Chimarra of Sabah and Sarawak, Northern Borneo (Trichoptera: Philopotamidae)

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

* 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 1010 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. digitata. These ters defining the group below, we have not attempted include well-developed anterodorsal apodemes on to provide a list of the included taxa. In a larger segment IX and the presence of two elongate, basal, sense, the digitata group undoubtedly falls within and usually asymmetrically arranged endothecal a more inclusive group of species within which the spines (varying from 1–3, rarely absent). Since there characters defining the digitata group emerged. This is no evidence that New World species in the subge- larger group includes the type species for the genus, nus Chimarra constitute a monophyletic group, the Chimarra marginata (Linnaeus, 1767). This is evi- Chimarra aterrima group is here subsumed within denced by the similar venational characters C. margi- the Chimarra digitata group. Venational charac- nata shares with the digitata group. However, among ters typifying this group, and also applicable to other characters, C. marginata lacks the pair of elon- C. marginata, include a sinuously inflected forewing gate endothecal spines that generally characterizes Rs vein, such that the basal fork of the discal cell the digitata group. This more comprehensive group is distinctly asymmetrical, a short discal cell (length is referred to here as the marginata group. Its recog- usually about 2 times width), and forewing 2A vein nition is useful in that it clearly defines the subgenus intersecting 3A, and with a crossvein to 1A vein Chimarra, in the event that additional subgenera are proximal to the intersection (making the apex of 2A eventually recognized from within the subgenus as appear forked or Y-shaped) (Fig. 40A). The descrip- currently defined. We have restricted the definition of tion below applies to most of species of the Chimarra the digitata group to species with the set of apomor- digitata group; individual species may vary in one or phies discussed by Ross (1956) for Chimarra digitata more characters. and by Blahnik (1998) for New World representatives of the subgenus Chimarra. Other characters are Adult. Forewing with pronounced, sinuous inflec- discussed below. Some of the other Asian species dis- tion of Rs, sometimes with enlarged node at point cussed by Ross, such as C. sepulchralis Hagen and of inflection, extending into cell below; base of discal C. circularis Hagen, share this same basic set of cell forked asymmetrically, usually distinctly thick- characters and could be considered members of this ened and enlarged; discal cell short (length usually group. We have retained C. digitata as a name to 2–2.5 times width); m crossvein proximal to s and refer to this group because it lacks distinctive sec- r-m crossveins, s not hyaline; hind wing 2A complete, ondary apomorphies and is more representative of crossvein to 1A present (apparently forked apically). the morphology of the group as a whole. New World Male foretarsi and claws enlarged or not. members of the Chimarra digitata group were placed Male genitalia (Figs 2, 7). Segment VIII not highly into a number of small, well-defined species groups modified, tergum often slightly longer than ster- (Blahnik 1998). These can be considered subgroups num, sternum sometimes with short ventral proc- of the digitata group, as defined here. Australian spe- ess. Segment IX with anteroventral margin at least cies of Chimarra, some of which can be placed in somewhat expanded, sometimes distinctly so, mesal the Chimarra digitata group, were also organized margin sinuously invaginated or broadly rounded; into informal species groups by Cartwright (2002). anterodorsally, with pair of distinctly projecting As discussed below, a number of species from Bor- apodemes; dorsal margin short, but present, con- neo belong to the digitata group. Ultimately, these tinuous posteriorly with projecting membranous will probably also be organized into a number of mesal lobe of tergum X; ventral process posteriorly 112 Tijdschrift voor Entomologie, volume 152, 2009

a b c

Fig. 1. Chimarra antheae, male genitalia. – a, lateral; b, segment IX and tergum X, dorsal; c, inferior appendage, ventral; d, phallic apparatus, lateral; inset, dorsal of phallotremal sclerite complex.

projecting, varying from short, triangular to long, papillate projections in most species. Phallotremal acute. Membranous mesal lobe of segment X usu- sclerite complex typically composed of simple rod ally divided apically, secondarily sclerotized in a few and ring structure, usually much shorter than either species, never with sensilla. Lateral lobes of tergum phallotheca or endotheca, sometimes with accompa- X sclerotized, widely separated, usually more or less nying dorsolateral or apicolateral sclerites. vertical in orientation; sensilla reduced to 2 (nearly consistently), often on projecting process. Preanal Chimarra antheae sp. n. appendages short, setose, often somewhat flattened. Fig. 1 Inferior appendages extremely variable in shape in different species, usually with strong basal inflection, Type material. Holotype: ?, Malaysia, Sabah, as viewed laterally, and long (or short) dorsal pro- Kinabalu National Park, Headquarters, roadside, jection, dorsal projection usually distinctly narrower 06°00’N, 116°32’E, 1500 m, 12.v.1987, J. Huis- than basal part of appendage; in ventral view, with man, (UMSP000208644) (RMNH). distinct mesal curvature, often with mesal ridge, Paratypes. Malaysia, Sabah, 3.5 km SW Kg. Long tooth, tubercle, or spine-like projection from mesal Pa Sia, Sg. Ritan, 04°24’N, 115°42’E, 1160 m, 8– surface. Phallobase typically with slight constriction 9.iv.1987, J. Huisman, 1?, 1/ (UMSP); Crocker near base, forming slight bulge or “belly”; ventral Range, 5 km N Tenom, base of Sg. Ulu Noloyan, apex usually projecting (at least slightly). Endotheca 05°10’N, 115°56’E, 1010 m, 10–11.x.1986, J. Huis- with 2 short to elongate endothecal spines, asym- man, 2? (UMSP); Kinabalu National Park, Head- metrically arranged near base of structure (nearly quarters area, 1560 m, 9.ix.1983, Hevel & Steiner, symmetrically arranged in a few species, reduced 1? (NMNH); Kinabalu National Park, Headquar- to one in others, or with a 3rd, much reduced and ters, Sg. Silau-Silau & Sungai Liwagu, 11.viii.2005, more apical spine in a few species); endotheca often W. Mey, 3? (MNHU); SW Sabah, nr. Long Pa Sia elongate and much retracted into phallobase when (west), 1050 m, 25.xi-8.xii.1987, Achterberg, 1? not inflated; apex textured, or with small spines or (RMNH). Blahnik et al.: Chimarra of northern Borneo 113

Chimarra antheae is most similar to C. gyrospina, short, at midlength with pair of elongate lateral sp. n., C. phlegyas Malicky, and C. thyestes Malicky sclerites. in possessing a tergum X with a slender process at Etymology. Named for Anthea Phillips in recognition midlength on its dorsal margin, and in the shape of of her friendship and support during J. Huisman’s the inferior appendages, each of which is narrow api- research in Sabah. cally and has the basomesal margin produced and concavely cupped. The apex of the basal part of infe- Chimarra dejongi sp. n. rior appendage in C. antheae is distinctly longer than Fig. 2 in either C. phlegyas or C. thyestes and the ventral apex of the phallobase, while being sclerotized and Type material. Holotype: ?, Malaysia, Sabah, 60 km produced as in C. phlegyas, is not nearly so strongly W Lahad Datu, Danum Valley Field Centre, ventrally curved. It differs from C. gyrospina in that nature trail brooklet, 04°58’”N, 117°48’E, 180 m, the base of the inferior appendage, in lateral view, is 9–13.ix.1986, J. Huisman (UMSP000107302) less angular, and in having a single, elongate phallic (RMNH). spine rather than a pair of spines, one of which is Paratypes. Malaysia, Sabah, 65 km W Lahad helically curved. Datu, Sg. Purut camp, 04°57’N, 117°45’E, 200 m, Adult. Color (in alcohol) yellowish-brown. Length of 26–27.x.1987, Huisman & de Jong, 1?, 2/ forewing: male 4.5–5.3 mm, female 5.7 mm. Fore- (UMSP); Tawau Hills National Park, Sg. Gelas, wing venation: stem of Rs curved, with enlarged, 20.viii.2005, W. Mey, 3? (UMHU). sclerotized node at inflection; fork at base of discal cell distinctly thickened, asymmetric, length of dis- Chimarra dejongi is closely related to and very simi- cal cell about 2 times width; m crossvein proximal to lar to Chimarra phlegyas Malicky. Both species have crossveins s and r-m, crossvein s not hyaline; 2A vein nearly identically shaped inferior appendages and intersecting 3A vein (apparently forked apically). internal phallic structure, including the presence of Postocular parietal sclerite moderately elongate. one moderately lengthed phallic spine and two small, Maxillary palps relatively short, segment 3 distinctly tack-like spines distal to the phallotremal sclerite. longer than 2, 3 subequal to 5. Protarsal claws of Chimarra dejongi differs from C. phlegyas primarily male not or very little enlarged, symmetrical. in the structure of the ventral apex of the phallobase, Male genitalia. Abdominal segment VIII short; ter- which is elongate, strongly sclerotized, and ven- gum about as long as sternum, unmodified, entire, trally deflexed in C. phlegyas, but scarcely produced without mesal excavation; sternum VIII without and very weakly sclerotized in C. dejongi. An addi- posteroventral projection. Abdominal segment IX tional difference is that the elongate, narrow dorsal very short dorsolaterally, tergum very short dorsally, processes present on the lateral lobes of tergum X in with pair of short apodemes from anterolateral mar- C. phlegyas (and also C. antheae, sp. n., C. gyrospina, gin; anteroventral margin in lateral view distinctly sp. n., and C. thyestes Malicky) seem to be absent or produced, rounded; posterior margin straight; only minimally developed in C. dejongi. ventral process projecting posteriorly, short, acute Adult. Color (in alcohol) yellowish-brown. Length apically. Preanal appendage setose, short, some- of forewing: male 4.2–4.5 mm, female 4.7– what flattened. Inferior appendage about as long 4.8 mm. Forewing venation: stem of Rs curved, with as tergum X; in lateral view strongly dorsally enlarged, sclerotized node at inflection; fork at base inflected basoventrally, widest basally, very narrow of discal cell distinctly thickened, asymmetric, length distally, basoventral margin rounded; in ventral of discal cell about 2 times width; m crossvein proxi- view mesally curved, basomesal margin produced mal to crossveins s and r-m, crossvein s not hyaline; to form broad rounded lobe, its surface concave. 2A vein intersecting 3A vein (apparently forked api- Tergum X with sclerotized lateral lobes and pro- cally). Postocular parietal sclerite short. Maxillary jecting mesal membranous lobe; lateral lobe long, palps relatively short, segment 3 slightly longer than with apex broadly rounded, in lateral view simple, 2, 3 subequal to 5. Protarsal claws of male not or very rounded apically, lobe bearing 2 sensilla. Phallobase little enlarged, symmetrical. tubular, with pronounced basodorsal expansion, Male genitalia. Abdominal segment VIII short; ter- moderately elongate, ventral apex distinctly sclero- gum longer than sternum, unmodified; sternum tized, distinctly projecting, acute, ventrally deflexed. VIII without posteroventral projection. Abdominal Endotheca length not discernable (not expanded), segment IX short dorsolaterally, tergum short dor- with granularly textured region and one prominent, sally, with pair of distinct apodemes from antero- elongate endothecal spine. Phallotremal sclerite lateral margin; anteroventral margin in lateral view complex composed of rod and ring structure, rod distinctly produced, rounded; posterior margin 114 Tijdschrift voor Entomologie, volume 152, 2009

a b membranous mesal lobe of tergum X tergum VIII preanal appendage sclerotized lateral lobe membranous mesal lobe of tergum X of tergum X sensillla sclerotized lateral lobe of tergum X preanal appendage

sensillla c

segment IX segment IX inferior appendage

ventral process d

Fig. 2. Chimarra dejongi, male genitalia. – a, lateral; b, segment IX and tergum X, dorsal; c, inferior appendage, ventral; d, phallic apparatus, lateral; inset, dorsal of phallotremal sclerite complex and phallic spine.

Chimarra demeter Malicky straight; ventral process projecting posteriorly, short, Fig. 3 acute apically. Preanal appendage setose, short, somewhat flattened. Inferior appendage about as Chimarra demeter Malicky 2000: 32, holotype ?, Malaysia long as tergum X; in lateral view strongly dorsally (Sabah), collection of H. Malicky. inflected basoventrally, widest basally, narrowing distally, basoventral margin with subacute projec- Material examined. Malaysia, Sabah, Danum Valley Field tion, apex subacute; in ventral view mesally curved, Centre, Sg. Segama, 18.viii.2005, W. Mey, 1? (UMHU); basomesal margin produced to form broad rounded 60 km W Lahad Datu, Danum Valley Field Centre, lobe, its surface concave. Tergum X with sclerotized Sg. Palum Tambun & vicinity, 04°58’N, 117°48’E, 150 m, 9.ix.1986, J. Huisman, 7?, 6/ (RMNH); same locality, lateral lobes and projecting mesal membranous lobe; 19–26.iii.1987, J. Huisman, 59?, 59/ (UMSP); 60 km lateral lobe moderately long, with apex narrowed, W Lahad Datu, Danum Valley Field Centre, Sg. Palum subacute, lobe bearing 2 sensilla, sensilla at about Tambun, 04°57’30”N, 117°48’E, 180 m, 18.iii.1987, midlength, not or scarcely projecting, apex of ven- J. Huisman, 6?, 2/ (RMNH); Danum Valley Field tral margin slightly hooked. Phallobase tubular, with Centre, nature trail brooklet, 04°58’N, 117°48’E, 180 m, pronounced basodorsal expansion, moderately elon- 9.ix.1986, J. Huisman, 5?, 10/ (UMSP); Long Pa Sia, airstrip, 04°24’N, 115°43’E, 1000 m, 16.iv.1987, gate, ventral apex weakly sclerotized, slightly project- Huisman & van Tol, 2?, 7/ (RMNH); Kg. Long Pa Sia, ing, subacute, weakly ventrally deflexed. Endotheca confluence Sg. Pa Sia & Sg. Matang, 04°24’N, 115°43’E, length not discernable (not expanded), with granu- 1000 m, 13.iv.1987, J. Huisman, 2? (UMSP); 60 km W larly textured region and 3 endothecal spines, 1 mod- Lahad Dalu, Danum Valley Field Centre, Ulu Segama, erately long, probably basal, and 2 very short, prob- 04°58’N, 117°48’E, 150 m, 8–13.ix.1986, J. Huisman, ably apical. Phallotremal sclerite complex composed 2? (UMSP); nr. Danum Valley Field Centre, W10, ? of rod and ring structure, rod short, dorsolaterally, 150 m, 2–23.viii.1987, Achterberg & Kennedy, 1 (RMNH); 60 km W Lahad Datu, Danum Valley with pair of short curved sclerites. Field Centre, nr. Segama bridge, 04°58’N, 117°48’E, Etymology. Named in honor of Dr. Rienk de Jong, 150 m, 20.x.1987, Huisman & de Jong, 1? (RMNH); friend and colleague of J. Huisman at the National 10 km NNE Ranau, Kg. Randagung baru, Sg. Pangatan, Museum of Natural History Naturalis, Leiden. 06°02’N, 116°44’E, 350 m, 6.ii.1987, J. Huisman, 23?, 7/ (UMSP). Blahnik et al.: Chimarra of northern Borneo 115 a b c d

Fig. 3. Chimarra demeter, male genitalia. – a, lateral; b, segment IX and tergum X, dorsal; c, inferior appendage, ventral; d, inferior appendage, dorsal; e, phallic apparatus, lateral.

Among species from Borneo, Chimarra demeter is with pair of short apodemes from anterolateral mar- similar to both C. devogeli, sp. n. and C. drepane, sp. n. gin; anteroventral margin in lateral view distinctly However, the inferior appendages of C. demeter are produced, rounded; posterior margin nearly straight; distinctly shorter than either of these species and the ventral process projecting posteriorly, elongate, ventral process of segment IX is not as narrow basally. narrowing from base, acute apically. Preanal The most diagnostic and distinguishing character of appendage setose, short, somewhat flattened. C. demeter is the shape of the lateral lobes of tergum Inferior appendage longer than tergum X; in lateral X, each of which has a distinctive short, truncate view strongly dorsally inflected basoventrally, wid- projection on its dorsal margin bearing 2 sensilla and est basally, narrowing distally, basoventral margin a projecting, attenuate ventral apex that is slightly rounded, apex rounded; in ventral view mesally dorsally hooked. curved, mesal surface without processes. Moderately Adult. Color (in alcohol) brown. Length of fore- elongate, dorsally inflected from base, uniformly wing: male 4.4–5.2 mm, female 4.6–5.8 mm. Fore- mesally curved; appendage widest basally, nearly wing venation: stem of Rs curved, with enlarged, uniform in width distally, apices rounded. Tergum sclerotized node at inflection; fork at base of discal X with sclerotized lateral lobes and projecting mesal cell distinctly thickened, asymmetric, length of dis- membranous lobe; lateral lobe short, with projecting cal cell about 2.5 times width; m crossvein proximal ventral margin, lobe bearing 2 sensilla, sensilla on to crossveins s and r-m, crossvein s not hyaline; 2A short, truncate middorsal lobe, apex of ventral mar- vein intersecting 3A vein (apparently forked api- gin attenuate, slightly dorsally hooked. Phallobase cally). Postocular parietal sclerite short. Maxillary tubular, with pronounced basodorsal expansion, palps relatively short, segment 3 subequal to 2 and moderately elongate, ventral margin slightly bulged 5. Protarsal claws of male slightly enlarged, sym- basally, ventral apex distinctly sclerotized, distinctly metrical. projecting, acute. Endotheca length not discernable Male genitalia. Abdominal segment VIII short; ter- (not expanded), with granularly textured region and gum about as long as sternum, unmodified, entire, 1 or 2 short endothecal spines. Phallotremal sclerite without mesal excavation; sternum VIII with indis- complex composed of rod and ring structure, rod tinct posteroventral projection. Abdominal segment very short. IX short dorsolaterally, tergum very short dorsally, 116 Tijdschrift voor Entomologie, volume 152, 2009

a b c d

Fig. 4. Chimarra devogeli, male genitalia. – a, lateral; b, segment IX and tergum X, dorsal; c, inferior appendage, ventral; d, inferior appendage, dorsal; e, phallic apparatus, lateral.

Chimarra devogeli sp. n. J. Huisman, 1? (RMNH); nr. Danum Valley Field Figs 4, 39 Centre, W9, 210 m, 14–26.iii.1987, Achterberg, Type material. Holotype: ?, Malaysia, Sabah, 2?, 7/ (RMNH); Sungai Moyig at tributary, 8 mi Long Pa Sia, confluence Sg. Pa Sia & Sg. Matang, E Penampung, 1.x.1978, G. & C. Edmunds, 7? 04°24’N, 115°43’E, 1000 m, 1.iv.1987, J. Huisman (NMNH). (UMSP000107190) (RMNH). Paratypes. Malaysia, Sabah, Danum Valley Field Chimarra devogeli is most similar to C. tawitawi Mal- Centre, Sungai Kalisun, Sungai Palun Tambun, & icky from the Philippines, particularly in the overall Sungai Segama, 17–19.viii.2005, W. Mey, 24?, shape of the inferior appendages and general struc- 3/ (UMHU); Danum Valley Field Centre, Sungai ture of tergum X. It differs in that the dorsal projec- Kalisun, 2.xi.2003, W. Mey, 14?, 1/ (UMHU); tion from tergum X is more elongate, with distinct Kinabatangan, bridge, 22.viii.2005, W. Mey, 1? projecting apical sensilla, and the ventral apex of the (UMHU); Long Pa Sia, Sg. Matang, 04°24’N, phallobase has a distinct, sinuous deflection. Consid- 115°43’E, 6.xii.1987, Huisman & Achterberg, 5?, ering the overall similarity of the species, it is likely 5/ (UMSP); 60 km W Lahad Datu, Danum Valley that the record of C. tawitawi from Borneo by Mal- Field Centre, Sg. Palum Tambun & vicinity, 04°58’N, icky (2008) actually represents this species. Among 117°48’E, 150 m, 19–26.iii.1987, J. Huisman, 7?, species from Borneo, Chimarra devogeli is most simi- 4/ (UMSP); Long Pa Sia, confluence Sg. Pa Sia & lar to C. drepane, sp. n. and C. demeter Malicky. It Sg. Matang, 04°24’N, 115°43’E, 1000 m, 1.iv.1987, resembles both those species in having elongate nar- J. Huisman, 13? (RMNH); same locality, 5.iv.1987, row inferior appendages without mesal projections, J. Huisman, 3?, 8/ (RMNH); Long Pa Sia airstrip, and particularly in the structure of tergum X, which 04°24’N, 115°43’E, 1000 m, 7.xii.1987, J. Huis- has the ventral margin forming a sclerous projec- man, 2?, 3/ (UMSP); 75 km. W Lahad Datu, tion and a dorsal margin with a process bearing two confl. S. Sabran, S. Danum S/N, 04°34’12”N, sensilla. It differs in that the inferior appendages are 117°24’36”E, 200 m, 23.x.1987, J. Huisman, 1?, more elongate than C. demeter, but shorter than in 1/ (RMNH); Kg. Long Pa Sia, confluence Sg. Pa C. drepane, and in the shape of the dorsal process of Sia & Sg. Matang, 04°24’N, 115°43’E, 1000 m, tergum X, which is not upright and truncate as in 13.iv.1987, J. Huisman, 19? (UMSP); 60 km C. demeter. The slightly decurved and projecting W Lahad Dalu, Danum Valley Field Centre, Ulu apex of the phallobase seems to be a consistent dif- Segama, 04°58’N, 117°48’E, 150 m, 8–13.ix.1986, ference between this species and C. drepane. Blahnik et al.: Chimarra of northern Borneo 117

Adult. Color (in alcohol) light brown. Length of vicinity, 04°58’N, 117°48’E, 150 m, 19–26.iii.1987, forewing: male 4.7–5.0 mm, female 4.2–5.9 mm. J. Huisman, 1? (RMNH); Sapulut, kampong, road- Forewing venation: stem of Rs curved, with enlarged, side, 04°42’N, 116°29’E, 290 m, 4.v.1987, J. Huis- sclerotized node at inflection; fork at base of discal man, 1?, 1/ (UMSP). cell distinctly thickened, asymmetric, length of discal cell about 2.5 times width; m crossvein proximal to Chimarra drepane is probably most similar to crossveins s and r-m, crossvein s not hyaline; 2A vein C. monorum Chantaramongkol & Malicky, described intersecting 3A vein (apparently forked apically). from Thailand, particularly in the length and shape Postocular parietal sclerite short. Maxillary palps of the inferior appendages, but differs in details of moderately elongate, segment 3 subequal to 2 and 5. tergum X. Among species from Borneo, C. drepane Protarsal claws of male not or very little enlarged, is similar to C devogeli, sp. n. and C. demeter Malicky. symmetrical. It resembles those species in having elongate narrow Male genitalia. Abdominal segment VIII short; ter- inferior appendages without mesal projections, and gum about as long as sternum, unmodified; sternum particularly in the structure of tergum X, which has VIII with slight posteroventral projection. Abdomi- the ventral margin forming a sclerous projection and nal segment IX very short dorsolaterally, tergum very a dorsal margin with a process bearing two sensilla. short dorsally, with pair of distinct apodemes from It differs in that the inferior appendages are more anterolateral margin; anteroventral margin in lateral elongate than C. devogeli, and much more elongate view distinctly produced, angular; posterior margin than C. demeter, and in the shape of the dorsal proc- nearly straight; ventral process projecting posteriorly, ess of tergum X, which is not upright and truncate as elongate, narrow, acute apically. Preanal appendage in C. demeter. The shorter phallobase, with a nearly setose, short, rounded, broadest basally. Inferior straight projecting ventral apex seems to be a consist- appendage elongate, much longer than tergum X; ent difference between C. drepane and C. devogeli, in lateral view dorsally inflected basoventrally, wid- which it otherwise closely resembles. est basally, gradually narrowing distally, basoventral Adult. Color (in alcohol) light brown. Length of margin rounded, apex subacute; in ventral view forewing: male 3.8–4.6 mm, female 4.5–4.7 mm. uniformly mesally curved, mesal surface without Forewing venation: stem of Rs curved, with enlarged, processes. Tergum X with sclerotized lateral lobes sclerotized node at inflection; fork at base of discal and projecting mesal membranous lobe; lateral lobe cell distinctly thickened, asymmetric, length of dis- short, with projecting ventral margin, lobe bearing 2 cal cell about 2 times width; m crossvein proximal to sensilla, sensilla on projecting, digitate dorsal lobe, crossveins s and r-m, crossvein s not hyaline; 2A vein apex of ventral margin narrow, subacute. Phallobase intersecting 3A vein (apparently forked apically). tubular, with pronounced basodorsal expansion, Postocular parietal sclerite short. Maxillary palps moderately elongate, ventral margin slightly bulged moderately elongate, segment 3 subequal to 2 and basally, slightly deflexed preapically, ventral apex dis- 5. Protarsal claws of male not or very little enlarged, tinctly sclerotized, distinctly projecting, subacute. symmetrical. Endotheca length not discernable (not expanded), Male genitalia. Abdominal segment VIII short; ter- with granularly textured region and 2 short, sub- gum about as long as sternum, unmodified; sternum equal endothecal spines. Phallotremal sclerite com- VIII with slight posteroventral projection. Abdomi- plex composed of rod and ring structure, rod very nal segment IX very short dorsolaterally, tergum very short, preapically with short lateral sclerites. short dorsally, with pair of short apodemes from Etymology. Named in honor of Dr. Ed de Vogel, anterolateral margin; anteroventral margin in lateral Hortus Botanicus, University of Leiden, friend and view distinctly produced, angular; posterior margin colleague of J. Huisman. nearly straight; ventral process projecting posteriorly, elongate, narrow, acute apically. Preanal appendage Chimarra drepane sp. n. setose, short, somewhat flattened. Inferior appendage very elongate, much longer than tergum X; in lateral Fig. 5 view strongly dorsally inflected basoventrally, nearly uniformly wide, basoventral margin rounded, apex Type material. Holotype: ?, Malaysia, Sabah, rounded; in ventral view uniformly mesally curved, 10 km SE Ranau, Kg. Nalapak, Sg. Kananapun, mesal surface without processes. Tergum X with 05°59’N, 116°47’E, 350 m, 7.ii.1987, J. Huisman, sclerotized lateral lobes and projecting mesal mem- (UMSP000107169) (RMNH). branous lobe; in lateral view with projecting ventral Paratypes. Malaysia, Sabah, 60 km W Lahad Datu, margin, apex of ventral margin narrow, subacute; Danum Valley Field Centre, Sg. Palum Tambun & dorsal margin with projecting, rounded lobe with 118 Tijdschrift voor Entomologie, volume 152, 2009

a b c d

Fig. 5. Chimarra drepane, male genitalia. – a, lateral; b, segment IX and tergum X, dorsal; c, inferior appendage, ventral; d, inferior appendage, dorsal; e, phallic apparatus, lateral.

