Revision of the genus Wakarumbia ( Coleóptera : Lycidae )

A revision of the genus Wakarumbia Bocák, 1999 from Sulawesi is presented. Altogether 10 species are included in the genus Wakarumbia: the type species W. gracilis Bocák, 1999 is redescribed, a new combination of Wakarumbia celebensis (Kleine, 1933) is proposed (originally placed in Protaphes Kleine, 1926) and the following new species are described: Wakarumbia brendelli sp. n., W. brunnescens sp. n., W.flavohumeralis sp. n., W. grandis sp. n., W. nigra sp. n., W. oculata sp. n., W. pallescens sp. n. and W. similis sp. n. The important diagnostic characters are illustrated and all species are keyed. Relationship between species and eco­ logical data are briefly discussed.


INTRODUCTION
The genus Wakarumbia was proposed for one species collected on Buton Island, near the south-east coast of Su lawesi (Bocak, 1999).This genus is closely related to Hemiconderis Kleine, 1926 with which it forms the sub tribe Hemiconderina within the tribe Metriorrhynchini (Lycidae: Metriorrhynchinae).Thanks to M.J.D. Brendell and P.M. Hammond from the Natural History Museum, London, I was able to study the material collected in Northern Sulawesi during the Wallace Project organised in 1985 by the Royal Entomological Society of London (Knight, 1988).This material was collected by different methods and one of the main goals was to estimate insect diversity in this area and, therefore, the material was sorted by the staff of the Natural History Museum in Lon don on the basis of the external morphology and colour patterns into morphospecies.It is now possible to com pare this estimate with the real number of species.Addi tionally, studying Kleine's collection in Warsaw revealed a species belonging to the genus Wakarumbia, previously classified by Kleine (1933a, b) in Protaphes (Lycidae: Erotinae).

MATERIAL AND METHODS
All morphological measurements were made using the ocular grid of an Olympus SZX-12 binocular microscope.Data given in the paragraph "measurements" refer to the holotype (except W. celebensis), the sizes given in descriptions express the vari ability of specimens, when a series was available.Eye diameter was measured in lateral view and when the eye outline was not circular the diameter was measured at the widest point.The in terocular distance when measured from above was the minimum distance between eyes.The ratios were calculated by dividing points measured by ocular grid and, therefore, they can differ from those obtained by division of data given in descriptions in millimetres.Line illustrations were derived from microphoto graphs produced by an Olympus DP-10 digital camera.
Differential diagnosis.The genus Wakarumbia differs from the remaining Metriorrhynchini genera in the fully developed primary elytral costae, the absence of secon dary costae, the characteristic shape of phallus with unique complex spine-like structure in the basal part and the spermaduct attached to the characteristic pocket in the apical part of vagina .The closely related ge nus Hemiconderis Kleine, 1926 differs externally in having secondary elytral costae, a different arrangement of spines in male genitalia and a different shape of vagina (Bocak & Bocakova, 1990a).The taxa of the tribe Conderini have similar arrangement of pronotal costae but the relatively distant position of both groups is supported by numerous characters (Bocak & Bocakova, 1990b).

Distribution and relationship within genus
The genus Wakarumbia now comprises 10 closely re lated species known from a very restricted area of North ern Sulawesi (Dumonga-Bone National Park and Gunung Ambang Forest Reserve; 8 species), Southern Sulawesi (1 species collected at the end of last century) and from Bu ton Island (1 species).The closely related genus Hemi conderis is known from the Papuan Subregion only, and taking into account the fact that some parts of Sulawesi have a common tectonic history with the northern part of New Guinea, the genus Wakarumbia very probably origi nated in the area of islands which form the geologically older part of the present Papuan Region (Hall & Blundell, 1996).The colour patterns correspond with those of the syntopically occurring species of the genus Plateros Bourgeois (Platerodini) but differ substantially from those of Plateros species from Great Sundas.I have frequently noted during field work geographically restricted Mulle rian mimicry complexes in Lycidae and the Wakarumbia species belong to a group of Papuan mimicry complexes.
All species are very similar in external appearance and inconspicuously coloured.Small differences are found in the shape of the pronotum and the elytral reticulate cells, but both characters vary.The only reliable characters for identification and determining relationships are the shape of genitalia and size of the eyes in males.The primitive condition in the closely related genus Hemiconderis, as well as in whole tribe Metriorrhynchini, is the symmetri cal shape of the male phallus without any rotation of the apical part.A unique asymmetry was found in several species of Wakarumbia.The base of the ventral opening of the phallus can be asymmetrical with an unrotated api cal part as in W. niger  or the asymmetry of the basal part of opening can be more marked and accom panied by a distinct rotation of the apical part as in W. pallescens .There is a marked rotation of the phallus in W. grandis , W. celebensis Kleine (Fig. 24) and W. flavohumeralis (Fig. 27).The body colour pattern and the size of the eyes are variable in this group but are not similar to those in groups defined on the basis of male genitalia.A unique origin of the small eyes of the males in the group of darkly coloured species (W.nigra, W. similis and W. grandis) is improbable because this group includes species with both symmetrical and ro tated male genitalia.Unfortunately, there are no other re liable characters because females, whose genitalia usually provide such characters, are unknown for many species.

