Research Article |
Corresponding author: Yaheita Yokoi ( y.yokoi@kddnet.de ) Academic editor: Marianna Simões
© 2023 Yaheita Yokoi.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Yokoi Y (2023) Note on genitalia and taxonomy of the Callidiopini from the Philippines, with description of six new species and two subspecies (Coleoptera, Cerambycidae, Cerambycinae). Contributions to Entomology 73(1): 33-66. https://doi.org/10.3897/contrib.entomol.73.e101117
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Species of Ceresium and Examnes, Callidiopini, from the Philippines were reviewed. Six new species, Ceresium elongatipenne sp. nov., Ceresium holzschuhi sp. nov., Ceresium jimcopei sp. nov., Ceresium nigricolle sp. nov., Ceresium quinquemaculatum sp. nov., Ceresium crassum sp. nov. and one new subspecies, Ceresium huedepohli mindanao subsp. nov., are described. Of Examnes, one new subspecies, Examnes lumawigi subrugosus subsp. nov., is described. Male genitalia of eight Ceresium species and three Examnes species, in particular their endophalli and 8th sternites, are described and illustrated in detail. Female genitalia of three Ceresium species are described. The genitalia and the consequences for taxonomy are discussed.
Ceresium und Examnes Arten, Callidiopini, von den Philippinen wurden überprüft. 6 neue Arten, Ceresium elongatipenne sp. nov., Ceresium holzschuhi sp. nov., Ceresium jimcopei sp. nov., Ceresium nigricolle sp. nov., Ceresium quinquemaculatum sp. nov., Ceresium crassum sp. nov. und eine neue Unterart, Ceresium huedepohli mindanao subspec., werden beschrieben. Von Examnes wird eine neue Unterart, Examnes lumawigi subrugosus subsp. nov., beschrieben. Männliche Genitalien von acht Ceresium und drei Examnes Arten, insbesondere ihre Endophalli und 8ten Sternite, werden näher beschrieben und illustriert. Weibliche Genitalien von 3 Ceresium Arten werden beschrieben. Die Genitalien und die Folgerungen für die Taxonomie werden erörtert.
Apical process, Ceresium, 8th sternite, endophallus, Examnes, genitalia, Philippines
Callidiopini, Ceresium, Die Philippinen, 8ter Sternit, Endophallus, Examnes, Genitalien
The Callidiopini, Lacordaire, 1869, is a large tribe, mainly distributed in the Oriental and Austro-Asian Regions. While it is well explored with more than 400 hitherto described species in 68 genera, the knowledge of their genitalia has remained rudimentary for a long time. In recent years, however, many of their genitalia were newly described, in particular those of species belonging to the genera Ceresium Newman, 1842, Examnes Pascoe, 1869, Oxymagis Pascoe, 1866, Tethionea Pascoe, 1869, and Falsoibidion Pic, 1922. (
In the course of these observations, a few interesting aspects of their genitalia were disclosed. First, their 8th sternites are highly diversified in morphology. They are characteristic of each species. Second, their endophalli are even more significant for the taxonomy. With the exception of those Falsoibidion species, the endophalli are each furnished with a strongly sclerotized apical process surrounding the ejaculatory duct. These processes are extraordinarily intricate in structure, highly diversified, and characteristic of each species. In addition, the surface structure of the endophallic tube has proved to be another essential species indicator. Comparative observation of genitalia has thus become indispensable for the taxonomy of these genera and species. Therefore, investigation of genitalia should be further extended and intensified, involving more species and genera of the Callidiopini.
Ceresium as a genus was introduced by Newman 1942, with the descriptions of four species from the Philippines. Since then, a great number of species have been added to the genus. It is now by far the largest and most widely distributed genus of the Callidiopini. At present, it consists of more than 180 taxa in three subgenera, distributed in the Oriental and Austro-Asian Regions, spreading also into the insular and coastal regions of the Indian Ocean.
In contrast, the Ceresium fauna of the Philippines, the original locality of the genus, remained less explored for a long time, with only one species described in one-and-a-half centuries (
In this publication, six new species and one new subspecies of Ceresium are described from the Philippines. They include Ceresium elongatipenne sp. nov., Ceresium holzschuhi sp. nov., Ceresium jimcopei sp. nov., Ceresium nigricolle sp. nov., Ceresium quinquemaculatum sp. nov., Ceresium crassum sp. nov. and Ceresium huedepohli mindanao subsp. nov.