2 sensilla. Phallobase tubular, with pronounced Borneo, it is easily recognized by the position of the basodorsal expansion, relatively short, slightly sensilla of tergum X, which are on a rounded proc- expanded apically, ventral margin slightly bulged ess near its ventral margin, and by the general shape basally, ventral apex distinctly sclerotized, distinctly of the inferior appendages, which also have a small projecting, acute. Endotheca length not discernable mesal projection. The single, elongate phallic spine (not expanded), with apparent tract of numerous is also a distinctive character, although possibly vari- minute spines and 2 prominent endothecal spines, able. It is elongate in the holotype illustration by 1 moderate in length and 1 very short. Phallotremal Kimmins and also in the specimens from Sabah, but sclerite complex composed of rod and ring structure, less elongate in the specimen illustrated by Malicky rod very short, preapically with short lateral scler- (1989, fig. 8a-c) from Sumatra. ites. Adult. Color (in alcohol) light brown. Length of Etymology. This species is named Chimarra drepane forewing: male 4.2–4.5 mm. Forewing venation: from the Greek word drepane, a sickle, used as a noun stem of Rs curved, with enlarged, sclerotized node at in apposition, and referring to the elongate, mesally inflection; fork at base of discal cell distinctly thick- curved and sickle-shaped inferior appendages. ened, asymmetric, length of discal cell about 2 times width; m crossvein proximal to crossveins s and r-m, s Chimarra dulitensis Kimmins crossvein not hyaline; 2A vein intersecting 3A vein (apparently forked apically). Postocular parietal scle- Fig. 6 rite short. Maxillary palps relatively short, segment Chimarra dulitensis Kimmins, 1955: 378, holotype ?, Ma- 3 subequal to 2 and 5. Protarsal claws of male not or laysia (Sarawak), BMNH; Malicky 1989: 135, Indo- very little enlarged, symmetrical. nesia (Sumatra). Male genitalia. Abdominal segment VIII short; tergum longer than sternum, unmodified; sternum Material examined. Malaysia, Sabah, SE Sabah nr. Da- VIII without posteroventral projection. Abdomi- num Valley Field Center, 240 m, 18.x-22.xi.1987, Achter- nal segment IX short dorsolaterally, tergum short berg, 1? (RMNH); Long Pa Sia, confluence Sg. Pa Sia & dorsally, with pair of short apodemes from antero- Sg. Matang, 04°24’N, 115°43’E, 1000 m, 10.iv.1987, lateral margin; anteroventral margin in lateral view J. Huisman, 1? (UMSP); 60 km W Lahad Datu, Danum Valley Field Centre, nr. Segama bridge, 04°58’N, 117°48’E, produced, angular; posterior margin nearly straight; 150 m, 20.x.1987, Huisman & de Jong, 1? (UMSP). ventral process projecting posteriorly, moderately elongate, narrowing from base, acute apically. Pre- Chimarra dulitensis is a distinctive member of the anal appendage setose, short, somewhat flattened. Chimarra digitata lineage. Among species from Inferior appendage longer than tergum X; in lateral Blahnik et al.: Chimarra of northern Borneo 119 a b c d

Fig. 6. Chimarra dulitensis, male genitalia. – a, lateral; b, segment IX and tergum X, dorsal; c, inferior appendage, ventral; d, inferior appendage, dorsal; e, phallic apparatus, lateral. view dorsally inflected basoventrally, widest basally, Marei-Parei, Sg. Tawubang, 06°04’N, 116°30’E, narrowing distally, basoventral margin rounded, 1030 m, 11.iii.1987, J. Huisman, 2? (UMSP). apex rounded; in ventral view mesally curved, mesal surface with indistinct, broadly rounded projection Chimarra fuilianae is similar to C. tityos Malicky at midlength. Tergum X with sclerotized lateral lobes and C. terramater Malicky, but possibly most closely and projecting mesal membranous lobe; with apex related to C. xiphosella. All of these species have the broadly rounded, lobe bearing 2 sensilla, sensilla on character combination of a tergum X with a project- slightly projecting, ventrolateral lobe. Phallobase ing and somewhat upturned ventral margin, sensilla tubular, with pronounced basodorsal expansion, on a rounded projection at midlength on the same moderately elongate, ventral margin slightly bulged structure, an inferior appendage with a projection basally, ventral apex weakly sclerotized, slightly from its mesal margin, and a relatively elongate ven- projecting. Endotheca length not discernable (not tral process on segment IX. Among these species, expanded), granularly textured region not discern- C. fuilianae is distinguished by the relatively narrow able, with 1, prominent, elongate spine. Phallotre- and rounded basal margin of its inferior appendage, mal sclerite complex composed of elongate ring, rod as well as the overall shape of tergum X and the shape very reduced. and length of the ventral process of segment IX. Adult. Color (in alcohol) yellowish-brown. Length Chimarra fuilianae sp. n. of forewing: male 3.8–4.1 mm. Forewing venation: stem of Rs curved, with enlarged, sclerotized node Fig. 7 at inflection; fork at base of discal cell distinctly thickened, asymmetric, length of discal cell about Type material. Holotype: ?, Malaysia, Sabah, 12 km 2 times width; m crossvein proximal to crossveins NNE Ranau, Poring Hot Spring, Sg. Kepungit, s and r-m, crossvein s not hyaline; 2A vein intersect- 06°03’N, 116°42’E, 550 m, 24.i.1987, J. Huisman ing 3A vein (apparently forked apically). Postocular (UMSP000107250) (RMNH). parietal sclerite short. Maxillary palps moderately Paratypes. Malaysia, Sarawak, Gunung Gading elongate, segment 3 distinctly longer than 2, 3 sub- National Park, Sungai Lundu, 23–26.x.2003, W. Mey, equal to 5. Protarsal claws of male not or very little 5? (UMHU); Sabah, 9.5 km NNW Kundassang, enlarged, symmetrical. 120 Tijdschrift voor Entomologie, volume 152, 2009

a b c d

endotheca endothecal spines phallotremal sclerite phallotheca complex

Fig. 7. Chimarra fuilianae, male genitalia. – a, lateral; b, segment IX and tergum X, dorsal; c, inferior appendage, ventral; d, inferior appendage, dorsal; e, phallic apparatus, lateral.

Male genitalia. Abdominal segment VIII short; ter- slender, curved apical spine. Phallotremal sclerite gum about as long as sternum, unmodified; sternum complex composed of elongate ring, rod not evident, VIII without posteroventral projection. Abdominal apically with pair of short, curved, lateral sclerites. segment IX short dorsolaterally, tergum short dor- Etymology. Named in honor of Fui-Lian Tan, friend sally, with pair of distinct apodemes from anterola- of J. Huisman. teral margin; anteroventral margin in lateral view distinctly produced, angular; posterior margin slightly Chimarra gyrospina sp. n. produced; ventral process projecting posteriorly, Fig. 8 elongate, nearly uniform in width, rounded apically. Preanal appendage setose, short, rounded or subquadrate, somewhat flattened, slightly narrowed basally. Type material. Holotype: ?, Malaysia, Sabah, Inferior appendage longer than tergum X; in lateral ½ way, road Melligan to Long Pa Sia, 04°35’N, view dorsally inflected basoventrally, widest basally, 115°42’E, 1200 m, 15.xii.1986, J. Huisman narrowing distally, basoventral margin rounded, (UMSP000107290) (RMNH). apex subacute; in ventral view mesally curved, bas- Paratypes. Malaysia, Sabah, same data as holotype, omesally with slightly angulate projection, mesal 1/ (RMNH). surface with tooth-like projection at midlength. Ter- gum X with sclerotized lateral lobes and projecting Chimarra gyrospina is similar to C. antheae, sp. n., mesal membranous lobe; lateral lobe with project- C. thyestes Malicky, and C. phlegyas Malicky in posing ventral margin, lobe bearing 2 sensilla, sensilla sessing a tergum X with a slender process at midlength on slightly projecting, rounded middorsal lobe, apex on its dorsal margin, and in the shape of the infe- of ventral margin relatively broad, slightly hooked. rior appendages, which are very narrow apically and Phallobase tubular, with pronounced basodorsal have the basomesal margin produced and concavely expansion, moderately elongate, ventral margin cupped. It is easily distinguished by the overall shape slightly bulged basally, ventral apex weakly sclero- of the inferior appendages and differs from the other tized, slightly projecting, acute. Endotheca elongate, species mentioned in that the apical phallic spine tubular, with granularly textured region; 3 endothe- is very distinctly curled, the ventral apex of the cal spines, 2 subequal, short, basal spines and 1 short, phallobase is less developed, and one of the sensilla Blahnik et al.: Chimarra of northern Borneo 121 a b c

Fig. 8. Chimarra gyrospina, male genitalia. – a, lateral; b, segment IX and tergum X, dorsal; c, inferior appendage, ventral; d, phallic apparatus, lateral.

of tergum X is located on its dorsal process. lateral view strongly dorsally inflected basoventrally, Adult. Color (in alcohol) yellowish-brown. Length of widest basally, very narrow distally, basoventral mar- forewing: male 4.5 mm, female 5.3 mm. Forewing gin acute, apex subacute; in ventral view mesally venation: stem of Rs curved, with enlarged, sclero- curved, basomesal margin produced to form broad tized node at inflection; fork at base of discal cell rounded lobe, its surface concave. Tergum X with distinctly thickened, asymmetric, length of discal sclerotized lateral lobes and projecting mesal mem- cell about 2.5 times width; m crossvein proximal to branous lobe; lateral lobe long, with apex broadly crossveins s and r-m, crossvein s not hyaline; 2A vein rounded, in lateral view with elongate, digitate intersecting 3A vein (apparently forked apically). process at midlength on dorsal margin, lobe bear- Postocular parietal sclerite short. Maxillary palps ing 2 sensilla, one sensillum located basolaterally, moderately elongate, segment 3 distinctly longer other on elongate dorsal process. Phallobase tubular, than 2, 3 subequal to 5. Protarsal claws of male not with pronounced basodorsal expansion, moderately or very little enlarged, symmetrical. elongate, ventral apex weakly sclerotized, slightly Male genitalia. Abdominal segment VIII short; ter- projecting, acute. Endotheca length not discern- gum about as long as sternum, unmodified; sternum able (not expanded), granularly textured region not VIII without posteroventral projection. Abdominal discernable, with 2 endothecal spines, 1 long basal segment IX short dorsolaterally, tergum short dor- spine and 1 long, slender, curved apical spine. Phal- sally, with pair of short apodemes from anterola- lotremal sclerite complex composed of rod and ring teral margin; anteroventral margin in lateral view structure, rod very short. slightly produced, angular; posterior margin slightly Etymology. This species is named Chimarra gyrospina produced; ventral process projecting posteriorly, from the Latin word gyrus, meaning to turn around short, narrowing from base, acute apically. Pre- or circle, and referring to the helically bent apical anal appendage setose, short, somewhat flattened. phallic spine. Inferior appendage about as long as tergum X; in 122 Tijdschrift voor Entomologie, volume 152, 2009

a b c

Fig. 9. Chimarra phlegyas, male genitalia. – a, lateral; b, segment IX and tergum X, dorsal; c, inferior appendage, ventral; d, phallic apparatus, lateral.

Chimarra phlegyas Malicky Jong, 1?, 1/ (RMNH); 60 km W Lahad Datu, Da- Figs 9, 40 num Valley Field Centre, EO trail, Sg. Segama, 04°58’N, 117°48’E, 150 m, 18.x.1987, Huisman & de Jong,, 2?, ? Chimarra phlegyas Malicky, 2008: 840, holotype , In- 1/ (RMNH); Sg. Moyig at tributary, 8 mi E Penampung, donesia (Eastern Kalimantan), Zoologische Museum 1.x.1978, G. & C. Edmunds, 1? (NMNH). Lausanne. Material examined. Malaysia, Sabah, 11 km S Nabawan, Chimarra phlegyas is very similar to C. dejongi, sp. n. pond on roadside, 04°57’N, 116°27’E, 400 m, 15.xi.1987, Both species have nearly identically shaped inferior J. Huisman, 1? (RMNH); Danum Valley Field Center, appendages and internal phallic structure, includ- ? Sg. Palun Tambun, 17.viii.2005, W. Mey, 4 (UMHU); ing the presence of one moderately lengthed phal- Tawau Hills National Park, Sg. Gelas, 20.viii.2005 & 3.iii.2006, W. Mey, 8? (UMHU); Kinabalu National Park, lic spine and two small, tack-like spines distal to the Poring Hot Springs, 30.x.2003, W. Mey, 1?, (UMHU); phallotremal sclerite. Chimarra phlegyas differs from 60 km W of Lahad Datu, DVFC, brooklet above lotus C. dejongi primarily in the structure of the ventral pond, 04°58’N, 117°48’E, 150 m, 19–28.x.1987, Huisman apex of the phallobase, which is elongate, strongly & de Jong, 46?, 26/ (UMSP); 60 km W Lahad Datu, sclerotized, and ventrally deflexed in C. phlegyas, Danum Valley Field Centre, Sg. Palum Tambun & vicinity, but scarcely produced and very weakly sclerotized in 04°58’N, 117°48’E, 150 m, 19–26.iii.1987, J. Huisman, 5? (RMNH); Long Pa Sia airstrip, 04°24’N, 115°43’E, C. dejongi. An additional difference is that the elon- 1000 m, 15.iv.1987, Huisman & van Tol, 2? (RMNH); gate, narrow dorsal process present on the lateral Poring Hot Spring, 11 km NNE Ranau, Sg. Montokungon lobe of tergum X in C. phlegyas seems to be absent or crossing road, 06°02’30”N, 116°43’E, 450 m, 28.i.1987, only minimally developed in C. dejongi. J. Huisman, 1? (UMSP); 20 km NNE Ranau, Kg. Taku- Adult. Color (in alcohol) yellowish-brown. Length tan, Sg. Mokodou, 06°06’N, 116°45’E, 325 m, 4.ii.1987; of forewing: male 4.0–5.0 mm, female 4.2–5.2 mm. ? J. Huisman, 1 (RMNH); 12 km NNE Ranau, Poring Forewing venation: stem of Rs curved, with enlarged, Hot Spring, staff quarters, 06°03’N, 116°42’E, 550 m, 1.ii.1987, Huisman & de Jong, 25? (UMSP); 60 km W sclerotized node at inflection; fork at base of discal Lahad Dalu, Danum Valley Field Centre, Ulu Segama, cell distinctly thickened, asymmetric, length of discal 04°58’N, 117°48’E, 150 m, 8–13.ix.1986, J. Huisman, cell about 2.5 times width; m crossvein proximal to 3? (RMNH); 60 km W Lahad Datu, Danum Valley Field crossveins s and r-m, crossvein s not hyaline; 2A vein Centre, nr. Segama bridge, 04°58’N, 117°48’E, 150 m, intersecting 3A vein (apparently forked apically). 20.x.1987, Huisman & de Jong, 5?, 2/ (UMSP); 12.5 km Postocular parietal sclerite short. Maxillary palps S Nabawan, on road to Kg. Pamuntariah, 1st bridge, 04°57’N, 116°27’E, 400 m, 16.xi.1987, J. Huisman, 1? relatively short, segment 3 slightly longer than 2, 3 (RMNH); 39 km on road Keningau-Nabawan, Kg. Sook, subequal to 5. Protarsal claws of male not or very 05°12’N, 116°19’E, 350 m, 17.xi.1987, Huisman & de little enlarged, symmetrical. Blahnik et al.: Chimarra of northern Borneo 123

Male genitalia. Abdominal segment VIII short; ter- Poring Hot Spring, 12 km NNE Ranau, Sg. Kepungit, gum about as long as sternum, unmodified; sternum E park boundary, 06°03’N, 116°42’E, 480 m, 26.xi.1986, VIII without posteroventral projection. Abdominal J. Huisman, 3? (UMSP); Long Pa Sia, airstrip, 04°24’N, 115°43’E, 1000 m, 16.iv.1987, Huisman & van Tol, 1? segment IX short dorsolaterally, tergum short dor- (RMNH); 12 km NNE Ranau, Poring Hot Spring, staff sally, with pair of distinct apodemes from anterola- quarters, 06°03’N, 116°42’E, 550 m, 1.ii.1987, Huis- teral margin; anteroventral margin in lateral view dis- man & de Jong, 1? (UMSP); 12 km NNE Ranau, Por- tinctly produced, rounded; posterior margin nearly ing Hot Spring, Sg. Tananasad, 06°03’N, 116°42’E, straight; ventral process projecting posteriorly, short, 560 m, 31.i.1987, J. Huisman, 3? (UMSP); Kg. Long broad basally, subtriangular. Preanal appendage Pa Sia, confluence Sg. Pa Sia & Sg. Matang, 04°24’N, ? setose, short, somewhat flattened. Inferior append- 115°43’E, 1000 m, 13.iv.1987, J. Huisman, 1 (UMSP); Interior Zone, Tenom, 16 km NE Agr. Res. Station, 200 m, age about as long as tergum X; in lateral view strongly 14–16.xii.1989, J. Huisman, 1?, 2/ (RMNH); Sg. Mala- dorsally inflected basoventrally, widest basally, grad- bit, rd. Long Pa Sia to Long Semado, 04°21’N, 115°41’E, ually narrowing distally, basoventral margin with 1175 m, 18–19.xii.1986, J. Huisman, 1?, 1/ (UMSP). subacute projection, apex rounded; in ventral view mesally curved, basomesal margin produced to form Chimarra terramater is similar to C. tityos Malicky, broad rounded lobe, its surface concave. Tergum X C. fuilianae, sp. n., and C. xiphosella, sp. n. All of with sclerotized lateral lobes and projecting mesal these species have the character combination of hav- membranous lobe; lateral lobe moderately long, with ing the lateral lobe of tergum X with a projecting projecting ventral margin, in lateral view with elon- and somewhat upturned ventral margin, sensilla on a gate, digitate process at midlength on dorsal margin, rounded projection at midlength on the same struc- lobe bearing 2 sensilla, sensilla located basolaterally, ture, an inferior appendage with a projection from apex of ventral margin slightly hooked. Phallobase its mesal margin, and a relatively elongate ventral tubular, with pronounced basodorsal expansion, rel- process on segment IX. Like C. tityos, C. terramater atively short, ventral apex distinctly sclerotized, dis- has the apex of the lateral lobes of tergum X sharply tinctly projecting, acute, strongly ventrally deflexed. upturned and acute (typically more so than in the Endotheca length not discernable (not expanded), illustration provided by Malicky), but differs in granularly textured region not discernable, with lacking an acutely projecting basoventral projection 3 endothecal spines, 1 moderately long, probably on the inferior appendages. The overall morphol- basal, and 2 very short, probably apical. Phallotremal ogy, and especially the inferior appendages are most sclerite complex composed of rod and ring structure, similar to C. fuilianae, but in the material examined rod short, at midlength with pair of sclerites. the basal part of the inferior appendage always seems to have an angular inflection. This is not apparent in Chimarra terramater Malicky the original illustration of the species, but the other characters, especially the overall shape of tergum X, Fig. 10 seem to conform closely to those of the specimen Chimarra terramater Malicky, 2008: 841, holotype ?, In- illustrated here. Chimarra terramater also seems to donesia (Eastern Kalimantan), Zoologische Museum differ diagnostically from C. fuilianae in that the Lausanne. ventral process of segment IX is distinctly shorter. Adult. Color (in alcohol) yellowish-brown. Length Material examined. Malaysia, Sarawak, Gunung Gad- of forewing: male 3.5–4.5 mm, female 3.9–4.6 ing National Park, Sg. Lundu, 23–26.x.2003, W. Mey, mm. Forewing venation: stem of Rs curved, with 72? (UMHU); Sabah, Poring Hot Spring, 12 km NNE enlarged, sclerotized node at inflection; fork at base Ranau, Sg. Montokungon, 06°02’N, 116°42’E, 525 m, of discal cell distinctly thickened, asymmetric, length 30.i.1987, J. Huisman,, 1?, 12/ (RMNH); Crocker Range National Park, Sg. Myoog, 9.xi.2003, W. Mey, of discal cell about 2 times width; m crossvein proxi- 2? (UMHU); West Coast Zone, Sg. Kepungit water- mal to crossveins s and r-m, crossvein s not hyaline; fall, 06°03’N, 116°42’E, 550 m, 5.xii.1986, J. Huisman, 2A vein intersecting 3A vein (apparently forked api- 4?, 1/ (UMSP); Long Pa Sia, confluence Sg. Pa Sia & cally). Postocular parietal sclerite short. Maxillary Sg. Matang, 04°24’N, 115°43’E, 1000 m, 10.iv.1987, palps moderately elongate, 3 distinctly longer than J. Huisman, 1? (UMSP); 3.5 km SW Kg. Long Pa Sia 2, 5 slightly shorter than 3. Protarsal claws of male & Sg. Ritan, 04°24’N, 115°42’E, 1160 m, 8–9.iv.1987, J. Huisman, 5? (UMSP); same locality, 14–16.xii.1989, not or very little enlarged, symmetrical. J. Huisman, 3/ (RMNH); Poring Hot Spring, 11 km NNE Male genitalia. Abdominal segment VIII moderate in Ranau, Sg. Montokungon crossing road, 06°02’30”N, length; tergum about as long as sternum, unmodified; 116°43’E, 450 m, 28.i.1987, J. Huisman, 1? (RMNH); sternum VIII with slight posteroventral projection. 20 km NNE Ranau, Kg. Takutan, Sg. Mokodou, 06°06’N, Abdominal segment IX short dorsolaterally, tergum 116°45’E, 325 m, 4.ii.1987, J. Huisman, 1? (RMNH); short dorsally, with pair of distinct apodemes from 124 Tijdschrift voor Entomologie, volume 152, 2009

a b c d

Fig. 10. Chimarra terramater, male genitalia. – a, lateral; b, segment IX and tergum X, dorsal; c, inferior appendage, ventral; d, inferior appendage, dorsal; e, phallic apparatus, lateral.