Diversity and ecology
Most lycid species were collected individually or by sweeping of the lower tree strata of the tropical forest.The fauna of the higher strata is very poorly collected and generally large samples collected by quantitative methods are not available for most regions.The material described here indicates that only a small part of the diversity has been collected and that our knowledge can be substan tially improved when new collecting methods are used.Only 6 specimens of the genus Wakarumbia are cited in the literature (Kleine, 1933a, Mroczkowski, 1959, Bocak, 1999) and no additional specimens are present in the visited European museums.
The expedition of the Royal Entomological Society of London (Project Wallace 1985: Knight, 1988) to the Northern Sulawesi province of Indonesia collected 27 specimens which were sorted by the staff of the Natural History Museum in London on the basis of external char acters into 8 species.Closer examination based on male genitalia and the size of male eyes gave identical results in three species (all of them were represented by one specimen only), two species were split in two, and two species are lumped into one.In addition, the material in cluded three females that belong to different not yet de scribed species, one of them was originally designated a separate species by BMNH staff and two were excluded from the longer series of originally designated species.Although they can be distinguished from all known spe cies by their colour patterns and minute external charac ters, I decided not to describe them as new species because of potential problems with their identification when more species are collected in the future.Altogether I recognised 11 species in this material, 8 of which are described in this article.
Many of the specimens bear labels with data about col lecting methods and habitat; collecting method was avail able for 19 of 27 specimens.Although this information is very limited, it reveals something about the ecology of the group.The most successful method was a Malaise trap, which caught 11 specimens.This indicates a quite high flight activity.As 10 of the 11 specimens were females it is possible they were searching for oviposition sites.The remaining specimens were collected either on leaves (2 k, 12) or they were fogged from the high strata of the can opy (2 k, 3 2).No species for which a series of specimens was collected were taken exclusively by only one of these methods, therefore, I can not allocate individual species to the stratum categories used by Hammond et al. (1997) (tree crown specialist, stratum generalist or lower level specialist).It is very probable that adults of Wakarumbia species migrate between tree crowns and the lowest stra tum, where they seek decaying wood, which is the larval habitat for most Lycidae.
Three species, W. pallescens and two undescribed fe males (altogether 9 specimens), were collected in lowland forest (200-400 m a. s. l.), the others (16 specimens) were collected in lower montane forests (980-1,400 m), and two specimens at a higher elevation (1,780 m, Gng.Muajat summit area).The species distribution supports the re sults presented by Stork & Brendell (1990).There is no overlap between the Wakarumbia fauna of the lowland and lower montane forest, and the highest diversity is found in the mid-altitude zone.The absolute number of specimens is highest in the mid-altitude habitats, but as I do not have any data on the total material collected at dif ferent elevations during the British expedition, no conclu sion can be drawn from these data.My personal experience indicates that lycids are most common in Su matra and Malaysia in the lower montane Lithocarpus-Castanopsis forests, and it is likely that a similar distribution occurs in Northern Sulawesi even though the species composition of the forests there is different.Note.The species of the genus Wakarumbia differ mostly in the size of the eyes in males (they are uniform in females) and shape of male genitalia.The differences in colouration are insuf ficient in many cases for reliable identification.The characters available in both sexes were used in the key as far as possible, but some species had to be based on male characters only.
Female.Unknown.Differential diagnosis.W. brendelli differs from simi lar W. pallescens in the shape of male genitalia (compare Figs 18 and 22, 23) and transverse reticulate cells on ely tra.
Measurements.Male.Length of body 6.1 mm, width at hu meri 1.33 mm, length of pronotum 0.87 mm, width of pronotum 1.13 mm, interocular distance 0.39 mm, maximum diameter of eyes 0.52 mm.
Wakarumbia brunnescens sp.n.Description Male.Body small, slender, only very slightly widened posteriorly.Thorax brown, distinctly lighter pro-and mesothorax, head dark brown, only frontal part slightly lighter, pronotum, scutellum and elytra light brown with elytra slightly infuscated at apex.Head small, shining, with long pubescence, eyes large, hemispherically promi nent, maximum eye diameter 1.19-1.39times the intero cular distance.Antennae longer than three quarters of elytral length, strongly compressed, segments parallel sided.Pronotum trapezoidal, 1.38 times wider than long, with acutely projected hind angles (Fig. 10).Scutellum flat, bilobed at apex.Elytra flat, parallel-sided, with pri mary costae 2 and 4 strong over whole length, costae 1 and 3 shortened, in apical part interrupted or considerably weakened in some parts.Legs slender, strongly com pressed.Male genitalia slender, with robust basal part and with very narrow ventral bridge (Figs 13,14).
Female.Interocular distance 1.03 times longer than maximum eye diameter.
Differential diagnosis.W. brunnescens is the only lightly coloured species other than W. pallescens and W. brendelli.In this species the colouration of the pronotum and elytra varies between light and moderately dark shades of brown, but it is never clearly yellow as in W. pallescens and W. brendelli, and the male genitalia of W. brunnescens differ substantially in shape (Figs 13,14) from those in the other species.
Measurements.Male.Length of body 6.2 mm, width at hu meri 1.38 mm, length of pronotum 0.92 mm, width of pronotum 1.28 mm, interocular distance 0.40 mm, maximum diameter of eyes 0.47 mm.
Etymology.The specific name refers to the brown coloura tion of the dorsum of its body.