As to the male genitalia of Ceresium, 28 species from the Oriental and Austro-Asian Regions have been previously described (
The female genitalia are less indicative than those of the males. Most of their parts are not sclerotized and thus susceptible to deformation before and during the examination. However, sclerotized parts as 8th and 9th sternites and tergites as well as spermathecae and vaginal plates often reveal characteristics typical of the species, providing us with useful keys for the taxonomy. In this publication, female genitalia of Ceresium elongatipenne sp. nov., Ceresium quinquemaculatum sp. nov. and Ceresium crassum sp. nov. are described. For comparison, those of Ceresium furtivum Pascoe, 1869, and Ceresium striatocolle Holzschuh, 2011, are additionally illustrated.
Examnes Pascoe, 1869, is a much smaller genus than Ceresium and less widely distributed, consisting of 13 hitherto described species from the Oriental and Austro-Asian Regions. Four species of this genus were already examined as to male genitalia (
The observations of genitalia beyond the individual descriptions are discussed at the end, together with their consequences for taxonomy.
The specimens examined were either directly compared to the holotypes or to their original descriptions and photographs.
For the examination of genitalia, the specimens were first softened, then submerged in KOH solution for 12 hours and subsequently cleansed in water. They were photographed by a digital camera with a macro lens. Serial photos were stacked onto one image. Drawings were made by Adobe Illustrator. Median lobe, tegmen, 8th sternite and tergite as well as the endophallus were observed for each male specimen. In addition, when relevant, 9th sternites or tergites were described. In the previous publications of the author, the apical sclerotization of the endophallus was referred to as “ejaculatory duct complex”. In this publication, it is instead referred to as “apical process of endophallus.” In general, the side on which the ejaculatory duct or its protective capsule is located, is referred to as “dorsal”.
For genital analysis, males are preferred. Where male specimens were not available or a female is additionally available, female genitalia were examined, in particular their 8th and 9th sternites and tergites, spermathecae, spermathecal ducts and bursae copulatrix. Measurements were taken with an ocular micrometer.
The abbreviations used in the descriptions are as follows: TL – total length (from apical margin of clypeus to abdominal apex); HW – head width across eyes; PL – length of pronotum; PW – maximum width of pronotum; PA – apical width of pronotum; PB – basal width of pronotum; EL – length of elytra; EW – humeral width of elytra. In addition to these commonly applied ratios, EL/TL is also indicated for numeric comparisons of elytral length in relation to body.
Data on the collection labels are below reprinted verbatim.
Holotypes and paratypes of the species described in this publication are preserved in the Senckenberg Museum, Frankfurt.
CCH Collection Carolus Holzschuh, Villach, Austria.
CJC Collection Jim Cope, San Jose, California, USA.
CHM Collection Hiroshi Makihara, Chiba, Japan.
CTN Collection Tatsuya Niisato, Tokyo, Japan.
CNO Collection Nobuo Ohbayashi, Kanagawa, Japan.
CYY Collection Yaheita Yokoi, Ratingen, Germany.
Holotype
♂ (
Paratype
♀ (
The name of this species refers to its elongated elytra.
Slender. Colour testaceous. Elytra parallel-sided, distinctly elongated. Pronotal disc densely punctured. Blade of 8th sternite extremely short. 8th tergite strongly elongated.
Holotype ♂: TL = 10.8 mm; EL/TL = 0.73; EL/EW = 3.27; HW/PW = 0.97; PL/PW = 1.06; PA/PW = 0.73; PB/PW = 0.88. Paratype ♀: TL = 10.8 mm; EL/TL = 0.74; EL/EW = 3.30; HW/PW = 1.0; PL/PW = 1.06; PA/PW = 0.76; PB/PW = 0.92.
Colour universally testaceous, thorax slightly darker; setae pale yellowish to whitish.
Head nearly as broad as pronotum, glossy; punctures dense and regular; setae stout, curved, disorderly, pointing in different directions; median furrow from frons to occiput. Frons longer than broad; rather flattened; not steep; apex nearly as broad as base; sides moderately emarginated. Vertex moderately concave. Occiput finely sub-rugose. Eyes separated from one another by 5/9 the width of occiput or 3/2 that of the upper eye-lobes. Antennal supports broad, flattened, impunctate, each sub-encircled by an arcuate row of diminutive setiferous punctures. Antennae reaching elytral apices at antennomere X. Length of antennomeres relative to scape: III = 1.0; IV = 0.95; V = 1.4; VI = 1.25; VII = 1.25; VIII = 1.15; IX = 1.04; X = 0.95; XI = 1.05. Scapes sub-arcuate and strongly clavate.