Chimarra tityos Malicky anterolateral margin; anteroventral margin in lateral Fig. 11 view distinctly produced, angular; posterior margin nearly straight; ventral process projecting posteriorly, Chimarra tityos Malicky, 2008: 839, holotype ?, Indo- elongate, tapering, rounded to subtruncate apically. nesia (Eastern Kalimantan), Zoologische Museum Preanal appendage setose, short, somewhat flattened. Lausanne. Inferior appendage slightly longer than tergum X; in lateral view strongly dorsally inflected basoventrally, Material examined. Malaysia, Sarawak, Gunung Gad- widest basally, narrowing distally, basoventral margin ing National Park, Sg. Lundu, 23.-26.x.2003, W. Mey, rounded, apex acute; in ventral view mesally curved, 61? (MNHU); Long Rapun, 16 km N Bairo, Sg. Dapur, basomesally with slightly angulate projection. Ter- 03°53’N, 115°35’E, 1200 m, 19–22.ii.1987, J. Huisman, 1? (UMSP). Sabah, 60 km W Lahad Datu, Danum Val- gum X with sclerotized lateral lobes and projecting ley Field Centre, Sg. Palum Tambun & vicinity, 04°58’N, mesal membranous lobe; lateral lobe moderately 117°48’E, 150, 9.xi.1986, J. Huisman 1? (RMNH); long, with projecting ventral margin, lobe bearing Crocker Range National Park, Sg. Myoog, 8.xi.2003, 2 seta-like sensilla, sensilla on slightly projecting, W. Mey, 9?, 6/ (MNHU); Ranau, 20 km E Sg. Bayaan, short, rounded middorsal lobe, apex of ventral mar- 6.xi.2003, W. Mey, 4?, 1/ (MNHU); Danum Val- gin distinctly hooked dorsally. Phallobase tubular, ley Field Centre, Sg. Palum Tambun, 18–19.viii.2005, W. Mey, 19? (MNHU); Danum Valley Field Centre, with pronounced basodorsal expansion, moderately Sg. Kalisun, 17.viii.2005, W. Mey, 14?, 1/ (MNHU); elongate, ventral margin slightly bulged basally, ven- same locality, 2.xi.2003, W. Mey, 1?; Danum Valley tral apex weakly sclerotized. Endotheca elongate, Field Centre, Tembaling trail, 18.viii.2005, W. Mey, 1? tubular, with granularly textured region and 2 short, (MNHU); Tawau Hills National Park, Sg. Tawau, 19– subequal endothecal spines. Phallotremal sclerite 21.viii.2005, W. Mey, 8?, 2/ (MNHU); Long Pa Sia, complex composed of rod and ring structure, rod Sg. Matang, 04°24’N, 115°43’E, 6.xii.1987, Huisman & ? / moderately elongate. Achterberg, 1 , 1 (RMNH); SE Sabah nr. Danum Val- ley Field Center, 240 m, 18.x-22.xi.1987, Achterberg, 1?, 1/ (RMNH); 60 km W Lahad Datu, Danum Valley Field Centre, Sg. Palum Tambun & vicinity, 04°58’N, 117°48’E, 150, 9.xi.1986, J. Huisman, 2? (RMNH); same locality, Blahnik et al.: Chimarra of northern Borneo 125 a b c d

Fig. 11. Chimarra tityos, male genitalia. – a, lateral; b, segment IX and tergum X, dorsal; c, inferior appendage, ventral; d, inferior appendage, dorsal; e, phallic apparatus, lateral.

19–26.iii.1987, J. Huisman, 3?, 1/ (UMSP); Long in having an angulate, sclerotized mesal projection Pa Sia airstrip, 04°24’N, 115°43’E, 1000 m, 15.iv.1987, from the inferior appendage and in having the lateral Huisman & van Tol, 1? (UMSP); 60 km W Lahad lobes of tergum X with upturned apices. Distinctive Datu, Danum Valley Field Centre, nature trail brooklet, differences include a preapical ventral projection 04°58’N, 117°48’E, 180 m, 9–13.ix.1986, J. Huisman, 5? (UMSP); Poring Hot Spring, 11 km NNE Ranau, from the phallobase, a more prominent apicoventral Sg. Montokungon crossing road, 06°02’30”N, 116°43’E, projection from the inferior appendage, a less elon- 450 m, 25.xi.1986, J. Huisman, 18? (UMSP); same local- gate ventral process on segment IX, and a distinctive ity, 28.i.1987, J. Huisman, 12?, 23/ (UMSP); Poring Hot spinose tract on the endotheca. Spring, 12 km NNE Ranau, confluence Sg. Kepungit & Adult. Color (in alcohol) light brown. Length of Sg. Langanan, 06°03’N, 116°43’E, 450 m, 8.xii.1986, forewing: male 4.0–4.8 mm, female 4.0–5.2 mm. J. Huisman, 1?, 1/ (RMNH); 20 km NNE Ranau, Kg. Takutan, Sg. Mokodou, 06°06’N, 116°45’E, 325 m, Forewing venation: stem of Rs curved, with enlarged, 4.ii.1987, J. Huisman, 2?, 2/ (UMSP); 10 km SE sclerotized node at inflection; fork at base of discal Ranau, Kg. Nalapak, Sg. Kananapun, 05°59’N, 116°47’E, cell distinctly thickened, asymmetric, length of discal 350 m, 7.ii.1987, J. Huisman, 7?, 4/ (UMSP); Long cell about 2.5 times width; m crossvein proximal to Pa Sia, airstrip, 04°24’N, 115°43’E, 1000 m, 16.iv.1987, crossveins s and r-m, crossvein s not hyaline; 2A vein Huisman & van Tol, 1? (RMNH); 2 km SW Long Pa intersecting 3A vein (apparently forked apically). Sia, Long Rurun (Sg. Ritan), 04°23’N, 115°42’E, 1040 m, Postocular parietal sclerite short. Maxillary palps 5.xii.1987, Huisman & Achterberg, 2?, 1/ (RMNH); 12 km NNE Ranau, Poring Hot Spring, staff quarters, moderately elongate, segment 3 distinctly longer 06°03’N, 116°42’E, 550 m, 1.ii.1987, Huisman & de than 2, 3 subequal to 5. Protarsal claws of male not Jong, 1? (RMNH); Sg. Moyig at tributary, 8 mi E Penam- or very little enlarged, symmetrical. pung, 1.x.1978 G. & C. Edmunds, 17? (NMNH); Ten- Male genitalia. Abdominal segment VIII short; ter- om, 1.ix.1983, Steiner & Hevel, 3? (NMNH); Ranau, gum about as long as sternum, unmodified, entire, 13.viii.1983, Steiner & Hevel, 3?, 7/ (NMNH). without mesal excavation; sternum VIII with slight posteroventral projection. Abdominal seg- Chimarra tityos is most similar to C. fuilianae, sp. n. ment IX short dorsolaterally, tergum short dorsally, and C. terramater Malicky. It resembles these species with pair of distinct apodemes from anterolateral 126 Tijdschrift voor Entomologie, volume 152, 2009

a b c d

Fig. 12. Chimarra xiphosella, male genitalia. – a, lateral; b, segment IX and tergum X, dorsal; c, inferior appendage, ventral; d, inferior appendage, dorsal; e, phallic apparatus, lateral.

Chimarra xiphosella sp. n. margin; anteroventral margin in lateral view pro- Fig. 12 duced, rounded; posterior margin nearly straight; ventral process projecting posteriorly, elongate, narrowing from base, acute apically. Preanal appendage Type material. Holotype: ?, Malaysia, Sabah, 9.5 km setose, short, rounded or subquadrate, somewhat NNW Kundassang, Marei-Parei, Sg. Tawubang, flattened, slightly narrowed basally. Inferior append- 06°04’N, 116°30’E, 1030 m, 11.iii.1987, J. Huis- age about as long as tergum X; in lateral view dor- man (UMSP000107141) (RMNH). sally inflected basoventrally, widest basally, narrowing distally, apex subacute; in ventral view mesally Chimarra xiphosella is similar to C. fuilianae, sp. n., curved, basally with acute projection from posterior C. terramater Malicky, and C. tityos Malicky. All margin, mesal surface with broad tooth-like projec- of these species have the character combination of tion at midlength. Tergum X with sclerotized lateral having the lateral lobe of tergum X with a project- lobes and projecting mesal membranous lobe; lat- ing and somewhat upturned ventral margin, sen- eral lobe moderately long, with projecting ventral silla on a rounded projection at midlength on the margin, lobe bearing 2 sensilla, sensilla on slightly same structure, an inferior appendage with a pro- projecting, short, rounded middorsal lobe, apex of jection from its mesal margin, and a relatively elon- ventral margin distinctly hooked dorsally. Phallobase gate ventral process on segment IX. Among these, tubular, with pronounced basodorsal expansion, C. xiphosella is particularly distinctive, and easily relatively short, ventral apex weakly sclerotized, with diagnosed by the shape of the inferior appendages, strongly deflexed ventral process. Endotheca length each of which has a prominent apicoventral pro- not discernable (not expanded), with tract of numer- jection, and by the very elongate ventral process of ous needle-like spines near apex; and 2 fine, short, segment IX. subequal endothecal spines. Phallotremal sclerite Adult. Characters unknown, only abdomen existing. complex composed of rod and ring structure, rod Male genitalia. Abdominal segment VIII short; very short, apically with pair of distinct, curved, lat- tergum about as long as sternum, unmodified; eral sclerites. sternum VIII without posteroventral projection. Blahnik et al.: Chimarra of northern Borneo 127

Abdominal segment IX relatively wide dorsolaterally, diagnostic character of this group is that the dor- tergum short dorsally, with pair of distinct apodemes somesal part of segment IX is obsolete or membra- from anterolateral margin; anteroventral margin nous, and there is no membranous mesal lobe of in lateral view distinctly produced, angular; poste- tergum X (Fig. 22B). Additionally, the ventral proc- rior margin slightly produced; ventral process pro- ess of segment IX is either short and deflected ven- jecting posteriorly, very elongate, nearly uniform trally (rather than posteriorly) (Fig. 15A) or absent in width, subtruncate apically. Preanal append- (Fig. 14A). A subgroup of the Chimarra tsudai age setose, short, rounded or subquadrate, some- Group, including many species from Borneo, has what flattened, slightly narrowed basally. Inferior tergum VIII highly modified and apically margined appendage about as long as tergum X; in lateral with spines or spine-like projections, thus super- view dorsally inflected basoventrally, widest basally, ficially resembling species in the subgenus Curgia narrow distally, basoventral margin with acute pro- (Fig. 16A, G). The forewing venation typically has jection, apex rounded; in ventral view somewhat the stem of Rs deflected (curved downward) and mesally curved, basally with strong, thumb-like the fork at the base of the discal cell more or less projection from posterior margin, mesal surface symmetrical (Figs 41–43). The forewing discal with tooth-like projection at midlength. Tergum cell is elongate (length often 3 times the width or X with sclerotized lateral lobes only, mesal mem- more) and both veins 2A and 3A are looped to 1A brane not evident, apparently lacking; lateral lobe (Figs 41–43). The latter character also occurs in the long, with projecting ventral margin, lobe bearing subgenera Curgia, Otarrha, and some Chimarrita, 2 sensilla, sensilla on slightly projecting, rounded as well as in species of the Chimarrhafra lineage of middorsal lobe, apex of ventral margin narrow, suba- Africa, but not typically in the Chimarra digitata cute. Phallobase tubular, with pronounced basod- group, in which 2A appears to be forked or Y-shaped orsal expansion, moderately elongate, ventral apex apically (Figs 39A, 40A). distinctly sclerotized, distinctly projecting, with Adult. Forewing Rs deflected or curved (rather than strongly deflexed, acute ventral process. Endotheca sinuous or S-shaped); basal fork of discal cell more elongate, tubular, with granularly textured region or less symmetrical; discal cell elongate, length often and 3 endothecal spines, 2 subequal, moderately 3 or more times width; m crossvein usually proxi- long basal spines and 1 very short, slender apical mal to s and r-m crossveins, sometimes nearly lin- spine. Phallotremal sclerite complex not evident in early arranged; s hyaline or not; 2A obsolete apically, specimen examined. apparently looped to 1A. ? with foretarsi and claws Etymology. This species is named Chimarra xiphosella, (probably) always enlarged. as a diminutive derived from the Greek word xiphos, Male genitalia (Fig. 15). Segment VIII unmodified a sword, and referring to the elongate ventral or modified; if modified, then often enlarged, with process of segment IX; to be treated as a noun in posteromesal margin invaginated and margined api- apposition. cally with projecting spine-like or scabrous processes. Segment IX with anteromesal margin slightly Chimarra tsudai group to distinctly projecting, usually rounded mesally (rather than sinuate); anterodorsal apodemes weakly The Chimarra tsudai group is more geographically developed or absent; dorsomesal margin obsolete, restricted than the Chimarra digitata group, ranging membranous; ventral process broad basally, short, from Pakistan to Japan and into Southeast Asia and ventrally deflected, absent in some species. Tergum parts of Indonesia. About 130 species, or over half X without membranous mesal lobe; lateral lobes the total Chimarra species from the Oriental region, divided into sclerotized lateral and mesal lobes, lat- can be confidently placed here. These are listed in eral lobes variable in shape, always with numerous Table 1. The most diagnostic attribute is that the lat- sensilla, mesal lobes membranous basally, forming eral lobes of tergum X of the male genitalia are sub- upturned digitate processes, sometimes with api- divided into sclerotized lateral and mesal lobes, with cal sensilla. Preanal appendages short, setose, often the mesal pair membranous basally and forming very small. Inferior appendages variable in length upright, digitate processes that straddle the dorsally and shape, usually relatively linear as viewed later- directed phallic apparatus (Fig. 15A, B). As noted ally, with little evidence of basal inflection; linear or by Ross (1956), the group is also characterized by mesally curved, as viewed ventrally, typically with having the lobes of tergum X with numerous sen- acute apical or preapical tooth (or teeth) or acute dor- silla (many more than 2). These are always located sal or dorsomesal projection. Phallic apparatus with on the lateral lobes (Fig. 14 A, B) and often on the phallobase tubular, long or short, ventral apex pro- sclerotized mesal lobes as well (Fig. 24A, B). Another jecting or not; endotheca variable in length, short or 128 Tijdschrift voor Entomologie, volume 152, 2009

Table 1. Chimarra tsudai group species.Placement in tsudai group not ascertainable from original description: Chimarra auronitens Ulmer, 1906; Chimarra pedalis Banks, 1931; Chimarra pilosella Navás, 1932; Chimarra tagalica Banks, 1937.

Chimarra aberrans Martynov, 1935 Chimarra jannekae Blahnik et al., sp. n. Chimarra alcicorne Malicky, 1995 Chimarra jaroschi Malicky, 1994 Chimarra aminadab Malicky, 1993 Chimarra jisipu Malicky, 1989 Chimarra ammi Malicky, 2008 Chimarra joliveti Jacquemart, 1979 Chimarra anam Malicky, 2008 Chimarra kailishchandrai Malicky, 1997 Chimarra aneca Malicky & Chantaramongkol, 1993 Chimarra karlijnae Blahnik et al., sp. n. Chimarra areli Malicky, 2008 Chimarra khasia Kimmins, 1957 Chimarra argeia Malicky & Chantaramongkol, 1997 Chimarra kinabaluensis Blahnik et al., sp. n. Chimarra arkit Malicky, 2008 Chimarra kumaonensis Martynov, 1935 Chimarra atara Malicky & Chantaramongkol, 1993 Chimarra lahuorum Chantaramongkol & Malicky, 1989 Chimarra atnia Malicky & Chantaramongkol, 1993 Chimarra lambi Blahnik et al., sp. n. Chimarra atripennis Banks, 1931 Chimarra lannaensis Chantaramongkol & Malicky, 1989 Chimarra auricoma Kimmins, 1957 Chimarra lichiuensis Hsu & Chen, 1996 Chimarra batukaua Malicky, 1995 Chimarra litugena Malicky & Chantaramongkol, 1993 Chimarra berenike Malicky, 1998 Chimarra litussa Malicky & Chantaramongkol, 1993 Chimarra biatec Malicky, 1993 Chimarra liwaguensis Blahnik et al., sp. n. Chimarra burmana Kimmins, 1957 Chimarra longispina Sun, 2007 Chimarra caduca Blahnik et al., sp. n. Chimarra maoi Sun & Malicky, 2002 Chimarra chanchuluni Blahnik et al., sp. n. Chimarra malaisei Kimmins, 1957 Chimarra concava Kimmins, 1957 Chimarra mars Malicky, 2007 Chimarra concolor Ulmer, 1905 Chimarra matura Malicky & Chantaramongkol, 1993 Chimarra confusa Ulmer, 1907 Chimarra megara Malicky, 2007 Chimarra crepidata Kimmins, 1957 Chimarra meorum Chantaramongkol & Malicky, 1989 Chimarra cumata Malicky & Chantaramongkol, 1993 Chimarra mlabriorum Chantaramongkol & Malicky, 1989 Chimarra cuspidata Blahnik et al., sp. n. Chimarra momma Malicky & Chantaramongkol, 1993 Chimarra cygnus Blahnik et al., sp. n. Chimarra mommaides Mey, 1998 Chimarra danumensis Blahnik et al., sp. n. Chimarra nahesson Malicky & Chantaramongkol, 1993 Chimarra denticula Blahnik et al., sp. n. Chimarra nepalensis Kimmins, 1964 Chimarra devva Malicky & Chantaramongkol, 1993 Chimarra nigra Kimmins, 1964 Chimarra dirke Malicky & Thamsenanupap, 2000 Chimarra nigrorosea Schmid, 1960 Chimarra discolor Kimmins, 1957 Chimarra noloyan Blahnik et al., sp. n. Chimarra excavata Kimmins, 1957 Chimarra nonna Malicky, 1993 Chimarra fansipangensis Mey, 1998 Chimarra nunenada Melnitsky, 2005 Chimarra fenestrata Kimmins, 1964 Chimarra opaca Mey, 1998 Chimarra flaviventris Kimmins, 1957 Chimarra oreithyia Malicky, 2007 Chimarra fluctuata Sun, 2007 Chimarra palawana Malicky, 1994 Chimarra fulmeki Ulmer, 1951 Chimarra pasiphae Malicky, 2007 Chimarra fusca Kimmins, 1957 Chimarra pataplan Malicky, 1989 Chimarra gemmal Malicky, 1989 Chimarra phillipsae Blahnik et al., sp. n. Chimarra gigama Malicky, 1989 Chimarra physanoton Blahnik et al., sp. n. Chimarra gunungkawi Malicky, 1995 Chimarra podarge Malicky & Thamsenanupap, 2007 Chimarra haimuoi Malicky, 1995 Chimarra pontos Malicky, 2007 Chimarra haimuoiba Malicky, 1995 Chimarra preapicalis Blahnik et al., sp. n. Chimarra haimuoibon Malicky, 1995 Chimarra prokrustes Malicky, 2008 Chimarra haimuoimot Malicky, 1995 Chimarra rama Malicky & Chantaramongkol, 1993 Chimarra haimuoinam Malicky, 1995 Chimarra ravanna Malicky & Chantaramongkol, 1993 Chimarra hezron Malicky, 1993 Chimarra schwendingeri Chantaramongkol & Malicky, Chimarra hoangliensis Mey, 2005 1989 Chimarra horok Malicky, 1989 Chimarra scolops Blahnik et al., sp. n. Chimarra houvichka Schmid, 1960 Chimarra scopulifera Kimmins, 1957 Chimarra htinorum Chantaramongkol & Malicky, 1989 Chimarra semiramis Malicky, 2007 Chimarra igvarvaria Melnitsky, 2005 Chimarra senticosa Sun & Malicky, 2002 Chimarra inthanonensis Chantaramongkol & Malicky, Chimarra shiva Malicky & Chantaramongkol, 1993 1989 Chimarra silausilau Blahnik et al., sp. n. Chimarra jacobsoni Ulmer, 1951 Chimarra sinitorum Blahnik et al., sp. n. Blahnik et al.: Chimarra of northern Borneo 129