Description
Male.Body medium sized, slightly dilated posteriorly.Thorax light brown, head brown, pronotum light brown with infuscated patch in middle of disc and slightly dark ened pronotal carinae, elytra dark brown, only small hu meral part slightly lighter, but transition between lighter and darker part very gradual and unclear.Abdomen dark brown, legs with pale trochanters and basal half of femora and dark rest of femora, tibiae and tarsi.Head small, shin ing, antennal tubercles small but followed by shallow transverse depression, eyes relatively large, hemispheri cally prominent, interocular distance equal to maximum eye diameter, antennae longer than three quarters of ely-tral length, weakly serrate.Pronotum trapezoidal, 1.22 times wider than long, with median areola connected with basal margin through costa (Fig. 5), basal angles only slightly prominent, disc shining, covered with short pu bescence.Scutellum flat, bilobed at apex, densely pubes cent.Elytra slender, flat, with primary costa 2 and 4 strong in whole length, costae 1 and 3 much weaker, but only very slightly shorter, transverse costae dense, form ing distinctly transverse elytral cells.Legs strongly com pressed, slender.Male genitalia slender, apical part rotated by 360 degrees against base of phallus (Fig. 24).
Female.Unknown.Differential diagnosis.W. celebensis is the only spe cies in the genus in which the diameter of eyes in the male is equal to the interocular distance, has a light brown pronotum and very small, light humeri compared with the dark elytra.An advanced structure of male genitalia of W. celebensis indicates a close relationship with W. grandis and W.flavohumeralis .
Measurements.Paratype, k.Length of body 7.1 mm, width at humeri 1.49 mm, length of pronotum 1.19 mm, width of pro notum 1.35 mm, interocular distance 0.50 mm, maximum di ameter of eyes 0.50 mm.
Remark.Wakarumbia celebensis was described by Kleine (1933a) in the genus Protaphes Kleine, 1926 (Taphina, Erotinae) on the basis of the shape of pronotal fields.Kleine did not dissect the genitalia the structure of which places this species in the subfamily Metriorrhynchinae.

Wakarumbiaflavohumeralis sp. n. Description
Male.Body small, slender, parallel-sided.Prothorax and mesothorax light brown, metathorax, head, abdomen and legs except trochanters and bases of femora distinctly darked.Pronotum and scutellum yellow, humeral quarter of elytra bright yellow, with light coloured elytral costae and dark bottom of the reticulate cells in more than hu meral third of elytra, rest of elytra dark brown.
Head small, shining, with sparse long pubescence, an tennal tubercles small, flat but followed by distinct trans verse depression.Eyes very large, hemispherically promi nent, maximum eye diameter 1.67 times interocular dis tance.Antennae strongly compressed, weakly serrate (Fig. 3).Pronotum trapezoidal, only 1.11 times wider at base than long at midline, with nearly rectangular hind angles (Fig. 9).Scutellum flat, densely pubescent, deeply bilobed at apex.Elytra slender, flat, with primary costa 2 and 4 strong over whole length, costae 1 and 3 much weaker, but only very slightly shorter, transverse costae dense, forming apparently transverse elytral cells.Legs slender, strongly compressed.Male genitalia slender with rotated apical part of phallus (Fig. 27).
Female.Unknown.Differential diagnosis.W.flavohumeralis belongs, ac cording to the male genitalia, to the group of advanced species along with W. grandis and W. celebensis.It dif fers from them in the distinctly larger eyes in the male and the colouration of pronotum and elytra.
Measurements.Male.Length of body 6.3 mm, width at hu meri 1.71 mm, length of pronotum 0.98 mm, width of pronotum 1.11 mm; interocular distance 0.33 mm, maximum diameter of eyes 0.55 mm.
Etymology.This species is named after the colour pattern on its elytra.