Pronotum (Fig.
A–I. Ceresium elongatipenne sp. nov. Holotype male. A. Habitus, dorsal view; B. Ditto, ventral; C. Pronotum; D. Prosternum and mesoventrite; E. 8th sternite, ventral view; F. 7th sternite and 8th tergite, ventral view; G. 8th sternite and tergite, ventral view; H. Median lobe and endophallus, lateral view; I. Apical process of endophallus, dorsal view.
Scutellum sub-circular; fringed by diminutive setae.
Elytra elongated in relation to the body; sub-parallel-sided; apices separately rounded; punctures regular, dense, setiferous; setae on disc finer than the others.
Legs rather slender; femora clavate from base.
Venter (Fig.
Antennae reaching elytral apices at the middle of antennomere XI. Abdomen more voluminous than in male.
Female genitalia
(Figs
Luzon, Philippines.
This new species can be distinguished from all congeneric species by the parallel-sided, strongly elongated elytra. They are not only proportionally elongated in themselves, but also in relation to body, with EL/TL = 0.73–0.74. As to male genitalia, the structures of its 8th sternite and tergite are, as above described, exceptional in Ceresium. Apical process of its endophallus is elaborate and characteristic of the species.
Holotype
♂ (
The name of this species is dedicated to Mr. Carolus Holzschuh of Villach, Austria. The holotype belonged to his private collection.
Slender. Colour fulvus to testaceous. Elytra elongated, apically tapering. Punctures on pronotal disc deep and regular. Apical middle of 8th sternite elevated, distinctly setose.
Holotype ♂: TL = 10.5 mm; EL/TL = 0.70; EL/EW = 3.25; HW/PW = 0.93; PL/PW = 1.12; PA/PW = 0.74; PB/PW = 0.94.
Colour fulvus; thorax and head testaceous; abdomen reddish dark yellow; setae pale yellowish to whitish.
Head narrower than pronotum; glossy; punctures dense and regular; setae dense, stout, curved, recumbent; median furrow stretching from frons to vertex. Frons transverse, rather flattened, not steep; apex as broad as base; sides moderately emarginated. Vertex moderately concave. Occiput finely sub-rugose. Eyes separated from one another by half the width of occiput or 11/5 that of the upper eye-lobes. Antennal supports flattened; each sub-encircled by an arcuate row of diminutive setiferous punctures. Antennae surpassing the pronotal base at antennomere IV. Length of antennomeres relative to scape: III = 0.95; IV = 0.9; V = 1.25; VI = 1.25. Scape sub-arcuate and strongly clavate. Remark: Antennae damaged; antennomeres VII–XI missing.
Pronotum (Fig.
Scutellum bell-shaped; with dense pubescence.
Elytra elongated, sides distinctly tapering toward apices; apices separately rounded; punctures deep, regular, setiferous; setae curved and adpressed.
Legs slender, femora clavate from base.
Venter (Fig.
Male genitalia
(Figs
Mindanao, Philippines.
This new species can be compared to Ceresium virens Heller, 1914, from Papua New Guinea. Both species are similar in body structure, in particular regarding their characteristic elongated, tapering elytra. The new species can be easily distinguished by the surface of pronotum and elytra: the punctures are regular and deep, whereas they are irregularly aligned and of varying depth in C. virens. In addition, the legs and antennae of the latter are longer and more slender. The new species shares also a few external characteristics with the above described C. elongatipenne sp. nov. Meso-and metathorax, abdomen and legs are essentially similar. In contrast, they obviously differ from each other by the structure and surface of the prothorax. The distinctions in genitalia are even more evident. The endophallus, apical process in particular, of C. holzschuhi is rather reminiscent of Ceresium sugiartoi Yokoi, Makihara & Noerdjito, 2019, or to those of Examnes species (See Discussion). The elevation in the apical middle of its 8th sternite with a cluster of aligned, erect setae thereupon is remarkable, a new observation in the genus.
Holotype
♂ (
The holotype could be examined by courtesy of Mr. Jim Cope, California, USA. The name of this species is dedicated to him.