Chimarra sita Malicky & Chantaramongkol, 1993 Chimarra thaumas Malicky, 2008 Chimarra spinifera Kimmins, 1957 Chimarra triangulata Hsu & Chen, 1996 Chimarra spitzeri Malicky, 1994 Chimarra tsudai Ross, 1956 Chimarra stenodactylus Blahnik et al., sp. n. Chimarra uncula Mey, 1998 Chimarra suadulla Malicky & Chantaramongkol, 1993 Chimarra uschtu Malicky, 1989 Chimarra supanna Malicky, 1993 Chimarra vantoli Blahnik et al., sp. n. Chimarra suryasena Schmid, 1960 Chimarra vanwelzeni Blahnik et al., sp. n. Chimarra suthepensis Chantaramongkol & Malicky, 1989 Chimarra vesta Malicky, 2007 Chimarra sythoffi Ulmer, 1951 Chimarra wushikangensis Hsu & Chen, 1996 Chimarra talos Malicky, 2007 Chimarra yaorum Chantaramongkol & Malicky, 1989

elongate, sometimes textured with small papillae or with highly modified, invaginated mesally and variously tracts of numerous short spines, variably positioned armed with spine-like projections. Generalized fea- in different species; phallotremal sclerite complex tures it possesses are a phallotremal sclerite that is not variable, either small, forming typical short rod and greatly enlarged, and an endotheca that is relatively ring structure, or enlarged and sclerotized, sometimes short and lacks distinctive spines. Species in this with accompanying dorsolateral or apical sclerites. group are all distinguished by characteristic differences in the shapes of the lateral and mesal lobes of Chimarra atripennis Banks tergum X and in the location of the mesal projection of the inferior appendages. In C. caduca, the mesal Chimarrha atripennis Banks, 1931: 425, holotype /, Ma- projection of the inferior appendages is located just laysia (Sabah), MCZT. past midlength. In this respect, it is similar to C. jannekae, sp. n. and C. cygnus, sp. n., both of which dif- This species was originally described from Mt. Kina- fer significantly in the shapes the lobes of tergum X, balu, based on specimens from Marei Parei, 5000 ft. as well as in details of the phallic apparatus. and Kiau, 3000 ft. Two specimens were indicated in Adult. Color (in alcohol) light brown. Length of the type series, but their sex was not stated. Also, forewing: male 5.5 mm. Forewing venation: stem of no illustration was provided. A specimen labeled as Rs only slightly curved; fork at base of discal cell not the holotype (MCZT 16506) was borrowed from thickened, nearly symmetrical, length of discal cell the Museum of Comparative Zoology, Harvard about 3 times width; m crossvein somewhat proxi- University, and examined by W. Mey (pers. com.). mal to crossveins s and r-m, crossvein s not hyaline; It is a female. The species is large (wing expanse 2A vein not intersecting 3A (apparently looped to 13 mm) and black in color. The description of the 1A). Postocular parietal sclerite moderately elongate. venation, Rs curved downward, discal cell elon- Maxillary palps moderately elongate, segment 3 sub- gate, makes it evident that this is a species in the equal to 2 and 5. Protarsal claws of male enlarged, Chimarra tsudai group. It is likely that it is one of the asymmetrical in shape and length. species described in this paper, but until females are Male genitalia. Abdominal segment VIII short; ter- definitively associated and described, its identity will gum about as long as sternum, modified, with small remain in doubt. mesal excavation, excavation bordered laterally with laciniate fringe, lateral fringe more elongate; sternum Chimarra caduca sp. n. VIII without posteroventral projection. Abdomi- nal segment IX short dorsolaterally; anteroventral Fig. 13 margin in lateral view slightly produced, rounded; posterior margin sinuously produced; ventral proc- Type material. Holotype: ?, Malaysia, Sabah, ess projecting ventrally, short, broad basally, broadly Crocker Range, 40 km S of Kota Kinabalu, Sin- rounded. Preanal appendage setose, short, rounded, suron Rd., 1500 m, 19.xii.1989, J. Huisman broadest basally. Inferior appendage about as long as (UMSP000107300) (RMNH). tergum X; in lateral view straight, nearly uniformly Paratypes. Malaysia, Sabah, Kinabalu National Park, wide, apex rounded; in ventral view nearly straight, headquarters, 1500 m, 10.viii.2005, W. Mey, 4? dorsomesally with broad, distinctly sclerotized, api- (UMHU). cally acute projection just beyond midlength. Ter- gum X with sclerotized lateral lobes and separate Chimarra caduca also belongs to the subgroup of mesal lobes, mesal membrane lacking; lateral lobe the Chimarra tsudai group in which tergum VIII is long, in lateral view narrow, apex cupped, slightly 130 Tijdschrift voor Entomologie, volume 152, 2009

a b c d

e f g

Fig. 13. Chimarra caduca, male genitalia. – a, lateral; b, segment IX and tergum X, dorsal; c, inferior appendage, ventral; d, inferior appendage, dorsal; e, phallic apparatus, lateral; f, inferior appendage, caudal; f, tergum VIII, dorsal.

downcurved, lobe bearing multiple sensilla; mesal Chimarra chanchuluni is a distinctive species with a lobe elongate, digitate, reclinate, posteriorly directed. number of diagnostic characters, including the shape Phallobase tubular, with pronounced basodorsal of the inferior appendages, which are somewhat expansion, very short. Endotheca relatively short, dorsoventrally flattened, mesally curved, as viewed granularly textured region not discernable, without ventrally, and have a small spine-like, preapical endothecal spines. Phallotremal sclerite composed of projection on the mesal surface. Other diagnostic rod and ring structure, rod very short, apically with characters include the shape of the mesal lobes of pair of short, curved, lateral sclerites. tergum X (each of which is divided into a pair of Etymology. This species is named Chimarra caduca sclerotized, spine-like processes), the elongate pre- from the Latin word caducus, meaning “that falls or anal appendages, and the form of the phallic appara- is inclined to fall”, in reference to the relatively pros- tus, which is laterally compressed apically, and lacks trate mesal lobes of tergum X of this species. endothecal spines. Adult. Color (in alcohol) yellowish-brown. Length Chimarra chanchuluni sp. n. of forewing: male 5.2–6.8 mm, female 5.8–7.2 mm. Forewing venation: stem of Rs curved, slightly scle- Fig. 14 rotized and thickened at inflection; fork at base of discal cell not thickened, nearly symmetrical, length Type material. Holotype: ?, Malaysia, Sabah, of discal cell about 4 times width; m crossvein proxi- Kinabalu National Park, Headquarters, Sg. Liwagu mal to crossveins s and r-m, crossvein s hyaline; 2A @ Silau-Silau trail, 06°00’N, 116°33’E, 1470 m, vein not intersecting 3A (apparently looped to 1A). 22.i.1987, Huisman & de Jong (UMSP000107181) Postocular parietal sclerite moderately elongate. Max- (RMNH). illary palps elongate, segment 3 slightly greater than Paratypes. Malaysia, Sabah, same data as holotype, 2, 5 shorter than 3. Protarsal claws of male enlarged, 1?, 4/ (RMNH); Crocker Range, 5 km N Tenom, asymmetrical in shape and length. base of Sg. ulu Noloyan, 05°10’N, 115°56’E, 1010 m, Male genitalia. Abdominal segment VIII short; 10–11.x.1986, J. Huisman, 6? (UMSP). tergum longer than sternum, unmodified, with Blahnik et al.: Chimarra of northern Borneo 131 a b c d

e g

Fig. 14. Chimarra chanchuluni, male genitalia. – a, lateral; b, segment IX and tergum X, dorsal; c, inferior appendage, ventral; d, inferior appendage, dorsal; e, phallic apparatus, lateral; f, phallic apparatus, dorsal; g, apex of expanded endotheca, lateral.

V-shaped mesal depression; sternum VIII with slight bearing minute spinules; other endothecal spines posteroventral projection. Abdominal segment IX absent. Phallotremal sclerite complex composed of relatively wide dorsolaterally, tergum obsolete dor- rod and ring structure, rod moderately elongate. somesally, without distinct anterolateral apodemes; Etymology. Named for Chan Chu Lun, friend of anteroventral margin in lateral view not, or scarcely J. Huisman. produced; posterior margin sinuously produced; ventral process absent or nearly so. Preanal append- Chimarra cuspidata sp. n. age setose, moderately elongate, broadest basally, Fig. 15 tapering and rounded apically. Inferior appendage slightly longer than tergum X; in lateral view straight, nearly uniformly wide, apex rounded; in Type material. Holotype: ?, Malaysia, Sabah, ventral view somewhat mesally curved, preapically 60 km W Lahad Datu, Danum Valley Field Cen- with small, acute, mesal projection. Tergum X with tre, Sg. Palum Tambun & vicinity, 04°58’N, sclerotized lateral and mesal lobes, mesal membrane 117°48’E, 150 m, 19–26.iii.1987, J. Huisman lacking; lateral lobe moderately long, in lateral (UMSP000107351) (RMNH). view narrow, apex slightly downcurved, lobe bear- Paratypes. Malaysia, Sarawak, Annah Rais. ing multiple sensilla; mesal lobe short, distinctly Sg. Semadang, 800 m, 27.x.2003, W. Mey, 1? sclerotized, bifid, both branches strongly hooked, (UMHU); Sabah, Crocker Range National Park, mesal branch dorsally directed, lateral branch later- Sg. Myoog, 9.xi.2003, W. Mey, 9? (UMHU); Kina- ally directed. Phallobase tubular, with pronounced balu National Park, Poring Hot Springs, 30.x.2003, basodorsal expansion, moderately elongate, relatively W. Mey, 2? (UMHU); Lohan near Ranau, narrow, laterally compressed apically, dorsal margin Sg. Lohan, 7.xi.2003, W. Mey, 5? (UMHU); 60 km extended apically, elongate, narrow, lightly sclero- W of Lahad Datu, DVFC, brooklet above lotus tized. Endotheca relatively short, dorsally with bulg- pond, 04°58’N, 117°48’E, 150 m, 19–28.x.1987, ing tract of minute spines, apicoventrally with lobe Huisman & de Jong, 1? (RMNH); 12 km NNE 132 Tijdschrift voor Entomologie, volume 152, 2009

a b c d tergum VIII sclerotized mesal lobe preanal of tergum X appendage sclerotized lateral lobe of tergum X

sensillla

segment IX

e inferior appendage

ventral process

g h

Fig. 15. Chimarra cuspidata, male genitalia. – a, lateral; b, segment IX and tergum X, dorsal; c, inferior appendage, ventral; d, inferior appendage, dorsal; e, phallic apparatus, lateral; f, apex of expanded endotheca, lateral; g, inferior appendage, caudal; h, tergum VIII, dorsal.

Ranau, Poring Hot Spring, staff quarters, 06°03’N, to crossveins s and r-m, crossvein s not hyaline; 2A 116°42’E, 550 m, 1.ii.1987, Huisman & de Jong, 3? vein not intersecting 3A (apparently looped to 1A). (UMSP); Kinabalu National Park, Poring, 570 m, Postocular parietal sclerite short. Maxillary palps 8.ix.1983, Hevel & Steiner, 2?, 5/ (NMNH). relatively short, segment 3 distinctly longer than 2, 3 slightly longer than 5. Protarsal claws of male Chimarra cuspidata is related to those species of the enlarged, asymmetrical in shape, scarcely different in Chimarra tsudai group with a distinctly modified, length. mesally invaginated, spinose tergum VIII. In C. cus- Male genitalia. Abdominal segment VIII short; ter- pidata, tergum VIII is relatively short, and the mesal gum about as long as sternum, modified, with invagination V-shaped and margined by spines of a V-shaped mesal excavation, excavation with short uniform length. It is easily diagnosed from any other laciniate fringe; sternum VIII without posteroventral members of this group by the shape of the lateral projection. Abdominal segment IX very short dorso- lobe of tergum X, which is distinctly inflated preapi- laterally, tergum obsolete dorsomesally, without dis- cally, as viewed laterally, and by the distinctive api- tinct anterolateral apodemes; anteroventral margin comesal process on the inferior appendages, which is in lateral view not, or scarcely produced; posterior truncately cusped or weakly bicusped. margin sinuously produced; ventral process project- Adult. Color (in alcohol) dark brown. Length of ing ventrally, short, broad basally, subtriangular. forewing: male 4.3–4.9 mm. Forewing venation: Preanal appendage setose, very small, rounded. Infe- stem of Rs curved, distinctly sclerotized and thick- rior appendage about as long as tergum X; in lateral ened at inflection; fork at base of discal cell some- view straight, nearly uniformly wide, apex rounded; what thickened, nearly symmetrical, length of dis- in ventral view nearly straight, apically with discal cell about 3 times width; m crossvein proximal tinctly sclerotized, weakly bicuspidate projection. Blahnik et al.: Chimarra of northern Borneo 133 a b c d

f g h

Fig. 16. Chimarra cygnus, male genitalia. – a, lateral; b, segment IX and tergum X, dorsal; c, inferior appendage, ventral; d, inferior appendage, dorsal; e, phallic apparatus, lateral; inset, apex of phallus, dorsal; f, variant form, detail of lateral; g, tergum VIII, dorsal; h, variant form, tergum VIII, dorsal.

Tergum X with sclerotized lateral lobes and separate 7.xi.1987, Huisman & de Jong (UMSP000107285) mesal lobes, mesal membrane lacking; lateral lobe (RMNH). long, in lateral view inflated at midlength, apex nar- Paratypes. Malaysia, Sabah, same data as holotype, row, bearing multiple sensilla; mesal lobe elongate, 1?, 1/ (UMSP); Kinabalu National Park, Head- digitate, posterodorsally curved. Phallobase tubular, quarters, Sg. Silau-Silau, 11.viii.2005, W. Mey, 2?, with pronounced basodorsal expansion, relatively 1/ (UMHU); Kinabalu National Park, tributary short, ventral apex distinctly sclerotized, distinctly to Sg. Liwagu, 1700 m, 26.ii.2006, W. Mey, 1? projecting, subacute, ventrally deflexed. Endotheca (UMHU); Kinabalu National Park, Headquarters, relatively short, with minutely spinulose region Sg. Tibabar, 06°02’N, 116°33’E, 1750 m, 12.i.1987, preapically, endothecal spines absent. Phallotremal J. Huisman, 1? (RMNH); Kinabalu National Park, sclerite complex composed of rod and ring structure, Headquarters, Sg. Liwagu, 1500 m, 10.viii.2005, rod moderately elongate. W. Mey, 1? (UMHU); Kinabalu National Park, Etymology. This species is named Chimarra cuspi- Sungai Mesilau, 1800 m, 12.viii.2005, W. Mey, data, for the cusped apicomesal projections on the 11?, 3/ (UMHU); Crocker Range, 5 km N inferior appendage. Tenom, base of Sg. ulu Noloyan, 05°10’N, 115°56’E, 1010, 10–11.x.1986, J. Huisman, 4? (RMNH); Chimarra cygnus sp. n. Kinabalu National Park, 12 km NNW Kundassang, Marei-Parei, trickle, 06°05’N, 116°31’E, 1670 m, Fig. 16 9.iii.1987, J. Huisman, 4?, 3/ (UMSP).

Type material. Holotype: ?, Malaysia, Sabah, Kina- Chimarra cygnus is related to those species of the balu Park, Carson’s Fall, 06°02’N, 116°33’E, 1900 m, Chimarra tsudai group with a modified, mesally 134 Tijdschrift voor Entomologie, volume 152, 2009

invaginated, spinose tergum VIII. It is most readily slightly narrowed basally. Inferior appendage slightly diagnosed by the characteristic shape of the mesal longer than tergum X; in lateral view straight, nearly lobes of tergum X, which are elongate, digitate, uniformly wide, apex rounded; in ventral view nearly and curved in a somewhat swan-neck like manner. straight, preapically with prominent, mesal, tooth- Useful ancillary characters for diagnosing this spe- like projection. Tergum X with sclerotized lateral lobes cies are the shape of the inferior appendage, which and separate mesal lobes, mesal membrane lacking; has a prominent, acute mesal projection preapically, lateral lobe moderately long, in lateral view simple, and the shape of the lateral lobes of tergum X, each rounded apically, with slight to distinct basodorsal of which is moderately elongate and has a rounded projection, lobe bearing multiple sensilla; mesal lobe basal projection. A distinctive variant of this species very elongate, narrow, digitate, strongly posterodor- is illustrated in Figs 16F and 16H. The holotype sally curved. Phallobase tubular, with pronounced is the form in Fig. 16A-E, G. Differences between basodorsal expansion, short, ventral apex distinctly these two forms include the spinose development of sclerotized, distinctly projecting, forming laterally tergum VIII, the development of the ventral apex compressed process, variable in shape. Endotheca of the phallobase, and shape of the basal projec- relatively short, granularly textured region not dis- tion of the lateral lobe of tergum X. However, the cernable; without endothecal spines. Phallotremal overall differences are minor. We believe this varia- sclerite complex composed of rod and ring structure, tion to be intraspecific. Specimens from the Crocker rod moderately elongate, dorsolaterally, with pair of Range were distinctly smaller in size and account elongate curved sclerites. for the wide variation in size found in this species. Etymology. This species is called Chimarra cygnus, The closest relationship of C. cygnus is probably to from the Latin cygnus, a swan, used as a noun in C. lambi, sp. n. and C. danumensis, sp. n., both of apposition, and referring to the swan-necked shape which also have elongate phallotremal sclerites with of the mesal processes of tergum X in this species. accompanying dorsolateral sclerites. However, both of those species have the mesal projection of the infe- Chimarra danumensis sp. n. rior appendages very differently formed. The shape Fig. 17 of the inferior appendages of C. cygnus is probably most similar to C. jannekae, sp. n. and C caduca, sp. n., both of which differ significantly in the shape Type material. Holotype: ?, Malaysia, Sabah, and orientation of the lateral and mesal lobes of 60 km W Lahad Datu, Danum Valley Field Cen- tergum X. tre, Sg. Palum Tambun & vicinity, 04°58’N, Adult. Color (in alcohol) brown. Length of forewing: 117°48’E, 150 m, 19–26.iii.1987, J. Huisman male 5.4–7.4 mm. Forewing venation: stem of Rs (UMSP000107255) (RMNH). curved, slightly sclerotized and thickened at inflec- Paratypes. Malaysia, Sabah, same data as holotype, tion; fork at base of discal cell distinctly thickened, 4? (UMSP); Danum Valley Field Centre, Sg. Palun nearly symmetrical, length of discal cell about 3 times Tambun, 18.viii.2005, W. Mey, 3? (UMHU); width; m crossvein proximal to crossveins s and r-m, 60 km W Lahad Datu, Danum Valley Field Cen- crossvein s not hyaline; 2A vein not intersecting 3A tre, nr. Segama bridge, 04°58’N, 117°48’E, 150 m, (apparently looped to 1A). Postocular parietal scle- 20.x.1987, Huisman & de Jong, 1? (UMSP). rite moderately elongate. Maxillary palps elongate, segment 3 slightly greater than 2, 3 subequal to 5. Chimarra danumensis is related to those species of Protarsal claws of male enlarged, asymmetrical in the Chimarra tsudai group with a modified, mesally shape and length. invaginated, spinose tergum VIII. The closest rela- Male genitalia. Abdominal segment VIII short; ter- tionship of C. danumensis is probably to C. lambi, gum about as long as sternum, modified, with mesal sp. n. and C. cygnus, sp. n, both of which also have excavation, excavation bordered posterolaterally with an elongate phallotremal sclerite with accompanying short laciniate fringe, or sometimes an elongate, dorsolateral sclerites. In C. danumensis this structure bifurcating process with marginal spines; sternum is very elongate, nearly the length of the phallobase. VIII without posteroventral projection. Abdominal In overall features, C. danumensis is probably most segment IX relatively wide dorsolaterally, tergum similar to C. lambi, but can be easily distinguished obsolete dorsomesally, without distinct anterola- by the diagnostic bicusped projection from the mesal teral apodemes; anteroventral margin in lateral view surface of the inferior appendages, which in C. danu- slightly produced, rounded; posterior margin sinu- mensis is located preapically rather than apically. ously produced; ventral process projecting ventrally, Adult. Color (in alcohol) brown. Length of fore- very short. Preanal appendage setose, small, rounded, wing: male 4.6–5.0 mm. Forewing venation: stem Blahnik et al.: Chimarra of northern Borneo 135 a b c d

f e

Fig. 17. Chimarra danumensis, male genitalia. – a, lateral; b, segment IX and tergum X, dorsal; c, inferior appendage, ventral; d, inferior appendage, dorsal; e, phallic apparatus, lateral; inset, phallotremal sclerite, dorsal; f, tergum VIII, dorsal. of Rs curved, slightly sclerotized and thickened at sclerotized lateral lobes and separate mesal lobes, inflection; fork at base of discal cell not thickened, mesal membrane lacking; lateral lobe moderately nearly symmetrical, length of discal cell about 3 long, in lateral view narrow, parallel sided, rounded times width; m crossvein proximal to crossveins s and apically, lobe bearing multiple sensilla; mesal lobe r-m, crossvein s hyaline; 2A vein not intersecting 3A elongate, digitate, posterodorsally directed. Phal- (apparently looped to 1A). Postocular parietal scle- lobase tubular, with pronounced basodorsal expan- rite short. Maxillary palps relatively short, segment sion, moderately elongate. Endotheca length not 3 slightly longer than 2, 3 subequal to 5. Protarsal discernable (not expanded); without endothecal claws of male enlarged, asymmetrical in shape and spines. Phallotremal sclerite complex composed of length. rod and ring structure, rod elongate, dorsolaterally, Male genitalia. Abdominal segment VIII short; ter- with pair of elongate curved sclerites. gum distinctly longer than sternum, modified, with Etymology. This species is named Chimarra danu- broad, subquadrate mesal excavation, excavation api- mensis after the Danum Valley Field Centre, where colaterally with laciniate fringe; sternum VIII with- the holotype specimen was collected. out posteroventral projection. Abdominal segment IX relatively wide dorsolaterally, tergum obsolete Chimarra denticula sp. n. dorsomesally, without distinct anterolateral apo- Fig. 18 demes; anteroventral margin in lateral view not, or scarcely produced; posterior margin distinctly produced; ventral process projecting ventrally, broad Type material. Holotype: ?, Malaysia, Sabah, Crocker basally, subtriangular. Preanal appendage setose, Range, 5 km N Tenom, base of Sg. ulu Noloyan, short, rounded, broadest basally. Inferior appendage 05°10’N, 115°56’E, 1010 m, 10–11.x.1986, J. Huis- about as long as tergum X; in lateral view straight, man (UMSP000107331) (RMNH). nearly uniformly wide, apex rounded; in ventral view Paratypes. Malaysia, Sabah, same data as holotype, somewhat mesally curved, preapically with distinctly 4? (UMSP). sclerotized, bicuspidate projection. Tergum X with 136 Tijdschrift voor Entomologie, volume 152, 2009

a b c d

Fig. 18. Chimarra denticula, male genitalia. – a, lateral; b, segment IX and tergum X, dorsal; c, inferior appendage, ventral; d, inferior appendage, dorsal; e, phallic apparatus, lateral.

Chimarra denticula resembles those species of the stem of Rs curved, slightly sclerotized and thickened Chimarra tsudai group from Sabah without a modi- at inflection; fork at base of discal cell somewhat fied tergum VIII, but with an acute projection or thickened, nearly symmetrical, length of discal cell tooth on the dorsal margin of the inferior appendages, about 3 times width; m crossvein proximal to cross- evident in lateral view (including C. karlijnae, sp. n., veins s and r-m, crossvein s not hyaline; 2A vein not C. prokrustes Malicky, C. scolops, sp. n., C. sinitorum, intersecting 3A (apparently looped to 1A). Posto- sp. n., and C. stenodactylus, sp. n.). All of these species cular parietal sclerite moderately elongate. Maxillary also have a phallus with numerous small spines and palps relatively short, segment 3 slightly longer than a narrow sclerotized dorsal extension, which extends 2, 2 subequal to 5. Protarsal claws of male enlarged, over the basal part of the endotheca. They are addi- asymmetrical in shape and length. tionally characterized by a mesal lobe of tergum X Male genitalia. Abdominal segment VIII moderate that is distinctly sclerotized and with an apex that in length; tergum longer than sternum, unmodified, is acute and dorsally curved. Among these species, entire; sternum VIII without posteroventral projec- C. denticula is most easily diagnosed by the short and tion. Abdominal segment IX relatively wide dorsola- distinctively shaped lateral and mesal lobes of tergum terally, tergum obsolete dorsomesally, without distinct X. It is also distinctive in having an additional short anterolateral apodemes; anteroventral margin in lat- acute projection or tooth located preapically on the eral view not, or scarcely produced; posterior margin mesal surface of each inferior appendage. Chimarra distinctly produced; ventral process absent or nearly stenodactylus also has short lateral and mesal lobes of so. Preanal appendage setose, very small, rounded. tergum X, but these are differently shaped, and the Inferior appendage longer than tergum X; in lateral base of the dorsal projection on the inferior append- view straight, nearly uniformly wide, apex rounded, age of this species is very narrow. dorsal margin with basally wide, acute projection Adult. Color (in alcohol) yellowish-brown. Length at midlength; in ventral view somewhat mesally of forewing: male 4.6–5.5 mm. Forewing venation: curved, mesal surface with small, preapical tooth-like Blahnik et al.: Chimarra of northern Borneo 137 a b c d

f e

Fig. 19. Chimarra jannekae, male genitalia. – a, lateral; b, segment IX and tergum X, dorsal; c, inferior appendage, ventral; d, inferior appendage, dorsal; e, phallic apparatus, lateral; f, tergum VIII, dorsal.