Wakarumbia gracilis Bocák, 1999
Redescription Male.Body small, slender, yellow to brown, only ely tra and antennae dark brown, individual parts of body tend to be infuscate to a variable extent, especially basal part of pronotum except margin and costae, tibiae and femora in apical part and tarsi.Head small, shining, with sparse, short pubescence, antennal tubercles inconspicu ous, frons flat to slightly concave, eyes hemispherically prominent, diameter of eyes 1.24-1.30times the distance between them, head including eyes slightly wider than frontal margin of pronotum, maxillary palpi slender, api cal segment compressed, wider, obliquely cut at apex, la bial palpi short, apical segment wide, cut at apex, apical margin with 4-6 tubercle-shaped projections.Antennae not longer than three fifths of the length of elytra, com pressed, covered with dense, dark pubescence.Disc of pronotum shining, with sparse pubescence.Pronotal carinae forming narrow rhomboidal areola in middle of the disc.Scutellum longer than wide, flat, deeply emarginate at apex, densely pubescent.Elytra very slightly widened posteriorly, with four completely developed primary costae, costa 2 and 4 strong over whole length, costa 1 and 3 much weaker, stronger and shining at humeri only, very weak and even occasionally interrupted posteriorly, transverse costae well developed, areolae mostly quadrate.Legs apparently flattened, trochanters long, slender, without any projections.Male genitalia slender, symmetrical, quite robust at base and narrowed in middle, internal sac indistinct, without setae or pigmentation.
Female.Vagina long, with very short and indistinct median sac.Paired glands attached in middle part of va gina.Interocular distance 1.03 times maximum eye di ameter.
Differential diagnosis.W. gracilis is easily distin guishable by the body colouration from the other species of the genus.The symmetrical structure of the male geni talia indicates a basal position in this genus along with W. niger and a close relationship to W.brunnescens.
Measurements.Male.Length of body 4.6 mm, width at hu meri 0.88 mm, length of pronotum 0.62 mm, width of pronotum 0.76 mm, interocular distance 0.36 mm, maximum diameter of eyes 0.29 mm.

Wakarumbiagrandis sp. n. Description
Male.Body medium sized, slender, slightly widened posteriorly.Whole body dark brown to black, only tro chanters, basal part of femora, mandibles, bases of palpi and very narrow outer margins of pronotum lighter brown.Head small, shining, with small antennal tubercles and shallow transverse depression behind them, eyes small, hemispherically prominent, interocular distance 1.15 times greater than maximum eye diameter.Antennae strongly compressed, antennal segments robust, segment 3 only 1.65 times longer than wide, antennae as long as half of elytral length.Pronotum trapezoidal, 1.23 times 3/4 at base than maximum length, with prominent hind angles (Fig. 7).Scutellum deeply bilobed at apex, shining, with dense, short pubesce.Elytra flat, with strong primary costae 2 and 4 and much weaker costae 1 and 3, transverse costae dense, forming reticulate cells twice as wide as long, costae with fine, short setae.Legs slender, strongly compressed.Male genitalia slender, apical part of phallus strongly rotated (Figs 25,26).
Female.Unknown.Differential diagnosis.W. grandis, W. similis, W. ocu lata and W. nigra are the only species with completely dark brown to black dorsal part of body.W. grandis has the largest body and differs in the strongly rotated apical part of phallus (Figs 25,26), the long tubular basal part of phallus, and the slender and distinctly transverse elytral reticulate cells.
Measurements.Male.Length of body 7.5 mm, width at hu meri 1.53 mm, length of pronotum 1.05 mm, width of pronotum 1.29 mm; interocular distance 0.54 mm, maximum diameter of eyes 0.44 mm.
Etymology.The specific name "grandis" refers to its being the largest species in the genus.