Colour testaceous. Pronotum resembles that of Ceresium vestigiale Pascoe, 1866. Body, legs and antennae slenderer; antennae longer. Apical process of endophallus exceptionally large, in baculi-form; surface of endophallic tube scaly in part.
Holotype ♂: TL = 10.0 mm; EL/TL = 0.65; EL/EW = 2.90; HW/PW = 0.90; PL/PW = 1.04; PA/PW = 0.66; PB/PW = 0.88.
Colour testaceous; setae pale yellowish to whitish.
Head lightly narrower than pronotum, glossy; puncture dense, large, regular, setiferous; setae stout, curved, sub-erect, disorderly. Frons transverse, not steep; the middle moderately elevated, glossy, obtusely punctate. Vertex concave; with a shallow median furrow; setae pointing to antennal supports. Punctures on occiput obtuse; setae adpressed, pointing to vertex. Eyes separated from one another by half the width of occiput or twice that of the upper eye-lobes. Antennal supports broad, flattened, glabrous, nitid, though each sub-encircled by an arcuate row of diminutive setiferous punctures. Antennae reaching elytral apices by 9th article. Length of antennomeres relative to scape. III = 1.0; IV = 0.85; V = 0.80; VI = 1.1; VII = 1.05; VIII = 1.05; IX = 1.05; X = 0.9; XI = 0.80. Scapes sub-arcuate and moderately clavate.
Pronotum slightly longer than broad; apical half inflated; constricted just before apex; apex thus narrower than base. Puncture on disc dense, regular, setiferous; setae curved, pointing to the middle; median stripe broad, nearly half the length of pronotum. Sides each uneven with irregular elevations, of which the one near the latero-apical corner is the largest.
Scutellum sub-circular, with several short setae.
Elytra gradually tapering from humeri; apices separately rounded; disc well-flattened; punctures dense, regular, setiferous, attenuating towards apices; setae stout, curved and recumbent, though short and fine on disc.
Legs rather slender; femora clavate from base.
Venter (Fig.
Male genitalia
(Figs
Luzon, Philippines.
This new species can be distinguished from the other known species by the combination of its smaller size, universally testaceous colour, slender antennae, apically inflated pronotum and regular puncture on disc. The structure of, and integument on, pronotum can be compared to those of C. vestigiale from Borneo. The new species differs, however, in other external characteristics. Further, its male genitalia is clearly distinct. The apical process of its endophallus, as described above, is indeed exceptional. This type of the process has not been observed before in the genus Ceresium. At the same time, the surface of its endophallus is also atypical. The endophallus of this species is remarkable.
Ceresium huedepohli Yokoi, 2019: 23. Type locality: “Espana, Sibuyan Is., Romblon, Philippines”.
Holotype
♂ (
The name of this subspecies refers to its distribution.
Holotype ♂: TL = 8.8 mm; EL/TL = 0.70; EL/EW = 3.13; HW/PW = 1.04; PL/PW = 1.17; PA/PW = 0.62; PB/PW = 0.80.
(Figs
Mindanao, Philippines.
This subspecies differs from the nominotype in a few external characteristics. Its elytra are longer with EL/TL = 0.70, while antennae are shorter. The differences in male genitalia are more evident. Even though they are basically of similar design, a few essential deviations can be identified as above. The new subspecies, from Mindanao, is geographically separated from the nominotype from Sibuyan Island.
Holotype
♂ (
The name of this species refers to the colour of thorax.
Bi-coloured, head and prothorax dark, remainder of the body yellowish-brown. Pronotum rounded; disc encircled by a ring of whitish pubescence. Antennae slender and long.
Holotype ♂: TL = 15.4 mm; EL/TL = 0.63; EL/EW = 2.94; HW/PW = 0.86; PL/PW = 1.04; PA/PW = 0.65; PB/PW = 0.90.
Body yellowish-brown; head, thorax and apices of femora black; setae whitish.
Head narrower than pronotum; glossy; punctures dense, setiferous; setae stout, curved, disorderly, pointing to different directions. Frons transverse, not steep, emarginated on sides, elevated in middle, median furrow short. Vertex broad, moderately concave. Occiput finely sub-rugose; setae pointing to vertex. Eyes separated from one another by 2/5 the width of occiput or 5/3 of the upper eye-lobes; lower lobes densely fringed with short, sub-erect setae. Antennal supports broad, flattened, sub-nitid, though each sub-encircled by an arcuate row of diminutive setiferous punctures. Antennae surpassing elytral apices by 10th article. Length of antennomeres relative to scape: III = 0.84; IV = 0.80; V = 1.2; VI = 1.25; VII = 1.25; VIII = 1.2; IX = 1.13; X = 1.0; XI = 1.1. Scapes weakly clavate, slightly arcuate.