Chimarra jannekae sp. n. projection. Tergum X with sclerotized lateral lobes Figs 19, 41 and separate mesal lobes, mesal membrane lacking; lateral lobe short, in lateral view simple, rounded apically, distinctly downcurved, lobe bearing mul- Type material. Holotype: ?, Malaysia, Sabah, Crocker tiple sensilla; mesal lobe short, digitate, distinctly Range, 40 km S of Kota Kinabalu, Sinsuron Rd., sclerotized, dorsally directed, apex with approxi- 1500, 20.xii.1989, J. Huisman, (UMSP000107161) mately 2 spine-like projections. Phallobase tubular, (RMNH). with pronounced basodorsal expansion, moderately Paratypes. Malaysia, Sabah, same data as holotype, elongate, relatively narrow, ventral apex lightly scle- 7? (UMSP); Kinabalu National Park, Sg. Liwagu, rotized, projecting, dorsal margin extended apically, 1100 m, 12.viii.2005, W. Mey, 2? (UMHU); Kina- elongate, narrow, lightly sclerotized. Endotheca balu National Park, Sg. Liwagu, 1500 m, 10.viii.2005, elongate, tubular; with several tracks of small W. Mey, 1? (UMHU); Kinabalu National Park, endothecal spines, basal track of few scattered spines, tributary to Sg. Liwagu, 1200 m, 13.viii.2005, dorsal track of very short spines at midlength, lateral W. Mey, 1? (UMHU); Kinabalu National Park, track of small spines at midlength, and apical track Sg. Tibabar @ Liwagu trail, 06°02’N, 116°33’E, of slightly longer spines. Phallotremal sclerite com- 1750 m, 2.x.1986, J. Huisman, 1? (RMNH); plex composed of rod and ring structure, rod short. Kinabalu National Park, Headquarters area, 1560 m, Etymology. This species is named Chimarra denticula, 13.ix.1983, Hevel & Steiner, 2? (NMNH). as a diminutive form derived from the Latin word dens, a tooth, treated as a noun in apposition, and Chimarra jannekae belongs to a distinctive subgroup referring to the distinctive small preapical projection of the Chimarra tsudai group in which tergum VIII on the mesal surface of the inferior appendage in this is highly modified, invaginated mesally and variously species. armed with spine-like projections. Among species from Sabah, it resembles C. caduca, sp. n and C. cygnus, 138 Tijdschrift voor Entomologie, volume 152, 2009

a b c d

Fig. 20. Chimarra karlijnae, male genitalia. – a, lateral; b, segment IX and tergum X, dorsal; c, inferior appendage, ventral; d, inferior appendage, dorsal; e, phallic apparatus, lateral; f, phallic apparatus, dorsal; g, expanded endotheca, lateral.

sp. n. in the shape of the inferior appendages. Each apicolaterally with laciniate fringe, lateral margin inferior appendage in all of these species has a mesal forming downturned digitate projection. Preanal spine-like process at just past midlength. Addition- appendage setose, very small, rounded. Inferior ally, the phallic apparatus in all of these species lacks appendage about as long as tergum X; in lateral accessory spines. Diagnostic features of C. jannekae view straight, nearly uniformly wide, apex rounded; include the very elongate lateral lobes of tergum X in ventral view nearly straight, mesal surface with and the elongate, downcurved and distinctly sclero- tooth-like projection just beyond midlength. Ter- tized apex of the phallobase. gum X with sclerotized lateral lobes and separate Adult. Color (in alcohol) brown. Length of fore- mesal lobes, mesal membrane lacking; lateral lobe wing: male 6.0–6.6 mm. Forewing venation: stem long, in lateral view narrow, apex slightly ventrome- of Rs curved, slightly sclerotized and thickened sally curved, left and right lobes slightly asymmetri- at inflection; fork at base of discal cell somewhat cal; mesal lobe elongate, digitate, posterolaterally thickened, nearly symmetrical, length of discal cell directed, apex acute. Phallobase tubular, with pro- about 3 times width; m crossvein proximal to cross- nounced basodorsal expansion, moderately elongate, veins s and r-m, crossvein s not hyaline; 2A vein not ventral apex distinctly sclerotized, distinctly project- intersecting 3A (apparently looped to 1A). Postocular ing, subacute, strongly ventrally deflexed. Endotheca parietal sclerite moderately elongate. Maxillary palps relatively short, with granularly textured region, moderately elongate, segment 3 slightly greater than without endothecal spines. Phallotremal sclerite 2, 3 subequal to 5. Protarsal claws of male enlarged, complex composed of rod and ring structure, rod asymmetrical in shape and length. moderately elongate, with apicolateral sclerites. Male genitalia. Abdominal segment VIII short; Etymology. Named for Janneke Holzenthal, young- tergum distinctly longer than sternum, modified, est daughter of J. Huisman and R. Holzenthal. with broad, U-shaped mesal excavation, excavation Blahnik et al.: Chimarra of northern Borneo 139

Chimarra karlijnae sp. n. claws of male enlarged, asymmetrical in shape and Fig. 20 length. Male genitalia. Abdominal segment VIII moderate Type material. Holotype: ?, Malaysia, Sabah, 12 km in length; tergum longer than sternum, unmodi- NNE Ranau, Poring Hot Spring, Sg. Kepungit II, fied, with V-shaped mesal depression; sternum 06°04’N, 116°41’E, 700 m, 26.i.1987, J. Huisman VIII without posteroventral projection. Abdominal (UMSP000107256) (RMNH). segment IX relatively wide dorsolaterally, tergum Paratypes. Malaysia, Sabah, same data as holotype, obsolete dorsomesally, without distinct anterolateral 5/ (UMSP); Crocker Range, 5km N Tenom, base apodemes; anteroventral margin in lateral view of Sg. ulu Noloyan, 05°10’N, 115°56’E, 1010 m, slightly produced; posterior margin slightly produced; 10–11.x.1986, J. Huisman, 1? (RMNH); Por- ventral process absent or nearly so. Preanal append- ing Hot Spring, 12 km NNE Ranau, Sg. Kepungit age setose, very small, rounded. Inferior appendage waterfall, 06°03’N, 116°42’E, 550 m, 4.xii.1986, much longer than tergum X; in lateral view straight, J. Huisman, 1? (UMSP); 18 km on road Kenin- widest mesally, narrow distally, apex rounded, dorsal gau-Kimanis, gravel quarry, 05°26’N, 116°05’E, margin with very basally wide, apically acute projec- 1050 m, 20.i.1987, J. Huisman, 3? (UMSP); tion at midlength, the projection directed posteri- 12 km NNE Ranau, Poring Hot Spring, Sg. Tan- orly and imparting a crescentic appearance to apex anansad, 06°03’N, 116°42’E, 560 m, 28.viii.1986, of inferior appendage; in ventral view somewhat J. Huisman, 1? (RMNH); Poring Hot Spring, mesally curved, mesal surface without processes. 12 km NNE Ranau, Sg. Kipogoh, 06°03’N, Tergum X with sclerotized lateral lobes and separate 116°42’E, 550 m, 9.xii.1986, J. Huisman, 3? mesal lobes, mesal membrane lacking; lateral lobe (UMSP); 12 km NNE Ranau, Poring Hot Spring, short, in lateral view simple, rounded apically, lobe staff quarters, 06°03’N, 116°42’E, 550 m, 1.ii.1987, bearing multiple sensilla; mesal lobe short, digitate, Huisman & de Jong, 1? (RMNH). distinctly sclerotized, dorsally directed, with basal sensillate lobe and dorsally hooked apical projection. Chimarra karlijnae resembles those species of the Phallobase tubular, with pronounced basodorsal Chimarra tsudai group from Sabah that lack a modi- expansion, moderately elongate, relatively narrow, fied tergum VIII and have an acute projection or ventral apex lightly sclerotized, slightly projecting, tooth on the dorsal margin of the inferior append- dorsal margin extended apically, elongate, narrow, ages, evident in lateral view (including C. denticula, lightly sclerotized. Endotheca relatively short, with sp. n., C. prokrustes Malicky, C. scolops, sp. n., basal tract of short spines and also apicoventral tract C. sinitorum, sp. n., and C. stenodactylus, sp. n.). All of short spines. Phallotremal sclerite complex com- of these species also have a phallus with numerous posed of rod and ring structure, rod relatively short. small spines and a narrow sclerotized dorsal exten- Etymology. Named for Karlijn Holzenthal, oldest sion that extends over the basal part of the endotheca. daughter of J. Huisman and R. Holzenthal. They are additionally characterized by a mesal lobe of tergum X that is distinctly sclerotized and with an Chimarra kinabaluensis sp. n. apex that is acute and dorsally curved. Among these Fig. 21 species, C. karlijnae is easily diagnosed by the shape of the inferior appendage, which has the dorsal process enlarged, posteriorly directed and subequal in Type material. Holotype: ?, Malaysia, Sabah, length and development to the ventral margin, thus Kinabalu National Park, Headquarters, confluence producing a distinctive C-shaped apex to the inferior Sg. Liwagu & Sg. Silau-Silau, 06°00’N, 116°33’E, appendage. 1450 m, 27.vii.1986, J. Huisman (UMSP000107146) Adult. Color (in alcohol) light brown. Length of (RMNH). forewing: male 4.9–5.6 mm, female 5.0–5.8 mm. Paratypes. Malaysia, Sabah, Kinabalu National Forewing venation: stem of Rs curved, slightly scle- Park, Headquarters, Sg. Liwagu @ Silau-Silau trail, rotized and thickened at inflection; fork at base of 06°00’N, 116°33’E, 1470 m, 24.viii.1986, J. Huis- discal cell somewhat thickened, nearly symmetrical, man, 1? (UMSP); same locality, 10.i.1987, J. Huis- length of discal cell about 4 times width; m crossvein man, 1? (RMNH); Kundassang, Sg. Mesilau, 1st proximal to crossveins s and r-m, crossvein s not hya- bridge after golf course, 06°01’N, 116°36’E, 1650 m, line; 2A vein not intersecting 3A (apparently looped 3.x.1986, J. Huisman, 2? (UMSP). to 1A). Postocular parietal sclerite moderately elongate. Maxillary palps moderately elongate, segment Chimarra kinabaluensis is another species belonging to 3 slightly greater than 2, 3 subequal to 5. Protarsal the subgroup of the Chimarra tsudai group in which 140 Tijdschrift voor Entomologie, volume 152, 2009

a b c d

Fig. 21. Chimarra kinabaluensis, male genitalia. – a, lateral; b, segment IX and tergum X, dorsal; c, inferior appendage, ventral; d, inferior appendage, dorsal; e, phallic apparatus, lateral; f, tergum VIII, dorsal.

tergum VIII is highly modified, invaginated mesally iniate fringe; sternum VIII without posteroventral and variously armed with spine-like projections. It is projection. Abdominal segment IX relatively wide perhaps most similar to C. preapicalis, sp. n., which has dorsolaterally, tergum obsolete dorsomesally, with- a similarly shaped segment IX. The most useful diag- out distinct anterolateral apodemes; anteroventral nostic character is the shape of the inferior append- margin in lateral view slightly produced; posterior ages, which are very short, and possess an acute mesal margin slightly produced; ventral process projecting projection just preapically on the mesal margin. The ventrally, very short, broad basally, broadly rounded. greatly projecting tergum VIII, bordered by uniformly Preanal appendage setose, very small, rounded. Infe- sized spines is also a useful diagnostic character. Other rior appendage short, about as long as tergum X; in features it possesses a phallotremal sclerite that is not lateral view straight, widest basally, gradually nar- greatly enlarged, and an endotheca that is relatively rowing distally, apex rounded; in ventral view nearly short and lacks distinctive spines. straight, preapically with very prominent, basally Adult. Color (in alcohol) light brown. Length of broad, ventromesal, tooth-like projection. Tergum forewing: male 5.0–6.4 mm. Forewing venation: X with sclerotized lateral lobes and separate mesal stem of Rs curved, slightly sclerotized and thickened lobes, mesal membrane lacking; lateral lobe moder- at inflection; fork at base of discal cell not thickened, ately long, in lateral view simple, rounded apically, nearly symmetrical, length of discal cell about 3 times lobe bearing multiple sensilla; mesal lobe elongate, width; m crossvein somewhat proximal to crossveins s digitate, slightly posterodorsally curved. Phallobase and r-m, crossvein s not hyaline; 2A vein not intersect- tubular, with pronounced basodorsal expansion, ing 3A (apparently looped to 1A). Postocular parietal short. Endotheca relatively short, granularly textured sclerite moderately elongate. Maxillary palps elongate, region not discernable; without endothecal spines. segment 3 subequal to 2 and 5. Protarsal claws of male Phallotremal sclerite complex composed of elongate enlarged, asymmetrical in shape and length. ring, rod very reduced, preapically with short lateral Male genitalia. Abdominal segment VIII moderate sclerites. in length; tergum distinctly longer than sternum, Etymology. This species is named Chimarra kinabalu- modified, with V-shaped mesal excavation, exca- ensis for Kinabalu National Park, where the holotype vation bordered posterolaterally by spine-like lac- specimen was collected. Blahnik et al.: Chimarra of northern Borneo 141 a b c d

e f

h g

Fig. 22. Chimarra lambi, male genitalia. – a, lateral; b, segment IX and tergum X, dorsal; c, inferior appendage, ventral; d, inferior appendage, dorsal; e, inferior appendage, oblique caudal; f, phallic apparatus, lateral; g, phallic apparatus, dorsal; h, tergum VIII, dorsal.

Chimarra lambi sp. n. Chimarra lambi is another species belonging to the Fig. 22 distinctive subgroup of the Chimarra tsudai group in which tergum VIII is deeply invaginated mesally Type material. Holotype: ?, Malaysia, Sabah, Por- and variously armed with spine-like projections. ing Hot Spring, 12 km NNE Ranau, Sg. Kipogoh, It is related to subgroup within this latter group in 06°03’N, 116°42’E, 550 m, 1.ii.1987, J. Huisman which the phallotremal sclerite complex is greatly (UMSP000107253) (RMNH). enlarged and sclerotized and has elongate dorsola- Paratypes. Malaysia, Sabah, SW Sabah, nr. Long Pa teral sclerites. Other species from Sabah having this Sia (west), 1050 m, 25.xi-8.xii.1987, Achterberg, combination of characters include C. cygnus, sp. n. 1? (RMNH); 10 km S Long Pa Sia, Sg. Malabit, and C. danumensis, sp. n. Chimarra lambi is most 04°21’N, 115°41’E, 1180 m, 2.xii.1987, Huisman similar to C. danumensis, especially in having inferior & Achterberg, 2? (RMNH); 18 km on road Ken- appendages with the apex bicusped, but differs diag- ingau-Kimanis, gravel quarry, 05°26’N, 116°05’E, nostically in that the cusps are located more apically, 1050 m, 20.i.1987, J. Huisman, 2?, 4/ (UMSP); and also in that the lateral lobes of tergum X are 12 km NNE Ranau, Poring Hot Spring, Sg. Tan- more dorsoventrally flattened and the phallotremal anansad, 06°03’N, 116°42’E, 560 m, 28.viii.1986, sclerite complex is shorter. J. Huisman, 4? (UMSP); 2 km SW Long Pa Sia, Adult. Color (in alcohol) light brown. Length of Long Rurun (Sg. Ritan), 04°23’N, 115°42’E, forewing: male 5.0–5.6 mm, female 5.2–5.8 mm. 1040 m, 5.xii.1987, Huisman & Achterberg, 2? Forewing venation: stem of Rs curved, slightly scle- (RMNH); Poring Hot Spring, 12 km NNE Ranau, rotized and thickened at inflection; fork at base of Sg. Kipogoh, 06°03’N, 116°42’E, 550 m, 9.xii.1986, discal cell somewhat thickened, nearly symmetrical, J. Huisman, 2? (UMSP); 12 km NNE Ranau, Por- length of discal cell about 3 times width; m crossvein ing Hot Spring, staff quarters, 06°03’N, 116°42’E, proximal to crossveins s and r-m, crossvein s not hya- 550 m, Huisman & de Jong, 1? (UMSP). line; 2A vein not intersecting 3A (apparently looped 142 Tijdschrift voor Entomologie, volume 152, 2009

a c d

e f

Fig. 23. Chimarra liwaguensis, male genitalia. – a, lateral; b, segment IX and tergum X, dorsal; c, inferior appendage, ventral; d, inferior appendage, dorsal; e, phallic apparatus, lateral; f, tergum VIII, dorsal.

to 1A). Postocular parietal sclerite short. Maxillary lobe elongate, digitate, posterodorsally directed. palps moderately elongate, segment 3 slightly greater Phallobase tubular, with pronounced basodorsal than 2, 3 subequal to 5. Protarsal claws of male expansion, moderately elongate. Endotheca length enlarged, asymmetrical in shape and length. not discernable (not expanded); without endothecal Male genitalia. Abdominal segment VIII moderate spines. Phallotremal sclerite complex composed of in length; tergum distinctly longer than sternum, rod and ring structure, rod elongate, dorsolaterally, modified, with V-shaped mesal excavation, excava- with pair of elongate curved sclerites. tion apicolaterally with laciniate fringe; sternum Etymology. Named for Anthony Lamb, friend of VIII without posteroventral projection. Abdominal J. Huisman. segment IX relatively wide dorsolaterally, tergum obsolete dorsomesally, without distinct anterolateral Chimarra liwaguensis sp. n. apodemes; anteroventral margin in lateral view not, Fig. 23 or scarcely produced; posterior margin distinctly produced; ventral process projecting ventrally, short, broad basally, subtriangular. Preanal append- Type material. Holotype: ?, Malaysia, Sabah, Kina- age setose, short, rounded, broadest basally. Inferior balu National Park, Headquarters, Sg. Liwagu cross- appendage about as long as tergum X; in lateral view ing Silau-Silau trail, 06°00’N, 116°33’E, 1470 m, straight, nearly uniformly wide, apex with mesally 15.viii.1986, J. Huisman (UMSP000107130) directed process; in ventral view nearly straight, (RMNH). apex subtruncate; in caudal view with evident Paratypes. Malaysia, Sabah, same data as holotype, sclerotized, bicusped projection. Tergum X with 3/ (UMSP); Kinabalu National Park, Sg. Liwagu, sclerotized lateral lobes and separate mesal lobes, 1100 m, 12.viii.2005, W. Mey, 3? (MNHU); mesal membrane lacking; lateral lobe moderately Kinabalu National Park, Headquarters, Sg. Liwagu long, with apex broadly rounded, in lateral view sim- @ Silau-Silau trail, 06°00’N, 116°33’E, 1470 m, ple, dorsoventrally flattened, rounded apically, lobe 22.i.1987, Huisman & de Jong, 2? (UMSP); same bearing multiple short setae or seta-like sensilla; mesal locality, 10.ii.1987, J. Huisman, 1?, 16/ (UMSP); Blahnik et al.: Chimarra of northern Borneo 143 same locality, 15.xi.1986, J. Huisman, 1?, 7/ sensilla; mesal lobe elongate, digitate, reclinate, poste- (RMNH); Kinabalu National Park, Headquarters, riorly directed. Phallobase tubular, with pronounced on bridge to “Sg. Liwagu section I”, 06°00’N, basodorsal expansion, short. Endotheca relatively 116°33’E, 1480 m, 9.ii.1987, J. Huisman, 1?, 1/ short; without endothecal spines. Phallotremal scler- (RMNH); Kundassang, Sg. Mesilau, 1st bridge after ite complex composed of rod and ring structure, rod golf course, 06°01’N, 116°36’E, 1650 m, 3.x.1986, short, preapically with short lateral sclerites. J. Huisman, 7?, 2/ (UMSP); Kinabalu National Etymology. This species is named Chimarra liwaguen- Park, Headquarters area, 1560 m, 13.ix.1983, Hevel sis for the stream, Sungai Liwagu, where the holotype & Steiner, 1? (NMNH). specimen was collected.