Wakarumbia nigra sp.n. Description
Male.Body small, parallel-sided to slightly widened posteriorly.Whole body dark brown to black, only mouth parts, trochanters and bases of femora lighter.Head small, shining, antennal tubercles flat, eyes small, but hemi spherically prominent, interocular distance 1.25-1.28times maximum eye diameter.Antennae longer than two thirds of elytral length, strongly compressed, antennal segments nearly parallel-sided.Pronotum trapezoidal, 1.24 times wider than long, with acutely projected hind angles (Fig. 12).Scutellum, flat, bilobed at apex, shining.Elytra with equal strength of longitudinal costae in hu meral half.Primary costae slightly shortened, costae 1 and 3 weaker in apical part, transverse costae less dense and reticulate cells quadrate.Legs long, slender, strongly compressed.Male genitalia with basal part of phallus bulb-like (Figs 15,16).
Differential diagnosis.W. nigra belongs to the group of dark species and differs in having quadrate reticulate cells and a robust symmetrical phallus (Figs 15,16).It differs from the very similar W. similis in the shape of the male genitalia, bulb-like at base rather than slender (Fig. 17).
Measurements.Male.Length of body 5.55 mm, width at humeri 1.09 mm, length of pronotum 0.78 mm, width of pronotum 0.97 mm; interocular distance 0.41 mm, maximum diameter of eyes 0.32 mm.
Etymology.The specific name refers to the black colouration of the body.

Wakarumbia oculata sp. n. Description
Male.Body small, slender, nearly parallel-sided.Whole body dark brown to black, only trochanters, mouth parts and very narrow margins of pronotum brown to light brown.Head small, shining, sparsely pubescent, antennal tubercles well developed, with a shallow transverse depression behind them, eyes large, hemispherically prominent, maximum eye diameter 1.36 times interocular distance, antennae slender, weakly serrate, longer than two thirds of elytral length.Pronotum trapezoidal, 1.28 times wider than long (Fig. 11), disc only slightly shining, rather densely pubescent.Scutellum bilobed at apex, parallel-sided, slightly shining.Elytra with strong primary costae 2 and 4, much weaker and in apical third often interrupted costae 1 and 3, elytral reticulate cells slightly transverse.Legs slender, strongly compressed.Male genitalia bulb-like at base, symmetrical (Figs 19,20).
Female.Unknown.Differential diagnosis.W. oculata is similar to other dark species but differs in males that have much bigger eyes and in the shape of their phallus (Figs 19,20).
Measurements.Male.Length of body 6.35 mm, width at hu meri 1.30 mm, length of pronotum 0.89 mm, width of pronotum 1.14 mm; interocular distance 0.36 mm, maximum diameter of eyes 0.50 mm.
Etymology.The specific name "oculata" refers to the very large eyes ofmales.
Differential diagnosis.W. pallescens is similar to W. brunnescens and W. brendelli but differs in shape of male genitalia (Figs 22,23).It also differs from W. brunnes cens in being light yellow in colour and from W. brendelli in having quadrate elytral reticulate cells.Etymology.The specific name refers to the pale yellow col ouration.

Wakarumbia similis sp. n. Description
Male.Body small, parallel-sided to slightly widened posteriorly.Whole body dark brown to black except tro chanters and mouth parts.Both longitudinal and trans verse elytral costae slightly lighter than bottom of elytral cells.Head small, shining, with small antennal tubercles, eyes small, but hemispherically prominent, interocular distance 1.21-1.25 times maximum eye diameter.Anten nae slender, longer than three quarters length of elytra, antennal segments nearly parallel-sided.Pronotum trape zoidal, 1.21 times wider than long, with acutely promi nent hind angles.Scutellum flat, deeply bilobed.Elytra with primary costae 2 and 4 strong and reaching apex of elytra, costae 1 and 3 very weak, slightly shortened.Legs strongly compressed, slender.Male genitalia with slender base ofphallus (Figs.17).
Female.Interocular distance 1.52 times maximum eye diameter.Female genitalia with long, slender vagina, en larged basal part of spermaduct and slim spermatheca (Fig. 28).Differential diagnosis.W. similis differs from the very similar W. nigra in having a much slenderer basal part of phallus (compare Figs 15,16 and 17).
Measurements.Male.Length of body 5.0 mm, width at hu meri 1.0 mm, length of pronotum 0.74 mm, width of pronotum 0.89 mm; interocular distance 0.37 mm, maximum diameter of eyes 0.31 mm.
Etymology.W. similis is very similar to W. nigra sp.n. and the specific name refers to this fact.
Remark.The second male differs slightly from the other specimens as it lacks the lighter elytral costae and has more dense transverse costae.The male genitalia are almost identical with those of the holotype and therefore it is provisionally identified as W. similis but it is not in cluded in the type series.