan may Tijdschrift voor Entomologie 152: 109–166, Figs 1–46. [ISSN 0040–7496]. http://www.nev.nl/tve © 2009 Nederlandse Entomologische Vereniging. Published 1 August 2009. * corresponding author 110 Tijdschrift voor Entomologie, volume 152, 2009 actually be C. devogeli. Despite the known diversity of The material was predominantly collected with light Chimarra in the Oriental region, the overall number traps and preserved in 70% ethyl alcohol, transferred of species will undoubtedly expand considerably in to 80% ethyl alcohol on final curation. Some mate- the future, particularly with the description of the rial from the collections of the NMNH and MNHU fauna from India and other under-collected regions, is preserved on pins. Material collected by Wolfram including the Indonesian part of Borneo. The species Mey and deposited at the MNHU was identified by from northern Borneo are, in general, most similar him based on illustrations provided by the authors, and undoubtedly most closely related to species from but was not personally examined by the authors. Southeast Asia, particularly those recently described It was added to the type material to extend the distri- from Thailand and Vietnam (Chantaramongkol & butional records for the species and to make known Malicky 1989; Malicky & Chantaramongkol 1993a, the availability of this material for subsequent asso- 1993b, 1997, 2003; Malicky 1994, 1995; Malicky ciative and taxonomic work. & Prommi 2006; Malicky et al. 2000, 2005, 2004, 2006; Mey 1998a, b). As discussed below, most of these fall into two distinctive species groups, the Systematics Chimarra tsudai group and the Chimarra digitata Most of the species of Chimarra from Sabah can group, based on characters of the male genitalia and be divided into two distinctive groups. These cor- wing venation. respond to “lineages” discussed by Ross (1956), and Most of the material studied for this research was col- characterized by the species C. digitata Martynov and lected by Jolanda Huisman between June, 1986 and C. tsudai Ross. Both groups, as is typical of the species May, 1987 in Sabah, Sarawak, and Brunei, during currently placed in the subgenus Chimarra, are char- which time some 200 collection sites were visited. acterized by male genitalia with tergum X divided Habitats from which these collections were made and widely separated mesally into paired, sclerotized ranged from lowland to montane (50–3300 m) lateral lobes (Blahnik 1998, fig. 9A-B). Both groups primary forest; Chimarra species occurred between are widespread and species diverse in Asia, with 50–1900 m. members of the digitata group also occurring in the New World, Africa, and extending into some Pacific islands and eastern Australia. Additional “lineages” Materials and methods were also recognized by Ross (1956), including his Most specimens examined were cleared in 10–12% Vigarrha and Chimarrhafra lineages from the Philip- KOH, overnight at room temperature; additional pines and Africa, respectively, based on genera that specimens were cleared in lactic acid according to the Ross synonymized under Chimarra. Blahnik (1998) method described by Blahnik & Holzenthal (2004) provided evidence that all of these Old World species and Blahnik et al. (2007). Specimens were examined groups belong in the subgenus Chimarra. However, and illustrated using an Olympus SZX12 stereomi- some species in the subgenus Chimarra, including croscope at magnifications up to 200. Drawings three included in this paper, are not readily placed in were made by use of a 1010 mm ocular grid and any of these groups. These may either be primitive or inked on a light table. Terminology for genitalia aberrant species within the major groups discussed and wings follows that used by Blahnik (1998) for here, or representatives of additional species groups. Chimarra. Female identifications are based on their Two of these species, C. polyneikes Malicky and co-occurrence with males and similarity in size or col- C. noohi, sp. n., lack a curved and inflected stem of oration; they are included in the type series because of the Rs vein of the forewing and also have the s, r-m, the value of having presumptively associated material and m crossveins of the forewing linearly aligned and for future studies. The word Sungai (abbreviated Sg.) hyaline (Figs 44A, 45A). Based on this set of wing used in the locality data is the Malay word for a river venational characters, these two species are similar or stream. Holotypes are deposited in the Nation- to species in Ross’s Chimarrhafra group of the sub- aal Natuurhistorisch Museum Naturalis (formerly genus Chimarra, and also to species in the genus Rijksmuseum van Natuurlijke Historie), Leiden, Chimarrhodella Lestage and the New World endemic The Netherlands (RMNH). Paratypes are depos- Chimarra subgenera, Curgia Walker, Otarrha Blah- ited in the same institution and also in the collec- nik, and Chimarrita Blahnik. However, like other tions of the University of Minnesota, St. Paul, Min- species in the subgenus Chimarra, the two new spe- nesota, USA (UMSP), the United States National cies from Borneo have tergum X divided into widely Museum of Natural History, Washington, DC, separated lateral lobes. This character evidence sug- USA (NMNH), and the Museum für Naturkunde, gests that these species may be basal or primitive Humbolt-Universität, Berlin, Germany (MNHU). species within the subgenus Chimarra, and hence Blahnik et al.: Chimarra of northern Borneo 111 of special interest because of their value for infer- subgroups, but this will undoubtedly require a com- ring phylogenetic relationships within the subge- parative study of all of the Asian species as a prelimi- nus. Various species from other geographical regions nary step. undoubtedly also have primitive venational charac- teristics, but this information is not always included Chimarra digitata group in species descriptions. The tsudai group, as recognized here, is well-defined Ross (1956) used C. digitata to characterize a lineage and the species belonging to it are generally easy to of Chimarra with a character set that included, in the discern from literature descriptions. It is given for- male genitalia, exactly two sensilla on the lateral lobes mal recognition here as the Chimarra tsudai group. of tergum X and an apically divided membranous Species belonging to this group are listed in Table 1 mesal lobe. He proposed that a member of this line- and the characters defining it are discussed below. age gave rise to the Chimarra aterrima lineage (New The membership of the digitata group is more diffi- World members of the subgenus Chimarra). Blah- cult to define, in part because the characters defining nik (1998) discussed additional characters applica- it are not always easy to assess from species descrip- ble to New World species, some of which also apply tions. Thus, although we discuss the general charac- to Old World species, including C.