Chimarra liwaguensis is another species of the Chi- Chimarra noloyan sp. n. marra tsudai group in which tergum VIII is deeply Fig. 24 invaginated mesally and variously armed with spine- like projections. It is easily diagnosed by the shape of the inferior appendage, which, in ventral view, is Type material. Holotype: ?, Malaysia, Sabah, Crocker distinctly mesally curved, broad basally, narrow and Range, 5 km N Tenom, base of Sg. ulu Noloyan, rounded apically, and has an acute projection just 05°10’N, 115°56’E, 1010 m, 10–11.x.1986, J. Huis- past midlength from the ventral surface. The pos- man (UMSP000107155) (RMNH). teromesal excavation of tergum is also a useful diag- Paratypes. Malaysia, Sabah, Kinabalu National Park, nostic character in that it is deep and U-shaped and Sungai Liwagu, 1100 m, 12.viii.2005, W. Mey, 4?, margined by spines of a uniform length. Addition- 3/ (UMHU). ally, the phallic apparatus is simple in structure, with a very short phallobase and without internal spines. Chimarra noloyan also belongs to the distinctive sub- Adult. Color (in alcohol) brown. Length of fore- group of the Chimarra tsudai group in which tergum wing: male 6.3–7.0 mm, female 6.7–7.2 mm. Fore- VIII is invaginated mesally and variously armed with wing venation: stem of Rs curved, slightly sclerotized spine-like projections. Generalized features it pos- and thickened at inflection; fork at base of discal cell sesses, in common with some other members of this somewhat thickened, nearly symmetrical, length of group, are a phallotremal sclerite that is not greatly discal cell about 3 times width; m crossvein proxi- enlarged, and an endotheca that is relatively short mal to crossveins s and r-m, crossvein s not hyaline; and lacks distinctive spines. In C. noloyan the length 2A vein not intersecting 3A (apparently looped to of tergum VIII is short, thus the mesal invagination 1A). Postocular parietal sclerite elongate. Maxillary is not very deep and the posterior fringe of spine- palps moderately elongate, segment 3 subequal to 2, like projections are irregular in length and are not 5 very narrow, slightly shorter than 3. Protarsal claws formed into elongate processes. Chimarra noloyan is of male enlarged, asymmetrical in shape and length. most likely to be confused with species with similarly Male genitalia. Abdominal segment VIII moderate shaped inferior appendages, those with a mesal proc- in length; tergum distinctly longer than sternum, ess that is just preapical and not very strongly pro- modified, with broad, U-shaped mesal excavation, jecting, especially C. silausilau, sp. n. and C. vantoli, excavation with short laciniate fringe; sternum sp. n. In C. silausilau the apices of the mesal projec- VIII without posteroventral projection. Abdominal tions of the inferior appendage are more posteriorly segment IX relatively wide dorsolaterally, tergum directed and the lateral lobe of tergum X more nearly obsolete dorsomesally, without distinct anterola- subquadrate in lateral view. In C. vantoli the mesal teral apodemes; anteroventral margin in lateral view projections of the inferior appendage are almost slightly produced, rounded; posterior margin dis- apical and the laciniate fringe of tergum VIII more tinctly produced; ventral process absent or nearly elongate. An additional difference is that the mesal so. Preanal appendage setose, very small, rounded. lobes of tergum X in C. noloyan are shorter, flatter, Inferior appendage longer than tergum X; in lateral and broader than in related species. view slightly curved, nearly uniformly wide, apex Adult. Color (in alcohol) yellowish-brown. Length broadly rounded; in ventral view strongly mesally of forewing: male 5.0 mm. Forewing venation: stem curved, broad basally, ventromesally with broad, of Rs curved, slightly sclerotized and thickened at apically acute projection. Tergum X with sclerotized inflection; fork at base of discal cell somewhat thick- lateral lobes and separate mesal lobes, mesal mem- ened, nearly symmetrical, length of discal cell about brane lacking; lateral lobe moderately long, with 2.5 times width; m crossvein proximal to crossveins apex broadly rounded, in lateral view narrow, par- s and r-m, crossvein s not hyaline; 2A vein not inter- allel sided, rounded apically, lobe bearing multiple secting 3A (apparently looped to 1A). Postocular 144 Tijdschrift voor Entomologie, volume 152, 2009

a b c d

e f

Fig. 24. Chimarra noloyan, male genitalia. – a, lateral; b, segment IX and tergum X, dorsal; c, inferior appendage, ventral; d, inferior appendage, dorsal; e, phallic apparatus, lateral; f, tergum VIII, dorsal.

parietal sclerite moderately elongate. Maxillary palps a noun in apposition, for the type locality, Sungai ulu relatively short, segment 3 slightly longer than 2, Noloyan, where the holotype specimen was collected. 2 subequal to 5. Protarsal claws of male enlarged, asymmetrical in shape and length. Chimarra phillipsae sp. n. Male genitalia. Abdominal segment VIII short; ter- Figs 25, 42 gum about as long as sternum, modified, with narrow mesal excavation, excavation boardered laterally with laciniate fringe of variable length; sternum Type material. Holotype: ?, Malaysia, Sabah, Kina- VIII without posteroventral projection. Abdominal balu National Park, Headquarters, Sg. Liwagu cross- segment IX relatively wide dorsolaterally, tergum ing Silau-Silau trail, 06°00’N, 116°33’E, 1470 m, obsolete dorsomesally, without distinct anterolateral 15.xi.1986, J. Huisman (UMSP000107136) apodemes; anteroventral margin in lateral view not, (RMNH). or scarcely produced; posterior margin distinctly Paratypes. Malaysia, Sabah, Kinabalu National produced; ventral process projecting ventrally, Park, Headquarters area, 1560 m, 7.ix.1983, Hevel short, broad basally, subtriangular. Preanal append- & Steiner, 1? (NMNH); Kinabalu National Park, age setose, very small, rounded. Inferior appendage Sungai Liwagu, 1500 m, 10.viii.2005, W. Mey, 1? about as long as tergum X; in lateral view straight, (UMHU); Crocker Range, 40 km S of Kota Kina- nearly uniformly wide, apex rounded; in ventral view balu, Sinsuron Rd., 1500 m, 19.xii.1989, J. Huis- nearly straight, preapically with mesally directed, man, 1?, 1/ (UMSP); same locality, 20.xii.1989, tooth-like projection. Tergum X with sclerotized lat- J. Huisman, 1? (UMSP); Poring Hot Spring, eral lobes and separate mesal lobes, mesal membrane 12 km NNE Ranau, Sg. Kepungit, E park bound- lacking; lateral lobe moderately long, in lateral view ary, 06°03’N, 116°42’E, 480 m, 26.xi.1986, J. Huis- narrow, attenuate apically, lobe bearing multiple sen- man, 4? (UMSP); Poring Hot Spring, 12 km NNE silla; mesal lobe short, thumb-shaped, posterodor- Ranau, Sg. Kepungit waterfall, 06°03’N, 116°42’E, sally directed. Phallobase tubular, with pronounced 550, 4.xii.1986, J. Huisman, 2? (RMNH); basodorsal expansion, short. Endotheca relatively Kg. Kiau, Sg. Kadamaian, 06°02’N, 116°31’E, 720 m, short, with granularly textured region; without 26.viii.1986, J. Huisman, 1? (RMNH); 9.5 km endothecal spines. Phallotremal sclerite complex NNW Kundassang, Marei-Parei, Sg. Tawubang, composed of rod and ring structure, rod short, api- 06°04’N, 116°30’E, 1030 m, 11.iii.1987, J. Huis- cally with pair of short, curved, lateral sclerites. man, 14?, 1/ (RMNH). Etymology. This species is named Chimarra noloyan, as Blahnik et al.: Chimarra of northern Borneo 145 a c d

e f

Fig. 25. Chimarra phillipsae, male genitalia. – a, lateral; b, segment IX and tergum X, dorsal; c, inferior appendage, ventral; d, inferior appendage, dorsal; e, phallic apparatus, lateral; inset, phallotremal sclerite, dorsal; f, tergum VIII, dorsal.

Chimarra phillipsae is related to those species of the tergum about as long as sternum, modified, with Chimarra tsudai group with a modified, mesally small mesal excavation, excavation with short, paired, invaginated, spinose tergum VIII, although the mesal downturned laciniate mesal processes; sternum invagination and apical spines in this species are not VIII without posteroventral projection. Abdominal as pronounced as some other members of this group. segment IX relatively wide dorsolaterally, without Chimarra phillipsae is most readily diagnosed by the distinct anterolateral apodemes; anteroventral mar- distinctive shape of its inferior appendage, which has gin in lateral view slightly produced; posterior margin an apicoventral lobe, making the apex of the append- slightly produced; ventral process projecting ventrally, age appear broadly truncate in lateral view. Another very short, broad basally, broadly rounded. Preanal diagnostic feature is the distinctive shape of the mesal appendage setose, short, broadly rounded, broadest lobe of tergum X, which has a rounded basal enlarge- basally. Inferior appendage slightly longer than ter- ment and an extending, digitate apical projection. gum X; in lateral view straight, widest preapically, Adult. Color (in alcohol) brown. Length of forewing: apex slightly angulate, slightly upturned; in ventral male 5.4–6.2 mm, female 6.8–7.5 mm. Forewing view mesally curved, mesal surface with tooth-like venation: stem of Rs curved; fork at base of discal projection at midlength. Tergum X with sclerotized cell not thickened, nearly symmetrical, length of dis- lateral lobes and separate mesal lobes, mesal mem- cal cell about 3 times width; m crossvein somewhat brane lacking; lateral lobe moderately long, in lateral proximal to crossveins s and r-m, crossvein s hyaline, view simple, rounded apically, slightly downcurved, in part; 2A vein not intersecting 3A (apparently lobe bearing multiple sensilla; mesal lobe with looped to 1A). Postocular parietal sclerite elongate. rounded sensillate basal projection and elongate, nar- Maxillary palps moderately elongate, segment 3 sub- row ventral projection. Phallobase tubular, with pro- equal to 2, 5 slightly greater than 3, narrow. Protarsal nounced basodorsal expansion, short, ventral apex claws of male enlarged, asymmetrical in shape and lightly sclerotized, projecting. Endotheca about as length. long as phallobase; without endothecal spines. Phal- Male genitalia. Abdominal segment VIII short; lotremal sclerite complex composed of rod and ring 146 Tijdschrift voor Entomologie, volume 152, 2009

a b c d

e f

Fig. 26. Chimarra physanoton, male genitalia. – a, lateral; b, segment IX and tergum X, dorsal; c, inferior appendage, ventral; d, inferior appendage, dorsal; e, phallic apparatus, lateral; f, tergum VIII, dorsal.

structure, rod moderately elongate, apically with pair modified, invaginated mesally and variously armed of short, curved, lateral sclerites. with spine-like projections. Generalized features it Etymology. This species is also named for Anthea possesses are a phallotremal sclerite that is not greatly Phillips, friend of J. Huisman. enlarged, and an endotheca that is relatively short and lacks distinctive spines. It is most similar to a Chimarra physanoton sp. n. group of species from Sabah, including C. noloyan, sp. n., C. preapicalis, sp. n., C. silausilau, sp. n., and Fig. 26 C. vantoli, sp. n., all of which have similarly shaped inferior appendages, which are short, linear, with Type material. Holotype: ?, Malaysia, Sabah, Kun- an acute mesal projection located preapically, and a dassang kampong, Sg. Liwagu, on bridge, 06°00’N, tergum VIII that is relatively narrow. It is probably 116°34’E, 1185 m, 23.xi.1986, J. Huisman most similar to C. silausilau and C. vantoli, differing (UMSP000107246) (RMNH). distinctively from either in the shape of the lateral Paratypes. Malaysia, Sabah, same locality as holotype, lobes of tergum X, which are inflated basally, with a 23.viii.1986, J. Huisman, 1? (RMNH); SW Sabah, lateral crease, and with apices that are downturned nr. Long Pa Sia (west), 1050 m, 25.xi-8.xii.1987, and acutely narrowed. Like C silausilau, it has the Achterberg, 2?, 10/ (UMSP); Long Pa Sia airstrip, apicomesal projections of the inferior appendage dis- 04°24’N, 115°43’E, 1000 m, 2–3.xii.1989, J. Huis- tinctly posteriorly directed. man, 1? (UMSP); Crocker Range, 5km N Tenom, Adult. Color (in alcohol) brown. Length of fore- base of Sg. ulu Noloyan, 05°10’N, 115°56’E, wing: male 5.7–6.0 mm, female 5.7–6.2 mm. Fore- 1010 m, 10–11.x.1986, J. Huisman, 1? (RMNH); wing venation: stem of Rs curved, slightly sclerotized road, Long Pa Sia - Long Semado, Sg. Malabit, and thickened at inflection; fork at base of discal cell 06°21’N, 115°41’E, 1175 m, 3–4.xii.1987, J. Huis- somewhat thickened, nearly symmetrical; m crossvein man, 1?, 2/ (UMSP). proximal to crossveins s and r-m, crossvein s not hyaline; 2A vein not intersecting 3A (apparently looped Chimarra physanoton is another species belong- to 1A). Postocular parietal sclerite short. Maxillary ing to the distinctive subgroup of the Chima- palps relatively short, segment 3 slightly longer than rra tsudai group in which tergum VIII is highly 2, 3 subequal to 5. Protarsal claws of male enlarged, Blahnik et al.: Chimarra of northern Borneo 147 a b c d

e f

Fig. 27. Chimarra preapicalis, male genitalia. – a, lateral; b, segment IX and tergum X, dorsal; c, inferior appendage, ventral; d, inferior appendage, dorsal; e, phallic apparatus, lateral; inset, phallotremal sclerite, dorsal; f, tergum VIII, dorsal.

asymmetrical in shape, scarcely different in length. apically with pair of short, curved, lateral sclerites. Male genitalia. Abdominal segment VIII short; ter- Etymology. This species is named Chimarra physan- gum about as long as sternum, modified, with small oton, used as a noun in apposition, from the Greek mesal excavation, excavation bordered laterally with words physa, a bag or bellows, and noton, a back, laciniate fringe, mesal and lateral fringe more elon- and referring to the basally inflated tergum X in this gate; sternum VIII without posteroventral projection. species. Abdominal segment IX short dorsolaterally, tergum obsolete dorsomesally; anteroventral margin in lat- Chimarra preapicalis sp. n. eral view slightly produced; posterior margin sinu- Fig. 27 ously produced; ventral process projecting ventrally, short, broad basally, subtriangular. Preanal appendage setose, very small, rounded. Inferior appendage Type material. Holotype: ?, Malaysia, Sabah, about as long as tergum X; in lateral view straight, Crocker Range, 40 km S of Kota Kinabalu, Sin- nearly uniformly wide, apex rounded; in ventral suron Rd., 1500 m, 20.xii.1989, J. Huisman view nearly straight, apically with small, posterome- (UMSP000107216) (RMNH). sally directed, tooth-like projection. Tergum X with Paratypes. Malaysia, Sabah, Kinabalu National Park, sclerotized lateral lobes and separate mesal lobes, Sungai Liwagu, 1500 m, 10.viii.2005, W. Mey, 3? mesal membrane lacking; lateral lobe long, in lateral (UMHU); Kundassang kampong, Sg. Liwagu, on view inflated at midlength, apex attenuate, subacute, bridge, 06°00’N, 116°34’E, 1185 m, 23.xi.1986, lobe bearing multiple sensilla; mesal lobe elongate, J. Huisman, 1? (RMNH); Kinabalu National Park, digitate, slightly posterodorsally curved. Phallobase Headquarters, Sg. Liwagu, 06°01’N, 116°32’E, tubular, with pronounced basodorsal expansion, 1500 m, 11.ii.1987, J. Huisman, 1? (UMSP); 1 km relatively short, ventral apex distinctly sclerotized, S Kundasang, 1530 m, 22.viii.1983, Hevel & Steiner, projecting, subacute. Endotheca relatively short, 1? (NMNH). with granularly textured region, without endothecal spines. Phallotremal sclerite complex composed Chimarra preapicalis also belongs the distinctive of rod and ring structure, rod moderately elongate, subgroup of the Chimarra tsudai group in which 148 Tijdschrift voor Entomologie, volume 152, 2009

tergum VIII is highly modified, invaginated mesally bearing multiple sensilla; mesal lobe elongate, digi- and variously armed with spine-like projections. tate, posterolaterally directed, apex acute. Phallobase Generalized features it possesses are a phallotremal tubular, with pronounced basodorsal expansion, sclerite that is not greatly enlarged, and an endotheca short. Endotheca relatively short, with granularly that is relatively short and lacks distinctive spines, textured region; without endothecal spines. Phal- although the endotheca is “textured” with minute lotremal sclerite complex composed of rod and ring spines. It is most similar to a group of species from structure, rod moderately elongate, preapically with Sabah, including C. noloyan, sp. n., C. physanoton, sp. n., short lateral sclerites. C. silausilau, sp. n., and C. vantoli, sp. n., all of which Etymology. This species is named Chimarra preapi- have similarly shaped inferior appendages, which are calis because the mesal projections of the inferior short, linear, with the acute projections located api- appendages are located more preapically (farther comesally and a tergum VIII that is narrow and tends from the apex) than in closely related species. to have the laciniate fringe formed into 2 pairs of projections, margined with spines of variable length. Chimarra prokrustes Malicky From other species in this group it is most diagnosti- Fig. 28 cally distinguished by details in shape of the inferior appendages, which have the acute apicomesal pro- Chimarra prokrustes Malicky, 2008: 840, holotype ?, In- jections emerging more preapically (farther from the donesia (Eastern Kalimantan), Zoologische Museum apex) than most members of this group and are also Lausanne. more prominently developed. Details in the shape of the lateral lobes of tergum X and development of Material examined. Malaysia, Sabah, 18 km on road the armature of tergum VIII are also useful adjunct Keningau-Kimanis, gravel quarry, 05°26’N, 116°05’E, characters. 1050 m, 20.i.1987, J. Huisman, 1? (RMNH); 12 km Adult. Color (in alcohol) light brown. Length of NNE Ranau, Poring Hot Spring, Sg. Tananansad, 06°03’N, 116°42’E, 560 m, 28.viii.1986, J. Huisman, 1? (UMSP); forewing: male 4.8–5.6 mm. Forewing venation: 12 km NNE Ranau, Poring Hot Spring, Sg. Kepungit, stem of Rs curved, slightly sclerotized and thickened 06°03’N, 116°42’E, 550 m, 24.i.1987, J. Huisman, 1? at inflection; fork at base of discal cell somewhat (UMSP). thickened, nearly symmetrical, length of discal cell about 3 times width; m crossvein proximal to cross- Chimarra prokrustes resembles those species from veins s and r-m, crossvein s not hyaline; 2A vein not Borneo in the Chimarra tsudai group that have an intersecting 3A (apparently looped to 1A). Postocu- unmodified, but somewhat widened tergum VIII, lar parietal sclerite short. Maxillary palps moderately an acute projection or tooth on the dorsal margin elongate, segment 3 slightly greater than 2, 3 sub- of the inferior appendages, evident in lateral view, equal to 5. Protarsal claws of male enlarged, asym- and mesal lobes of tergum X that are sclerotized, metrical in shape and length. upturned and have an acute apex. Species in this Male genitalia. Abdominal segment VIII short; ter- group include C. denticula, sp. n., C. karlijnae, sp. n., gum longer than sternum, modified, with mesal C. scolops, sp. n., C. sinitorum, sp. n., and C. steno- excavation, excavation apicolaterally with bifur- dactylus, sp. n. All of these species also have a phallic cating, laciniate processes; sternum VIII without apparatus with numerous small spines and a narrow posteroventral projection. Abdominal segment IX sclerous dorsal extension, extending over the basal relatively wide dorsolaterally, tergum obsolete dor- part of the endotheca. As previously discussed, these somesally, without distinct anterolateral apodemes; species are all easily distinguished from one another anteroventral margin in lateral view slightly pro- by the characteristic and species-specific shapes of duced, rounded; posterior margin distinctly pro- the lateral and mesal lobes of tergum X, as well as duced; ventral process projecting ventrally, broad by differences in the shape of the inferior append- basally, subtriangular. Preanal appendage setose, ages. Among these species, Chimarra prokrustes is small, rounded, slightly narrowed basally. Infe- easily diagnosed by the very elongate and typically rior appendage slightly longer than tergum X; in undulate lateral lobes of tergum X, and by the wid- lateral view straight, widest basally, gradually nar- ened base of the dorsomesal process of the inferior rowing distally, apex rounded; in ventral view appendage. As compared to the illustration of the nearly straight, preapically with prominent, mesal, holotype, the form figured here seems to differ in tooth-like projection. Tergum X with sclerotized that the lateral lobes of tergum X are more irregu- lateral lobes and separate mesal lobes, mesal mem- lar in shape also have distinct, sensillate basolateral brane lacking; lateral lobe moderately long, in lat- protrusions, as well as inferior appendages, as viewed eral view narrow, apex slightly downcurved, lobe ventrally, with their apices less narrowed. Compared Blahnik et al.: Chimarra of northern Borneo 149 a b c d

Fig. 28. Chimarra prokrustes, male genitalia. – a, lateral; b, segment IX and tergum X, dorsal; c, inferior appendage, ventral; d, inferior appendage, dorsal; e, phallic apparatus, lateral; f, phallic apparatus, dorsal. to the differences that separate this species from its dorsoventrally; posterior margin distinctly pro- most closely related congeners, these discrepancies duced; ventral process absent or nearly so. Preanal are minor and probably not significant. appendage setose, short, rounded, broadest basally. Adult. Color (in alcohol) yellowish-brown. Length Inferior appendage longer than tergum X; in lateral of forewing: 4.4–5.0 mm. Forewing venation: stem view straight, widest at midlength, narrow distally, of Rs curved, slightly sclerotized and thickened at apex rounded, dorsal margin with very basally wide, inflection; fork at base of discal cell not thickened, abruptly apically acute projection at midlength; in nearly symmetrical, length of discal cell about ventral view somewhat mesally curved, mesal surface 3 times width; m crossvein proximal to crossveins without processes. Tergum X with sclerotized lateral s and r-m, crossvein s hyaline, in part; 2A vein not lobes and separate mesal lobes, mesal membrane intersecting 3A (apparently looped to 1A). Postocu- lacking; lateral lobe long, in lateral view narrow, lar parietal sclerite short. Maxillary palps relatively undulate, rounded apically, lobe bearing multiple short, segment 3 slightly longer than 2, 3 subequal sensilla; mesal lobe short, digitate, distinctly scle- to 5. Protarsal claws of male enlarged, asymmetrical rotized, dorsally directed, in lateral view, narrow, in shape and length. recurved, apex with single, acuminate spine-like Male genitalia. Abdominal segment VIII short; ter- projection. Phallobase tubular, with pronounced gum longer than sternum, unmodified; sternum basodorsal expansion, moderately elongate, relatively VIII without posteroventral projection. Abdominal narrow, dorsal margin extended apically, elongate, segment IX short dorsolaterally, tergum obsolete narrow, lightly sclerotized. Endotheca length not dorsomesally, without distinct anterolateral apo- discernable (not expanded); with numerous short demes; anteroventral margin in lateral view pro- endothecal spines. Phallotremal sclerite complex not duced, ventral margin rounded mesally as viewed evident in material examined. 150 Tijdschrift voor Entomologie, volume 152, 2009

a b c d

e g

Fig. 29. Chimarra scolops, male genitalia. – a, lateral; b, segment IX and tergum X, dorsal; c, inferior appendage, ventral; d, inferior appendage, dorsal; e, phallic apparatus, lateral; f, phallic apparatus, dorsal; inset phallotremal sclerite, ventral; g, expanded endotheca, lateral.