Pronotum slightly longer than broad; sub-spherical with sides almost evenly arcuate. Disc almost encircled by a ring of thick pubescence; bordered by a nitid, irregularly arcuate costa on each side; punctures dense, regular; setae short, curved; pointing to the middle; median stripe broad, about 3/10 the length of pronotum. Sides with a protuberance near each apical corner; punctures dense, variable; setae sparse, short, curved; (Fig.
Scutellum bell-shaped, pubescent.
Elytra tapering from humeri toward apices; apices separately sub-rounded. Punctures rather small, attenuating towards apices; setiferous. Setae short, stout and curved. Punctures and setae on disc finer.
Legs short and stout. Femora clavate from base on. Tibiae keeled (Fig.
Venter (Fig.
Male external genitalia
(Fig.
Luzon, Philippines.
The new species is bi-coloured. A similar colour pattern is also observed in Ceresium bicolor
Ceresium Aethiops Newman, 1842: 247. Type locality: Manila.
Ceresium Aethiops: White 1855: 244.
Ceresium aethiops:
Ceresium aethiops: Hüdepohl 1994: 93, 94.
1 ♂: “Mt. Banahao, P. l. Baker”; “1914, 14”. (
Venter: Fig.
A–I. Ceresium aethiops Newman. Male from Luzon. A. Habitus, dorsal view; B. Ditto, ventral; C. Prosternum and mesoventrite; D. Protibia, ventral view; E. Elytron; F. Median lobe and endophallus with apical process; G. Endophallus, part, surface; H. Tegmen, apical part of paramere, lateral view; I. Ditto, latero-ventro-frontal view.
Legs: Tibiae keeled (Fig.
Male genitalia
(Figs
Philippines (Manila); Luzon.
The external characteristics of this specimen from Mindanao correspond to the original description of holotype. It should be supplemented that the puncture on its elytra is unusually sporadic (Fig.
Regarding genitalia, the median lobe is more strongly arcuate than usual for a Ceresium species. Apical process of endophallus is characteristic of this species, though it is, with its large, deeply dehiscent apical sclerite, somewhat reminiscent of Ceresium ikuoyokoii Yokoi, 2019, from Sumatra. Multi-sinuate apex of 8th sternite is also characteristic. In addition, the thick, partly sclerotized membrane surrounding the basal parts of sternite and tergite is noteworthy.
Ceresium lingafelteri Vives, 2013: 63. Type locality: “Filipinas, Is. Luzon, Cordillera Prov., Apayo”.
1 ♂ “Calapan, Mindanao”; “Collection B. Schwarzer”; “Senckenberg Museum Frankfurt”. (
(Figs
A–I. Ceresium lingafelteri Vives. Male from Mindanao (New record). A. Habitus, dorsal view; B. Ditto, ventral view; C. Head and pronotum; D. Median lobe with endophallus, lateral view; E. Apical process of endophallus, ventral view; F. Tegmen, lateral view; G. Parameres, ventral view; H. 8th sternite, ventral view; I. 9th sternite (spicum gastrale) and tergite, dorsal view.
Luzon, Philippines; Mindanao (New record).
The setae of this specimen from Mindanao are whitish, and the pronotal sides have a few erect hairs. In other aspects, it corresponds to the original description of the holotype from Luzon.
The genitalia of this species are remarkable in several aspects. First, the dorsal plate of the median lobe is much more deeply dehiscent than usual. It is even more dehiscent than ventral plate, which is atypical. Second, tegmen is extraordinarily bi-sinuate in lateral view, while the external sides of parameres are each clothed with a cluster of bushy hairs. Third, the structure of 8th sternite is of a rare type, similar to that of C. elongatipenne sp. nov. Fourth, the peduncle of 9th sternite (speculum gastrale) is unusually long. These singularities have either not been observed before, or only rarely, in the genus Ceresium. Apical process of endophallus is characteristic of this species.
Ceresium longicorne Pic, 1926: 24. Type locality: “Formosa”.