Chimarra scolops sp. n. the inferior appendages, evident in lateral view, and Fig. 29 mesal lobes of tergum X that are sclerotized, upturned and have an acute apex. Species in this group include Type material. Holotype: ?, Malaysia, Sabah, Kina- C. denticula, sp. n., C. karlijnae, sp. n., C. prokrustes balu National Park, Headquarters, Sg. Liwagu cross- Malicky, C. sinitorum, sp. n., and C. stenodactylus, sp. ing Silau-Silau trail, 06°00’N, 116°33’E, 1470 m, n. All of these species also have a phallic apparatus 15.viii.1986, J. Huisman (UMSP000208645) with numerous small spines and a narrow sclerous (RMNH). dorsal extension, extending over the basal part of the Paratypes. Malaysia, Sabah, same data as holotype, endotheca. These species are all easily distinguished 2? (UMSP); Kinabalu National Park, headquarters, from one another by the characteristic and species- Sungai Silau-Silau, 11.viii.2005, W. Mey, 3?, 1/ specific shapes of the lateral and mesal lobes of ter- (UMHU); Kinabalu National Park, Sungai Liwagu, gum X, as well as by differences in the shape of the 1100 m, 12.viii.2005, W. Mey, 4? (UMHU); same inferior appendages. Among these species, Chimarra locality, 15.xi.1986, J. Huisman, 1? (RMNH); scolops is probably most similar to C. denticulata, but Crocker Range, 40 km S of Kota Kinabalu, Sin- differs in both the shape of the lateral lobes of ter- suron Rd., 1500 m, 18.xii.1989, J. Huisman, 2? gum X and in lacking a preapical “denticle” or tooth- (UMSP), same locality, 20.xii.1989, J. Huisman, like projection on the inferior appendage. 1? (RMNH). Adult. Color (in alcohol) yellowish-brown. Length of forewing: male 5.8–6.5 mm. Forewing venation: Chimarra scolops resembles those species in the stem of Rs curved, distinctly sclerotized and thick- Chimarra tsudai group from Sabah that have an ened at inflection; fork at base of discal cell distinctly unmodified, but somewhat widened tergum VIII, thickened, nearly symmetrical, length of discal cell an acute projection or tooth on the dorsal margin of about 2.5 times width; m crossvein proximal to Blahnik et al.: Chimarra of northern Borneo 151 a b c d

e f

Fig. 30. Chimarra silausilau, male genitalia. – a, lateral; b, segment IX and tergum X, dorsal; c, inferior appendage, ventral; d, inferior appendage, dorsal; e, phallic apparatus, lateral; inset, phallotremal sclerite, dorsal; f, tergum VIII, dorsal. crossveins s and r-m, crossvein s hyaline; 2A vein not Phallobase tubular, with pronounced basodor- intersecting 3A (apparently looped to 1A). Postocu- sal expansion, moderately elongate, ventral apex lar parietal sclerite moderately elongate, head very lightly sclerotized, slightly projecting, dorsal mar- large. Maxillary palps elongate, segment 3 subequal gin extended apically, elongate, narrow, lightly scle- to 2 and 5. Protarsal claws of male enlarged, asym- rotized. Endotheca elongate, with several tracts of metrical in shape and length. short spines, 1 basally, 1 dorsally in basal half, and Male genitalia. Abdominal segment VIII moderate 1 apically or preapically, composed of ventral group in length; tergum distinctly longer than sternum, of spines and paired lateral groups of several spines. unmodified; sternum VIII without posteroventral Phallotremal sclerite complex composed of rod and projection. Abdominal segment IX relatively wide ring structure, rod relatively short. dorsolaterally, tergum obsolete dorsomesally, with- Etymology. This species is named Chimarra scolops, out distinct anterolateral apodemes; anteroventral a noun in apposition, from the Greek word skolops, margin in lateral view not, or scarcely produced; a thorn, and referring to the thorn-like apices of the posterior margin slightly produced; ventral process mesal processes of tergum X in this species. absent or nearly so. Preanal appendage setose, short, rounded, broadest basally. Inferior appendage much Chimarra silausilau sp. n. longer than tergum X; in lateral view straight, nearly Fig. 30 uniformly wide, apex broadly rounded, dorsal margin with very basally wide, apically acute projection at midlength; in ventral view very slightly mesally Type material. Holotype: ?, Malaysia, Sabah, curved, nearly straight, mesal surface without proc- Kinabalu National Park, Sg. Silau-Silau, 06°00’N, esses. Tergum X with sclerotized lateral lobes and 116°32’12”E, 1560 m, bridge near Bukit Burung separate mesal lobes, mesal membrane lacking; lat- trail, 20–21.xi.1986, J. Huisman (UMSP000107237) eral lobe short, in lateral view irregular, rounded (RMNH). apically, distinctly downcurved, lobe bearing mul- Paratypes. Malaysia, Sabah, same data as holotype, tiple sensilla; mesal lobe short, digitate, distinctly 2?, 3/ (UMSP); Kinabalu National Park, Sg. Tiba- sclerotized, dorsally directed, with basal sensillate bar @ Liwagu trail, 06°02’N, 116°33’E, 1750 m, lobe and strongly dorsally hooked apical projection. 2.x.1986, J. Huisman, 1?, 3/ (RMNH). 152 Tijdschrift voor Entomologie, volume 152, 2009

Chimarra silausilau belongs to the distinctive sub- flattened. Phallobase tubular, with pronounced baso- group of the Chimarra tsudai group in which tergum dorsal expansion, short, ventral apex weakly sclero- VIII is highly modified, invaginated mesally and tized, slightly projecting. Endotheca relatively short, variously armed with spine-like projections. Gener- granularly textured region not discernable; without alized features it possesses are a phallotremal sclerite endothecal spines. Phallotremal sclerite complex that is not greatly enlarged, and an endotheca that is composed of rod and ring structure, rod short, api- relatively short and lacks distinctive spines, although cally with pair of short, curved, lateral sclerites. the endotheca is “textured” with minute spines. It Etymology. This species is named Chimarra silausilau, is most similar to a group of species from Sabah, as a noun in apposition, for the stream, Sungai Silau- including C. noloyan, sp. n., C. physanoton, sp. n., Silau, where the holotype specimen was collected. C. preapicalis, sp. n., and C. vantoli, sp. n., all of which have similarly shaped inferior appendages, Chimarra sinitorum sp. n. which are short, linear, and with the acute projec- Fig. 31 tions located apicomesally. All of these species also have a tergum VIII that is particularly narrow. Chimarra silausilau is probably most similar to Type material. Holotype: ?, Malaysia, Sabah, Poring C. physanoton and C. vantoli, differing from either Hot Spring, 12 km NNE Ranau, Sg. Montokungon, in having the apex of the lateral lobes of tergum X 06°02’N, 116°42’E, 525 m, 30.i.1987, J. Huisman more nearly subquadrate (less angular apically). Like (UMSP000107298) (RMNH). C. physanoton, it has the apicomesal projections of Paratypes. Malaysia, Sabah, same data as holotype, the inferior appendage posteriorly directed. 3?, 3/ (UMSP); 12 km NNE Ranau, Poring Adult. Color (in alcohol) light brown. Length of Hot Springs, Sg. Langanan, 980 m, 29.viii.1986, forewing: male 6.0–6.5 mm, female 6.5–6.8 mm. J. Huisman, 2?, 5/ (UMSP); Poring Hot Spring, Forewing venation: stem of Rs curved, slightly scle- 12 km NNE Ranau, confluence Sg. Kepungit & rotized and thickened at inflection; fork at base of Sg. Langanan, 06°03’N, 116°43’E, 450 m, 29.i.1987, discal cell somewhat thickened, nearly symmetrical, J. Huisman, 1? (RMNH); 12 km NNE Ranau, length of discal cell about 3 times width; m cross- Poring Hot Spring, Sg. Kepungit, 06°03’N, vein proximal to crossveins s and r-m, crossvein s 116°42’E, 480 m, 27.i.1987, J. Huisman, 1? not hyaline; 2A vein not intersecting 3A (apparently (RMNH); same locality, 550 m, 24.i.1987, J. Huis- looped to 1A). Postocular parietal sclerite moderately man, 20? (UMSP) Poring Hot Spring, 12 km NNE elongate. Maxillary palps moderately elongate, 3 dis- Ranau, Sg. Kepungit, E park boundary, 06°03’N, tinctly longer than 2, 5 slightly shorter than 3. Pro- 116°42’E, 480 m, 26.xi.1986, J. Huisman, 4?, 5/ tarsal claws of male enlarged, asymmetrical in shape (RMNH); Poring Hot Spring, 12 km NNE Ranau, and length. Sg. Kepungit waterfall, 06°03’N, 116°42’E, 550 m, Male genitalia. Abdominal segment VIII short; ter- 4.xii.1986, J. Huisman, 2? (UMSP); same locality, gum about as long as sternum, modified, with mesal 625 m, 29.viii.1986, J. Huisman, 1?, 3/; 12 km excavation, excavation apicolaterally with bifurcat- NNE Ranau, Poring Hot Spring, Sg. Tananansad, ing, laciniate processes, lateral process more elongate; 06°03’N, 116°42’E, 560 m, 28.viii.1986, J. Huis- sternum VIII without posteroventral projection. man, 2? (RMNH). Abdominal segment IX relatively wide dorsolaterally, tergum obsolete dorsomesally, without dis- Chimarra sinitorum resembles those species from tinct anterolateral apodemes; anteroventral margin Borneo in the Chimarra tsudai group that have an in lateral view slightly produced; posterior margin unmodified, but somewhat widened tergum VIII, distinctly produced; ventral process projecting ven- an acute projection or tooth on the dorsal margin trally, very short, broad basally, broadly rounded. of the inferior appendages, evident in lateral view, Preanal appendages setose, very small, rounded. and mesal lobes of tergum X that are sclerotized, Inferior appendage about as long as tergum X; in upturned and have an acute apex. Species in this lateral view straight, nearly uniformly wide, apex group include C. denticula, sp. n., C. karlijnae, rounded; in ventral view nearly straight, apically sp. n., C. prokrustes Malicky, C. scolops, sp. n., and with posteromesally directed, tooth-like projection. C. stenodactylus, sp. n. All of these species also have Tergum X with sclerotized lateral lobes and separate a phallic apparatus with numerous small spines and mesal lobes, mesal membrane lacking; lateral lobe a narrow sclerous dorsal extension, extending over moderately long, in lateral view simple, subquadrate, the basal part of the endotheca. These species are all apex rounded, lobe bearing multiple sensilla; mesal easily distinguished from one another by the charac- lobe short, digitate, posterodorsally directed, apex teristic and species-specific shapes of the lateral and Blahnik et al.: Chimarra of northern Borneo 153 a b c d

g f

Fig. 31. Chimarra sinitorum, male genitalia. – a, lateral; b, segment IX and tergum X, dorsal; c, inferior appendage, ventral; d, inferior appendage, dorsal; e, phallic apparatus, lateral; f, tergum VIII, dorsal. mesal lobes of tergum X, as well as by differences in absent or nearly so. Preanal appendage setose, small, the shape of the inferior appendages. Among these rounded, slightly narrowed basally. Inferior append- species, Chimarra sinitorum is distinctive both for age much longer than tergum X; in lateral view the shape of the inferior appendages, which have a straight, widest mesally, gradually narrowing distally, very broad enlargement basal to the acute mesal pro- apex rounded, dorsal margin with very basally wide, jection, and also for the narrow lateral lobe of tergum abruptly apically acute projection at midlength; in X and the strongly hooked mesal lobe with one or ventral view somewhat mesally curved, mesal surface two acute apical projections, the bidactylate condi- without processes. Tergum X with sclerotized lateral tion apparently most common. lobes and separate mesal lobes, mesal membrane lack- Adult. Color (in alcohol) yellowish-brown. Length ing; lateral lobe short, in lateral view narrow, apex of forewing: male 4.5–5.3 mm, female 5.0–5.8 mm. downturned, lobe bearing multiple sensilla; mesal Forewing venation: stem of Rs curved, slightly lobe short, digitate, distinctly sclerotized, dorsally sclerotized and thickened at inflection; fork at base of directed, in lateral view, narrow, apex with 1 or 2 discal cell distinctly thickened, nearly symmetrical, acuminate projections. Phallobase tubular, with pro- length of discal cell about 3 times width; m crossvein nounced basodorsal expansion, moderately elongate, proximal to crossveins s and r-m, crossvein s hyaline; relatively narrow, ventral apex lightly sclerotized, 2A vein not intersecting 3A (apparently looped to projecting, dorsal margin extended apically, elongate, 1A). Postocular parietal sclerite short. Maxillary palps narrow, lightly sclerotized. Endotheca moderately moderately elongate, segment 3 slightly greater than elongate, bulbously inflated, with minutely spinu- 2, 3 subequal to 5. Protarsal claws of male enlarged, lose ventral region and numerous short endothe- asymmetrical in shape and length. cal spines in several tracts, 1 basal, 1 apicodorsal, Male genitalia. Abdominal segment VIII short; ter- and 1 apicoventral. Phallotremal sclerite complex gum longer than sternum, unmodified; sternum composed of rod and ring structure, rod relatively VIII without posteroventral projection. Abdominal short, with short apicolateral sclerites. segment IX short dorsolaterally, tergum obsolete dor- Etymology. Named for Gabriel Sinit and his family, somesally, without distinct anterolateral apodemes; in recognition of the friendship and assistance they anteroventral margin in lateral view produced, pos- afforded to J. Huisman during her work and travels terior margin distinctly produced; ventral process on Borneo. 154 Tijdschrift voor Entomologie, volume 152, 2009

a b c d

Fig. 32. Chimarra stenodactylus, male genitalia. – a, lateral; b, segment IX and tergum X, dorsal; c, inferior appendage, ventral; d, inferior appendage, dorsal; e, phallic apparatus, lateral; f, expanded endotheca, dorsal.

Chimarra stenodactylus sp. n. C. prokrustes Malicky, C. scolops, sp. n., and C. sini- Fig. 32 torum, sp. n. All of these species also have a phallic apparatus with numerous small spines and a narrow Type material. Holotype: ?, Malaysia, Sabah, 60 km sclerous dorsal extension, extending over the basal W Lahad Datu, Danum Valley Field Centre, part of the endotheca. These species are all easily Sg. Palum Tambun & vicinity, 04°58’N, distinguished from one another by the characteristic 117°48’E, 150 m, 19–26.iii.1987, J. Huisman and species-specific shapes of the lateral and mesal (UMSP000107286) (RMNH). lobes of tergum X, as well as by differences in the Paratypes. Malaysia, Sabah, same data as holotype, shape of the inferior appendages. Among these spe- 1? (RMNH); Danum Valley Field Centre, nature cies, C. sinitorum is diagnosed by the shape of the trail brooklet, 04°58’N, 117°48’E, 180 m, 9.ix.1986, inferior appendages, which have a very narrow base J. Huisman, 2? (UMSP). to the acute mesal projection, and also by the very short, simple, rounded lateral lobes of tergum X. Chimarra stenodactylus resembles those species from Adult. Color (in alcohol) yellowish-brown. Length Borneo in the Chimarra tsudai group that have an of forewing: male 4.5–4.9 mm. Forewing venation: unmodified, but somewhat widened tergum VIII, stem of Rs curved, slightly sclerotized and thickened an acute projection or tooth on the dorsal margin at inflection; fork at base of discal cell distinctly thick- of the inferior appendages, evident in lateral view, ened, nearly symmetrical, length of discal cell about and mesal lobes of tergum X that are sclerotized, 2 times width; m crossvein proximal to crossveins s upturned and have an acute apex. Species in this and r-m, crossvein s hyaline; 2A vein not intersect- group include C. denticula, sp. n., C. karlijnae, sp. n., ing 3A (apparently looped to 1A). Postocular parietal Blahnik et al.: Chimarra of northern Borneo 155 sclerite short. Maxillary palps moderately elongate, provided with the type description, we are confi- segment 3 subequal to 2 and 5. Protarsal claws of dent of its determination. The form of the inferior male enlarged, asymmetrical in shape and length. appendages, in ventral view, and the dorsolateral pro- Male genitalia. Abdominal segment VIII moderate jections from the posterior margin of segment IX are in length; tergum distinctly longer than sternum, so characteristic and distinctive for the species that modified, with weakly developed V-shaped mesal it could hardly be anything else. Chimarra thaumas depression; sternum VIII without posteroventral is a very distinctive member of the Chimarra tsudai projection. Abdominal segment IX relatively wide group, with several characters that easily diagnose it. dorsolaterally, tergum obsolete dorsomesally, with- The apices of the inferior appendages are especially out distinct anterolateral apodemes; anteroventral diagnostic because each appendage has paired, acute, margin in lateral view slightly produced, ventral apicomesal projections. Segment VIII is also distinc- margin rounded mesally as viewed dorsoventrally; tive in being synsclerous, widened dorsally, and with posterior margin distinctly produced; ventral process a very deep posteromesal emargination that lacks absent or nearly so. Preanal appendage setose, small, accompanying spine-like processes. Other diagnostic rounded, slightly narrowed basally. Inferior append- features include the much enlarged and complex age longer than tergum X; in lateral view straight, phallotremal sclerite complex and the structure of nearly uniformly wide, apex broadly rounded, dor- tergum X, which has short lateral lobes, and divided sal margin with basally narrow, acute projection at mesal lobes, the lateral-most of which is heavily sclero- midlength; in ventral view somewhat mesally curved, tized, bulbous in shape, and with a mucronate apex. mesal surface without processes. Tergum X with scle- Adult. Color (in alcohol) yellowish-brown. Length rotized lateral lobes and separate mesal lobes, mesal of forewing: male 6.0–6.3 mm. Forewing venation: membrane lacking; lateral lobe short, in lateral view stem of Rs curved, slightly sclerotized and thick- simple, rounded apically, lobe bearing multiple sen- ened at inflection; fork at base of discal cell some- silla; mesal lobe short, digitate, distinctly sclerotized, what thickened, nearly symmetrical, length of dis- dorsally directed, apex with multiple, spine-like cal cell about 4 times width; 1st fork proximal to projections. Phallobase tubular, with pronounced crossvein s; m crossvein proximal to crossveins s and basodorsal expansion, moderately elongate, relatively r-m, crossvein s hyaline; 2A vein not intersecting 3A narrow, ventral apex lightly sclerotized, projecting, (apparently looped to 1A). Postocular parietal scler- dorsal margin extended apically, elongate, narrow, ite short. Maxillary palps moderately elongate, seg- lightly sclerotized. Endotheca elongate, tubular; with ment 3 subequal to 2 and 5. Protarsal claws of male 3 tracks of small endothecal spines, basal track of few enlarged, asymmetrical in shape and length. scattered spines, paired, dorsal track of very short Male genitalia. Abdominal segment VIII moderate spines at midlength, apical track of slightly longer in length; tergum distinctly longer than sternum, spines. Phallotremal sclerite complex composed of unmodified, with V-shaped mesal excavation; ster- rod and ring structure, rod very short. num VIII without posteroventral projection. Abdom- Etymology. This species is named C. stenodactylus from inal segment IX moderate in length, with rounded, the Greek words stenos, narrow, and dactylos, a finger, subspinose projections from the posterodorsal mar- used as a noun in apposition, and referring to the dor- gin, tergum obsolete dorsomesally, without distinct sal projection on the inferior appendage, which is very anterolateral apodemes; anteroventral margin in narrow basally compared to other related species. lateral view slightly produced, rounded; posterior margin slightly produced; ventral process absent or Chimarra thaumas Malicky nearly so. Preanal appendage setose, short, length about twice width. Inferior appendage longer than Figs 33, 43 tergum X; in lateral view straight, nearly uniformly Chimarra thaumas Malicky, 2008: 840, holotype ?, In- wide, apex rounded; in ventral view nearly straight, donesia (Eastern Kalimantan), Zoologische Museum slightly mesally curved apically, mesal surface with Lausanne. paired, acute preapical projections. Tergum X with sclerotized lateral lobes and separate mesal lobes, Material examined. Malaysia, Sabah, Crocker Range, mesal membrane lacking; lateral lobe very short, in 5 km N Tenom, base of Sg. ulu Noloyan, 05°10’N, lateral view simple, slightly displaced ventrally, lobe 115°56’E, 1010 m, 10–11.x.1986, J. Huisman 1? bearing multiple sensilla; mesal lobe short, distinctly ? (RMNH), 1 (UMSP). sclerotized, bifid, lateral branch rounded, mucronate, mesal branch digitate, slightly dorsally curved with Despite the several apparent discrepancies between several apical sensilla. Phallobase tubular, with pro- the specimen figured here and the illustration nounced basodorsal expansion, moderately elongate, 156 Tijdschrift voor Entomologie, volume 152, 2009

a b c d

Fig. 33. Chimarra thaumas, male genitalia. – a, lateral; b, segment IX and tergum X, dorsal; dashed line, segment VIII; c, inferior appendage, ventral; d, inferior appendage, dorsal; e, phallic apparatus, lateral; inset, phallotremal sclerite complex, dorsal.

ventral apex weakly sclerotized, slightly projecting. variously armed with spine-like projections. Gen- Endotheca length not discernable (not expanded). eralized features it possesses are a phallotremal scle- Phallotremal sclerite complex composed of elongate rite that is not greatly enlarged, and an endotheca sclerotized structure, more than half length of phal- that is relatively short and lacks distinctive spines, lotheca, apicolaterally with 2 small tack-like spines. although the endotheca is “textured” with minute spines. It is most similar to a group of species from Chimarra vantoli sp. n. Borneo, including C. noloyan, sp. n., C. preapicalis, sp. n., C. physanoton, sp. n., and C. silausilau, sp. n., Fig. 34 all of which have similarly shaped inferior appendages, which are short, linear, with acute projections Type material. Holotype: ?, Malaysia, Sabah, located apicomesally and a tergum VIII that is nar- Crocker Range, 40 km S of Kota Kinabalu, Sin- row and modified with variable lengthed spines. As suron Rd., 1500 m, 18.xii.1989, J. Huisman, in other members of this group, details in the struc- (UMSP000107245) (RMNH). ture of the inferior appendages and lobes of tergum Paratypes. Malaysia, Sabah, same data as holotype, X are important for distinguishing species. Chimarra 1? (UMSP); Kinabalu National Park, Headquar- vantoli is most similar to C. physanoton and C. silausi- ters, Sg. Silau-Silau, 11.viii.2005 & Sungai Liwagu, lau, and best distinguished by its nearly subquadrate 26.ii.2006, W. Mey, 6? (UMHU); Kinabalu inferior appendages, with the apicomesal projections National Park, Silau-Silau, 06°00’N, 116°32’12”E, neither posteriorly directed nor set off by an apical 1560 m, bridge near Bukit Burung trail, 20– notch. Other details distinguishing C. vantoli are 21.xi.1986, J. Huisman, 2? (UMSP). the apices of the lateral lobes of tergum X, which are more acute than C. silausilau, but less distinctively so Chimarra vantoli belongs to the distinctive sub- than C. physanoton. In the material examined, there group of the Chimarra tsudai group in which tergum was some variation in the development of the spined VIII is highly modified, invaginated mesally and processes of tergum VIII, either with the paired Blahnik et al.: Chimarra of northern Borneo 157 a b c d

f e

Fig. 34. Chimarra vantoli, male genitalia. – a, lateral; b, segment IX and tergum X, dorsal; c, inferior appendage, ventral; d, inferior appendage, dorsal; e, phallic apparatus, lateral; f, tergum VIII, dorsal. processes composed of variable lengthed spines, as in short, broad basally, subtriangular. Preanal append- the figure of the holotype specimen, or with each of age setose, very small, rounded. Inferior appendage the paired processes having short lateral spines along about as long as tergum X; in lateral view straight, its length. apex rounded; in ventral view nearly straight, apex Adult. Color (in alcohol) light brown. Length of subtruncate, with apicomesal tooth-like projection. forewing: male 5.9–6.3 mm, female 6.4–6.6 mm. Tergum X with sclerotized lateral lobes and separate Forewing venation: stem of Rs curved, distinctly mesal lobes, mesal membrane lacking; lateral lobe sclerotized and thickened at inflection; fork at base moderately long, in lateral view apex narrow, slightly of discal cell distinctly thickened, nearly symmetri- downcurved, lobe bearing multiple sensilla; mesal cal, length of discal cell about 2.5 times width; lobe elongate, digitate, posterodorsally directed, m crossvein proximal to crossveins s and r-m, cross- apex enlarged. Phallobase tubular, with pronounced vein s not hyaline; 2A vein not intersecting 3A basodorsal expansion, short. Endotheca relatively (apparently looped to 1A). Postocular parietal sclerite short, with granularly textured region; without moderately elongate. Maxillary palps moderately endothecal spines. Phallotremal sclerite complex elongate, segment 3 distinctly longer than 2, 3 sub- composed of rod and ring structure, rod short, api- equal to 5. Protarsal claws of male enlarged, asym- cally with pair of short, curved, lateral sclerites. metrical in shape and length. Etymology. Named for Dr. Jan van Tol, National Male genitalia. Abdominal segment VIII short; ter- Museum of Natural History Naturalis, friend and gum about as long as sternum, modified, with mesal colleague of J. Huisman. excavation, excavation with elongate laciniate fringe divided into 2 branches, sometimes with short lat- Chimarra vanwelzeni sp. n. eral fringe along each of the 2 branches; sternum Fig. 35 VIII without posteroventral projection. Abdominal segment IX relatively wide dorsolaterally, tergum obsolete dorsomesally, without distinct anterolateral Type material. Holotype: ?, Malaysia, Sabah, apodemes; anteroventral margin in lateral view not, Crocker Range, 40 km S of Kota Kinabalu, Sin- or scarcely produced; posterior margin sinuously suron Rd., 1500 m, 18.xii.1989, J. Huisman, produced; ventral process projecting ventrally, very (UMSP000107158) (RMNH). 158 Tijdschrift voor Entomologie, volume 152, 2009

a b c d

Fig. 35. Chimarra vanwelzeni, male genitalia. – a, lateral; b, segment IX and tergum X, dorsal; c, inferior appendage, ventral; d, inferior appendage, dorsal; e, phallic apparatus, lateral; inset, phallotremal sclerite and endothecal spines, dorsal.