Ceresium longicorne: Matsushita, 1932: 67, 68.
Ceresium japonicum Matsushita, 1932: 67, 68.
Ceresium tsushimanum Ohbayashi, 1961: 17.
Ceresium longicorne: Hua, 2002: 200.
1♂1♀, “Sant Lucia, central Palawan, Philippines; Apr. 11, 1989, T. Niisato leg.” (CTN); 1♀, ditto, with datum “Apr.10.1989” (CTN); 1♂2♀, “Trident Mines, Palawan, 3.v. 1989, A. Saito leg.” (CTN); 1♀, “(PHILIPPINES), Languan, 10m, N. Palawan Is. 1.IX. 1985, By light trap, M. Tomokuni” (CNO).
Figs
(Fig.
Taiwan; Japan; Korea; South China; Philippines; Palawan, Philippines (New record).
“Philippines” as a distribution is included in the catalogues by
Compared to the holotype, no essential difference was observed regarding external characteristics. Ceresium longicorne shares the characteristic puncture pattern on elytra with the now sympatric Ceresium immite Newman, 1842. In both species, the punctures in the apical halves of elytra are diminutive, in clear contrast to the normal ones in the basal halves, so that the surface of each elytron is optically divided in apical and basal halves. As this type of elytral surface is observed solely in these two cases, both now sympatric species are probably affiliated, though clearly differing in pronotal surface (Figs
More interesting for the taxonomy is the commonly shared structure of the head. The fronts of both C. longicorne and C. immite are short and steep (Figs
Holotype
♀ (
The name of this species refers to its altogether five maculae on pronotum and scutellum.
Colour yellowish brown. Pronotum evenly rounded; with four tomenta-maculae; surface longitudinally striate-retuculate. Scutellum with tomentum. Elytra apically tapering.
Holotype ♀: TL = 10.2 mm; EL/TL = 0.68; EL/EW = 2.70; HW/PW = 0.84; PL/PW = 0.95; PA/PW = 0.66; PB/PW = 0.82.
Colour yellowish brown; pronotum and head darker; setae pale-yellowish.
Head narrower than pronotum; glossy; densely punctured or sub-reticulate; setae sparse, short and fine. Frons transverse, sub-rectangular, flattened, not steep. Vertex broad, shallowly concave; reticulate (Fig.
Pronotum resembles that of C. striatocolle; more evenly rounded, sub-spherical; a little shorter than broad; with two pairs of tomenta-maculae; the latero-apical maculae large, oval, oblique, clearly bordered; the latero-basal ones smaller, with irregular boundaries; setae otherwise sparse, short, stout, disorderly. Surface of disc longitudinally striate-reticulate. Sides distinctly and evenly arcuate, though basal 1/7 constricted; punctures on sides large and deep.
Elytra short, tapering sub-linear toward apices; disc well-flattened. Punctures dense, apically attenuating; setae short and curved.
Legs short and stout; femora clavate from base on.
Venter (Fig.
Female genitalia
(Figs
Visaya, Philippines.
Pronotum of this new species is reminiscent of C. striatocolle from Borneo. Above all, the striate-reticulate disc and lateral tomento-maculae are commonly shared. However, the pronotal sides of the new species are more evenly rounded, while the maculae are larger and more clearly bordered. In addition, it obviously differs in body colour. Further, the head is more densely punctured or sub-reticulate, while the setae are shorter and finer. Female genitalia are analogous, differing in details (Fig.
Holotype
♀ (
The name of this species refers to the large and stout body.
Large. Antennae short. Body colour brownish; femora bi-coloured. Setae on head, pronotum and sides of abdomen erect to sub-erect.
Holotype ♀: TL = 17.5 mm; EL/TL = 0.68; EL/EW = 2.86; HW/PW = 0.78; PL/PW = 1.04; PA/PW = 0.64; PB/PW = 0.86.
Body colour reddish dark brown; pronotum, head, antennae and tibiae darker. Femora reddish brown; apices blackened. Setae whitish.
Head narrower than pronotum; glossy; punctures dense, large, deep, setiferous; setae stout, curved, sub-erect, disorderly, pointing to different directions. Frons transverse, reversed-sub-trapezoidal, not steep; the apical middle weakly elevated, sparsely punctured; apical corners sub-reticulate. Vertex flattened, feebly concave. Punctures on occiput large near eyes, obtuse otherwise. Eyes separated from one another by half the width of occiput or twice of the upper eye-lobes. Antennal supports flattened; punctate and setose. Antennae short, hardly reaching the 5th sternite; antennomeres III and IV each 4/5 as long as scape; V and VI equally as long; VII–XI gradually decreasing in length; XI as long as scape. Scapes sub-cylindrical; slightly arcuate.