Paratypes. Malaysia, Sabah, Kinabalu National Abdominal segment IX relatively wide dorsolater- Park, Headquarters, roadside, 06°00’N, 116°32’E, ally, tergum obsolete dorsomesally, without distinct 1500 m, 12.v.1987, J. Huisman, 1? (UMSP); West anterolateral apodemes; anteroventral margin in lat- Coast, Gn. Kinabalu, roadside, 1500 m, 26.xii.1989, eral view not, or scarcely produced; posterior mar- J. Huisman, 1? (UMSP). gin distinctly produced; ventral process absent or nearly so. Preanal appendage setose, relatively large, Chimarra vanwelzeni does not share an evident simi- subquadrate. Inferior appendage much longer than larity to any other species of the Chimarra tsudai tergum X; in lateral view slightly curved, nearly uni- group from Borneo. It is distinctive in the shape of its formly wide, narrow distally, apex acute; in ventral inferior appendages, which have their apices strongly view nearly straight basally, mesally curved apically, narrowed and mesally curved in ventral view. The apex elongate, narrow, strongly posteromesally bent. enlarged, short, sclerous phallotremal sclerite is also Tergum X with sclerotized lateral lobes and separate distinctive, as are the very short and subdivided lat- mesal lobes, mesal membrane lacking; lateral lobe eral lobes of tergum X. short, bifurcate, in lateral view lateral lobe bifurcate Adult. Color (in alcohol) light brown. Length of into a simple, obovate, strongly downcurved branch forewing: male 7.5–7.8 mm. Forewing venation: and short, digitate mesal branch, lobe bearing multi- stem of Rs curved, distinctly sclerotized and thick- ple sensilla; mesal lobe short, digitate, distinctly scle- ened at inflection; fork at base of discal cell some- rotized, dorsally directed, apex with approximately what thickened, nearly symmetrical, length of dis- 2 short spine-like projections. Phallobase tubular, cal cell about 4 times width; m crossvein proximal with pronounced basodorsal expansion, relatively to crossveins s and r-m, crossvein s not hyaline; 2A short. Endotheca relatively short; with patch of short vein not intersecting 3A (apparently looped to 1A). endothecal spines. Phallotremal sclerite complex Postocular parietal sclerite moderately elongate. Max- composed of short, highly modified rod and ring illary palps elongate, segment 3 slightly greater than structure, with distinctly sclerotized with bifurcating 2, 3 subequal to 5. Protarsal claws of male enlarged, apical branches. asymmetrical in shape and length. Etymology. Named for Dr. Peter van Welzen, Male genitalia. Abdominal segment VIII moderate National Herbarium of The Netherlands, Leiden in length; tergum longer than sternum, unmodified; Branch, University of Leiden, friend of J. Huisman. sternum VIII without posteroventral projection. Blahnik et al.: Chimarra of northern Borneo 159 a b

d c

Fig. 36. Chimarra noohi, male genitalia. – a, lateral; b, segment IX and inferior appendages, caudal; c, phallic apparatus, lateral; f, phallic apparatus, dorsal.

Unplaced species wing are similar to Edidiehlia. Venational characters of C. furti were not provided with its description. Simi- Chimarra noohi sp. n. lar venational characters, except for the absence of an anal loop are found in other species in the subgenus Figs 36, 44 Chimarra, notably the Chimarrhafra group from Africa and C. uvana Kimmins, 1957 from Sri Lanka. Type material. Holotype: ?, Malaysia, Sabah, Another species described below, C. ventritropis, sp. Long Pa Sia, confluence Sg. Pa Sia & Sg. Matang, n. also lacks an anal loop in the hind wing, but in 04°24’N, 115°43’E, 1000 m, 10.iv.1987, J. Huis- other respects bears no resemblance to C. noohi or the man (UMSP000107195) (RMNH). other 2 species discussed. A single foretibial spur pro- vides the character evidence for placing C. noohi in Chimarra noohi is a most unusual species of Chima- the genus Chimarra and the divided lobes of tergum rra. The shape of segment IX, development of the lat- X for placing it in the subgenus Chimarra. The same eral lobes of tergum X, and the shape of the inferior evidence would apply to Edidiehlia hiskia, but because appendages are all distinctive and diagnostic. That both it and C. furti are so unusual and different from the elongate, hornlike processes on the dorsolateral one another, as well as from C. noohi, and because we margin of segment IX are lateral lobes of tergum X have not examined these other species, we are not pro- is suggested by its 2 short setae (or seta-like sensilla). posing any nomenclatural changes. A somewhat similar development of tergum X, also Adult. Color (in alcohol) yellowish-brown. Length of with seta-like projections is found in Chimarra furti forewing: male 3.4 mm. Forewing venation: stem of Mey, 1998 from the Philippines and also in Edidiehlia Rs nearly straight; fork at base of discal cell some- hiskia Malicky, 1993 from Sumatra, although the lat- what thickened, nearly symmetrical, length of discal ter has 4 setae on each lobe (based on the illustration cell about 2.5 times width; m crossvein aligned with provided). Venational characters, forewing with the crossveins s and r-m, crossvein s hyaline; 2A vein s, r-m, and m veins unpigmented and nearly linearly intersecting 3A vein (apparently forked apically). arranged, Rs vein of the forewing lacking the typi- Postocular parietal sclerite short. Maxillary palps cal curvature found in most species of the subgenus relatively short, segment 3 slightly longer than 3, 5 Chimarra, and also the absence of an anal loop in slightly longer than 3. Protarsal claws of male not or the hind wing and the fused Sc and R1 of the hind very little enlarged, symmetrical. 160 Tijdschrift voor Entomologie, volume 152, 2009

a b c d

Fig. 37. Chimarra polyneikes, male genitalia. – a, lateral; b, segment IX and tergum X, dorsal; c, inferior appendage, ventral; d, inferior appendage, dorsal; e, phallic apparatus, lateral; inset, endothecal spines, dorsal.

Chimarra polyneikes Malicky Male genitalia. Abdominal segment VIII short; ter- Figs 37, 45 gum about as long as sternum, unmodified; sternum VIII with slight posteroventral projection. Abdomi- Chimarra polyneikes Malicky, 2008: 840, holotype ?, In- nal segment IX nearly trianguloid in lateral view, donesia (Eastern Kalimantan), Zoologische Museum wide ventrally, narrowing dorsally, tergum obsolete Lausanne. dorsomesally, with pair of short apodemes from anterolateral margin; ventral process absent or nearly Material examined. Malaysia, Sabah, Poring Hot Spring, so. Preanal appendage apparently absent. Inferior 12 km NNE Ranau, Sg. Kepungit, E park bound- appendage about as long as tergum X; in lateral view ary, 06°03’N, 116°42’E, 480 m, 26.xi.1986, J. Huisman ? with acute sclerotized ventral apex and reclinate dor- 1 (RMNH); 12 km NNE Ranau, Poring Hot Spring, Sg. Tananansad, 06°03’N, 116°42’E, 560 m, 28.viii.1986, sal process, narrowing to an acute, posteriorly curved J. Huisman, 4? (UMSP), 2? (RMNH). dorsal apex; mesal surface with acute, sclerotized, tooth-like projection. Tergum X with sclerotized lateral lobes only, mesal membrane lacking; lateral lobe Chimarra polyneikes has a simple morphology that elongate, narrow, apex acute, in lateral view arched might deflect attention from its very unusual com- at base and strongly ventrally curved, lobe bearing bination of characters. The shape of the inferior 2 short setae or seta-like sensilla, preapically. Phallo- appendages, with their chisel-like apices is diagnos- base tubular, with pronounced basodorsal expansion, tic and distinctive. The general form of the inferior moderately elongate, slightly invaginated apicoven- appendages, as well as the presence of 2 sensilla trally. Phallotremal sclerite complex composed of on the lateral lobes of tergum X and 2 endothecal elongate sclerotized structure more than half length spines in the phallic apparatus suggest a relation- of phallobase, complex apparently consisting of elon- ship to the Chimarra digitata group. However, the gate rod and ring structure, with accompanying pair shape of segment IX, without a projecting anter- of elongate, symmetrical, apicolateral sclerites. oventral margin, without pronounced anterodorsal Etymology. Named for Nooh Bin Dawa, in recogni- apodemes, and with a ventral process more ventrally tion of the help and friendship given to J. Huisman than posteriorly directed, would be unusual char- during her studies and collecting on Borneo. acters for a species in the Chimarra digitata group. Blahnik et al.: Chimarra of northern Borneo 161 a b c d

f g

Fig. 38. Chimarra ventritropis, male genitalia. – a, lateral; b, segment IX and tergum X, dorsal; c, inferior appendage, ventral; d, inferior appendage, dorsal; e, inferior appendage, caudal; f, phallic apparatus, lateral; g, phallic apparatus, dorsal; inset, phallotremal sclerite, dorsal.

Additionally, the lateral lobes of tergum X are incom- relatively wide dorsolaterally, tergum relatively long pletely divided dorsomesally, possibly a secondarily dorsally, continuous with base of tergum X, with- derived state and not reflective of a basal position out distinct anterolateral apodemes; anteroventral in the subgenus Chimarra as a whole. Venational margin in lateral view not, or scarcely produced; characters for C. polyneikes are primitive, with the Rs posterior margin nearly straight; ventral process vein of the forewing nearly straight, and the s, r-m, projecting ventrally, broad basally, subtriangular. and m veins linear and hyaline. Like members of the Preanal appendage setose, small, rounded, slightly Chimarra tsudai group, the 2A vein of the forewing is narrowed basally. Inferior appendage about as long looped to the 1A vein. This combination of charac- as tergum X; in lateral view, dorsally inflected baso- ters does not suggest a clear relationship to either of ventrally, widest basally, narrow distally, basoventral the main species lineages discussed in this paper. margin rounded, apex subquadrate; in ventral view Adult. Color (in alcohol) yellowish-brown. Length mesally curved, mesal surface with small, tooth-like of forewing: male 3.4–3.6 mm. Forewing venation: projection at midlength. Tergum X lightly sclero- stem of Rs nearly straight; fork at base of discal cell tized dorsally, apically cleft to from lateral lobes; distinctly thickened, nearly symmetrical, length lateral lobe moderately long, with apex narrowed, of discal cell about 3 times width; 1st fork slightly subacute, lobe bearing 2 sensilla, sensilla at about proximal to s crossvein; m crossvein aligned with midlength, on slightly projecting, lateral protrusions. crossveins s and r-m, crossvein s hyaline; 2A vein Phallobase tubular, with pronounced basodorsal not intersecting 3A (apparently looped to 1A). Hind expansion, moderately elongate, ventral margin wing venation: Sc and R1 fused, first fork absent. slightly bulged basally. Endotheca elongate, tubu- Postocular parietal sclerite short. Maxillary palps lar, granularly textured region not discernable; with relatively short, segment 3 slightly longer than 2, 3 2 elongate endothecal spines, nearly symmetrical in subequal to 5. Protarsal claws of male not or very lit- length and position, apices distinctly flattened. Phal- tle enlarged, symmetrical. lotremal sclerite complex composed of rod and ring Male genitalia. Abdominal segment VIII short; ter- structure, rod short. gum about as long as sternum, unmodified, entire, without mesal excavation. Abdominal segment IX 162 Tijdschrift voor Entomologie, volume 152, 2009

Fig. 39. Chimarra devogeli, wings. – a, forewing; b, hind a wing.

I Fig. 40. Chimarra phlegyas, R1 R2 Sc R3 wings. – a, forewing; b, hind C R R4 wing. a ic r s II R5 Rs r-m M1 III cu m m-cu M2 M3+4 V Cu1a 1A 2A 3A Cu2 Cu1b

b R C I Sc R1 R2 R3 r R4 s Rs II R5 cu-a r-m 1A m-cu III M1 2A M2 3A V M3+4 Cu1a 1A+2A Cu2 Cu1b

Chimarra ventritropis sp. n. Chimarra ventritropis is a distinctive and unusual Figs 38, 46 species of Chimarra, easily diagnosed by the distinctive ventral process of segment IX, dorsal spine-like Type material. Holotype: ?, Malaysia, Sabah, projections of segment IX, and the shape of the infe- Kinabalu National Park, Headquarters, Sg. Liwagu rior appendages. Like species in the Chimarra tsudai crossing Silau-Silau trail, 06°00’N, 116°33’E, group, it has multiple sensilla on the lateral lobes of 1470 m, 15.xi.1986, J. Huisman (UMSP000107152) tergum X, but lacks the sclerotized, digitate mesal (RMNH). lobes that characterize that group. The overall set Paratypes. Malaysia, Sabah, Kinabalu National Park, of characters it possesses suggests that it may repre- Sg. Liwagu, 1100 m, 12.viii.2005, W. Mey, 5? sent a species or lineage basal to the Chimarra tsudai (UMHU); Crocker Range, 40 km S of Kota Kina- group. balu, Sinsuron Rd., 1500 m, 20.xii.1989, J. Huis- Adult. Color (in alcohol) light brown. Length of man, 1? (UMSP). forewing: male 4.8–5.4 mm. Forewing venation: Blahnik et al.: Chimarra of northern Borneo 163

Fig. 41. Chimarra jannekae, wings. – a, forewing; b, hind wing. a

Fig. 42. Chimarra phillipsae, wings. – a, forewing; b, hind a wing.

Fig. 43. Chimarra thaumas, wings. – a, forewing; b, hind a wing.

b 164 Tijdschrift voor Entomologie, volume 152, 2009

Fig. 44. Chimarra noohi, wings. – a, forewing; b, hind wing. a

Fig. 45. Chimarra polyneikes, wings. – a, forewing; b, hind wing. a

Fig. 46. Chimarra ventritropis, wings. – a, forewing; b, hind wing. a

b Blahnik et al.: Chimarra of northern Borneo 165 stem of Rs only slightly curved; fork at base of dis- Dr. Wolfram Mey, Museum für Naturkunde, Hum- cal cell somewhat thickened, nearly symmetrical, bolt-Universität, Berlin for checking the utility of the length of discal cell about 3 times width; m crossvein illustrations and for extending the available records aligned with crossveins s and r-m, crossvein s hya- of examined material. We also thank Dr. Erik J. van line; 2A vein not intersecting 3A (apparently looped Nieukerken for his editorial services, useful sugges- to 1A). Postocular parietal sclerite short. Maxillary tions, and assistance in finding some of the literature palps elongate, segment 3 distinctly longer than 2, cited. This material is based upon work supported 5 longer than 3. Protarsal claws of male not or very by the National Science Foundation grant no. DEB little enlarged, symmetrical. 0117772. Additional support was provided by the Male genitalia. Abdominal segment VIII moderate University of Minnesota Insect Collection under in length; tergum distinctly longer than sternum, projects AES-MIN-17–015 and 17–017. This sup- unmodified; sternum VIII without posteroventral port is gratefully acknowledged. projection. Abdominal segment IX relatively wide dorsolaterally, tergum obsolete dorsomesally, without distinct anterolateral apodemes; anteroventral margin References in lateral view not, or scarcely produced; posterior Banks, N., 1931. Neuropteroid insects from North Bor- margin slightly produced; ventral process project- neo, particularly from Mt. Kinabalu. – Journal of the ing ventrally, forming ventrally rounded, posteriorly Federated Malay States Museums 16: 411–429. subacute lobe, somewhat constricted basally. Preanal Blahnik, R.J., 1998. A revision of the Neotropical species appendage setose, small, rounded, slightly narrowed of the genus Chimarra, subgenus Chimarra (Trichop- basally. Inferior appendage longer than tergum X; in tera: Philopotamidae). – Memoirs of the American Entomological Institute 59: vi+1–318. lateral view dorsally inflected basoventrally, nearly Blahnik, R.J. & R.W. Holzenthal, 2004. Collection and uniformly wide, basoventral margin rounded, apex curation of Trichoptera, with an emphasis on pinned subquadrate, with short apical and preapical teeth; material. – Nectopsyche, Neotropical Trichoptera in ventral view strongly mesally curved. Tergum X Newsletter 1: 8–20. http://www.entomology.umn. with sclerotized lateral lobes only, mesal membrane edu/museum/links/news.html [visited on 15 June lacking; lateral lobe very short, in lateral view sim- 2008]. ple, rounded apically, slightly downcurved, lobe Blahnik, R.J., R.W. Holzenthal & A. Prather, 2007. The bearing multiple sensilla. Phallobase tubular, with lactic acid method for clearing Trichoptera genitalia. pronounced basodorsal expansion, short, preapically – In: J. Bueno–Soria, R. Barba–Alvarez & B. Armitage with dorsolateral flange-like expansions. Endotheca (Eds.), Proceedings of the XIIth International Sympo- length not discernable (not expanded); without sium on Trichoptera: 9–14. The Caddis Press, Colum- endothecal spines. Phallotremal sclerite complex bus, Ohio, USA. composed of rod and ring structure, rod very short, Cartwright, D.I., 2002. The Australian species of Chi- preapically with short lateral sclerites. marra Stephens (Trichoptera: Philopotamidae). – Memoirs of the Museum of Victoria 59: 393–437. Etymology. This species is named C. ventritropis, a Chantaramongkol, P. & H. Malicky, 1989. Some Chi- noun in apposition, from the Latin words venter, marra (Trichoptera: Philopotamidae) from Thailand. belly, and tropis, a keel, and referring to the distinc- – Aquatic Insects 11: 223–240. tive ventral process of segment IX in this species. Kimmins, D.E., 1955. Results of the Oxford Univer- sity expedition to Sarawak, 1934. Order Trichoptera. – Sarawak Museum Journal (New Series) 6: 374–442. Acknowledgments Malicky, H., 1989. Köcherfliegen (Trichoptera) von Sumat- The fieldwork of Jolanda Huisman in East Malaysia ra und Nias: Die Gattungen Chimarra (Philopotami- was made possible through the financial support of dae) und Marilia (Odontoceridae), mit Nachträgen the Melchior Treub Foundation, the Uyttenboog- zu Rhyacophila (Rhyacophilidae). – Mitteilungen der aart-Eliasen Foundation, and the National Museum Schweizerischen Entomologischen Gesellschaft 62: of Natural History (Leiden, The Netherlands). 131–143. Jolanda Huisman is very grateful to Anthea Phillips- Malicky, H., 1993. Neue asiatische Köcherfliegen (Tri- Lamb, Fui-Lian Tan-Inger, Gabriel Sinit, Rienk de choptera: Rhyacophilidae, Philopotamidae, Ecno- midae und Polycentropodidae). – Entomologische Jong, Ed de Vogel, Chan Chu Lun, Anthony Lamb, Berichte Luzern 29: 77–88. Peter van Welzen, Jan van Tol, and Nooh Bin Dawa, Malicky, H., 1994. Neue Trichopteren aus Nepal, Viet- for their help and support, and to her daughters, nam, China, von den Philippinen und vom Bismarck– Karlijn and Janneke Holzenthal. Her gratitude is Archipel (Trichoptera). – Entomologische Berichte expressed in the etymologies of several of the spe- Luzern 31: 163–172. cies described above. Gratitude is also extended to 166 Tijdschrift voor Entomologie, volume 152, 2009

Malicky, H., 1995. Neue Köcherfliegen (Trichoptera, In- P. Thamsenanupap & D. Thapanya, 2004. 27 neue secta) aus Vietnam. – Linzer Biologische Beiträge 27: Köcherfliegen aus Thailand (Insecta, Trichoptera). 851–885. – Linzer Biologische Beiträge 36: 287–304. Malicky, H., 2000. Einige neue Köcherfliegen aus Sabah, Malicky, H., P. Chantaramongkol, P. Chaibu, P. Tham- Nepal, Indien und China (Trichoptera: Rhyacophi- senanupap & I. Thani, 2000. Acht neue Köcherfliegen lidae, Hydrobiosidae, Philopotamidae, Polycentro- aus Thailand. – Braueria 27: 29–31. podidae, Ecnomidae, Psychomyiidae, Hydropsychi- Malicky, H., P. Chantaramongkol, N. Changthong & dae, Brachycentridae, Odontoceridae, Molannidae). P. Thamsenanupap, 2005. Neun neue Köcherfliegen – Braueria 27: 32–39. aus Thailand (Trichoptera). – Linzer Biologische Bei- Malicky, H. 2008. Köcherfliegen (Insecta, Trichoptera) träge 37: 597–604. aus der Umgebung von Malinau (Kalimantan, Bor- Malicky, H., P. Chantaramongkol, N. Changthong, neo, Indonesien). – Linzer Biologische Beiträge 40: A. Nuntakwang & P. Thamsenanupap, 2006. Besch- 833–879. reibungen einiger Köcherfliegen aus Nord–Thailand Malicky, H. & P. Chantaramongkol, 1993a. Neue (Trichoptera). – Braueria 33: 39–42. Trichopteren aus Thailand. Teil 1: Rhyacophilidae, Malicky, H. & T. Prommi, 2006. Beschreibungen eini- Hydrobiosidae, Philopotamidae, Polycentropodidae, ger Köcherfliegen aus Süd–Thailand (Trichoptera). Ecnomidae, Psychomyidae, Arctopsychidae, Hydropsy- – Linzer Biologische Beiträge 38: 1591–1608. chidae. – Linzer Biologische Beiträge 25: 433–487. Mey, W., 1998a. Contribution to the knowledge of Malicky, H. & P. Chantaramongkol, 1993b. Neue Tri- the caddisfly fauna of the Philippines, III (Insecta: chopteren aus Thailand. Teil 2: Rhyacophilidae, Trichoptera). – Entomofauna 19: 1–32. Philopotamidae, Polycentropodidae, Ecnomidae, Mey, W., 1998b. Die Köcherfliegenfauna des Fan Si Pan– Psychomyidae, Xiphocentronidae, Helicopsychidae, Massivs in Nord–Vietnam. 3. Beschreibung weiterer Odontoceridae. – Linzer Biologische Beiträge 25: neuer Arten (Trichoptera). – Opuscula Zoologica 1137–1187. Fluminensia 165: 1–17. Malicky, H., & P. Chantaramongkol, 1997. Weitere neue Ross, H.H., 1956. Evolution and classification of the Köcherfliegen (Trichoptera) aus Thailand. – Linzer Bi- mountain caddisflies. – University of Illinois Press, ologische Beiträge 29: 203–215. Urbana, Illinois, USA. 213 pp. Malicky, H., & P. Chantaramongkol, 2003. Vierzehn neue Köcherfliegen aus Thailand (Insecta, Trichoptera). – Linzer Biologische Beiträge 35: 915–925. Malicky, H, P. Chantaramongkol, P. Bunlue, N. Chang- Received: 26 June 2008 thong, J. Nawvong, A. Nuntakwang, T. Prommi, Accepted: 11 May 2009