Pronotum slightly longer than broad, well rounded; sides almost evenly arcuate; punctures dense and regular; median stripe short and narrow; setae stout, curved, erect to sub-erect.
Elytra feebly tapering toward apices; basal middle elevated; punctures dense and regular, apically attenuating; setae short and curved.
Legs: Femora clavate from base; bi-coloured; apices blackened and obtusely punctured.
Venter (Fig.
A–F. Ceresium crassum sp. nov. Holotype female. A. Habitus, dorsal view; B. Ditto, ventral; C. Pronotum; D. Prosternum and mesoventrite; E. 8th sternite-tergite, apices, frontal view; F. Genitalia, clockwise from top: median oviduct; vaginal plates; bursa copulatrix; spermatheca; vagina; far left: 9th sternite.
Female genitalia
(Figs
Luzon, Philippines.
The new species is comparable to Ceresium furtivum Pascoe, 1869, from the Sunda Islands. However, it is much larger, and its antennae distinctly shorter. Further, its setae on head, pronotum and abdomen are erect to sub-erect instead of recumbent. In addition, the coloration pattern of femora is distinct, rather resembling the sympatric Ceresium femoratum Aurivillius, 1927. The latter differs, however, in other essential characters such as structure of body, legs and punctures. As to female genitalia, its 8th sternite and tergite are shorter than those of C. furtivum, not parallel-sided. Setae and puncture thereupon are also different. Above all, their apices are thickened (Figs
Examnes lumawigi Hüdepohl, 1990: 88. Type locality: ♂, “Luzon, Mountain province”; ♀, “Sibuyan, Espana”.
Examnes lumawigi:
Luzon and Sibuyan, Philippines.
Holotype
♂ (
The name of this subspecies refers to the pronotal surface.
This subspecies differs from the nominotype in the structure and integument of pronotum.
Holotype ♂: TL = 10.0 mm; EL/TL = 0.67; EL/EW = 2.85; HW/PW = 0.90; PL/PW = 1.02; PA/PW = 0.76; PB/PW = 0.92.
Body smaller. Pronotum with PL/PW = 1.02 slightly longer than wide, instead of “distinctly longer than wide”; surface obtusely rugose; several erect setae on each side (Fig.
(Figs
Luzon, Philippines.
The pronotal structure and surface of this subspecies differ substantially from those of the nominotype Examnes lumawigi lumawigi (Fig.
Apical process of its endophallus is essentially of similar structure as that of E. philippensis, nominotype of the genus Examnes. Differences are concerned with the structures of sides and base as well as of appendage (Figs
Examnes kawakamii
1♂: “Basilan Philippin”; “Coll-Schwarzer”; “Senckenberg-Museum, Frankfurt”. (
(Figs
Endophallus of this species is described above, as it was not included in the previous description of the holotype from Borneo (
Borneo; Basilan, Philippines (New Record).
No essential difference from the holotype from Borneo is observed. Though the apex of 8th sternite is less strongly emarginated, the deviation is tolerable as a variation within a species. Apical process of its endophallus is essentially of similar structure as that of E. philippensis. Differences are concerned with the structures of sides and base as well as of appendage (Figs
1♂ “(Bazilan)”; “Maloong”; “vii-viii. 1982 ; « K. Kuwasima »”. (CHM).
(Figs
This specimen from Basilan was examined for comparative purposes. Genitalia of this species is described above, as endophallus was not included in the previous description (
A–L. Ceresium elongatipenne sp. nov. Holotype male. Genitalia. A. Median lobe with endophallus, dorsal view; B. Ditto, lateral; C. Ditto, ventral; D. Endophallus, apical process, dorsal view; E. Ditto, lateral; F. Ditto, ventral; G. Tegmen, dorsal view; H. Ditto, lateral; I. Ditto, ventral; J. 8th sternite, ventral view; K. Ditto, with 8th tergite in the background; L. 7th and 8th tergite, dorsal view. Scale bars: 0.5 mm (D–F); 1 mm (A–C, G–L).
A–K. Ceresium holzschuhi sp. nov. Holotype male. Genitalia. A. Median lobe with endophallus, dorsal view; B. Ditto, lateral; C. Ditto, ventral; D. Endophallus, apical process, dorsal view; E. Ditto, lateral; F. Ditto, ventral; G. Tegmen, dorsal view; H. Ditto, lateral; I. Ditto, ventral; J. 8th sternite and tergite, ventral view; K. Ditto, fronto-lateral view. Scale bars: 0.2 mm (D–F); 1 mm (A–C, G–K).
A–K. Ceresium jimcopei sp. nov. Holotype male. Genitalia. A. Median lobe with endophallus, dorsal view; B. Ditto, lateral; C. Ditto, ventral; D. Endophallus, apical process, dorsal view; E. Ditto, lateral; F. Ditto, ventral; G. Tegmen, dorsal view; H. Ditto, lateral; I. Ditto, ventral; J. 8th sternite, ventral view; K. Ditto, with 8th tergite in the background. Scale bars: 0.5 mm (D–F); 1 mm (A–C, G–K).
Luzon, Bohol, Mindanao (Philippines), Basilan, Philippines (New record); Bali; Maluku; Waigeou; West Ilian; Papua (Indonesia); Papua-Newguinea; Vanikoro (Solomon Islands).
A–J. Ceresium huedepohli mindanao subsp. nov. Holotype male. Genitalia. A. Median lobe with endophallus, dorsal view; B. Ditto, lateral; C. Ditto, ventral; D. Endophallus, apical process, dorsal view; E. Ditto, lateral; F. Ditto, ventral; G. Tegmen, dorsal view; H. Ditto, lateral; I. Ditto, ventral; J. 8th sternite, with 8th tergite in the background, ventral view. Scale bars: 0.5 mm (D–F); 1 mm (A–C, G–J).
A–K. Ceresium aethiops Newman. Male from Luzon. Genitalia. A. Median lobe with endophallus, dorsal view; B. Ditto, lateral; C. Ditto, ventral; D. Endophallus, apical process, dorsal view; E. Ditto, lateral; F. Ditto, ventral; G. Tegmen, dorsal view; H. Ditto, lateral; I. Ditto, ventral; J. 8th sternite, ventral view; K. Ditto, with 8th and 9th tergite in the background and partly sclerotized membrane in the foreground. Scale bars: 0.5 mm (D–F); 1 mm (A–C, G–K).
The holotype was recorded from “Insularum Manillaum”. There was no further indication. The distribution in Basilan is thus treated as a new record within the Philippines.
A–L. Ceresium lingafelteri Vives. Male from Mindanao (New record). Genitalia. A. Median lobe with endophallus, dorsal view; B. Ditto, lateral; C. Ditto, ventral; D. Endophallus, apical process, dorsal view; E. Ditto, lateral; F. Ditto, ventral; G. Tegmen, dorsal view; H. Ditto, lateral; I. Ditto, ventral; J. 8th sternite, ventral view; K. Ditto, with 8th tergite in the background; L. 9th Sternite (spicum gastrale) and tergite, dorsal view. Scale bars: 0.5 mm (D–F); 1 mm (A–C, G–L).
A–K. Ceresium longicorne PIC. Male from Palawan, Philippines. (New record). Genitalia. A. Median lobe with endophallus, dorsal view; B. Ditto, lateral; C. Ditto, ventral; D. Endophallus, apical process, dorsal view; E. Ditto, lateral; F. Ditto, ventral; G. Tegmen, dorsal view; H. Ditto, lateral; I. Ditto, ventral; J. 8th sternite with 8th tergite in the background; K. Ditto with 9th sternite (spicum gastrale), with 7th tergite in the background. Scale bars: 0.5 mm (D–F); 1 mm (A–C, G–K).
A–L. Examnes lumawigi subrugosus subsp. nov. Holotype male. Genitalia. A. Median lobe with endophallus, dorsal view; B. Ditto, lateral; C. Ditto, ventral; D. Endophallus, apical process, dorsal view; E. Ditto, lateral; F. Ditto, ventral; G. Tegmen, dorsal view; H. Ditto, lateral; I. Ditto, ventral; J. 8th sternite, ventral view; K. Ditto, with 8th tergite in the background; L. 8th tergite, ventral view. Scale bars: 0.5 mm (D–F); 1 mm (A–C, G–L).
Essentially, the external characteristics of this specimen from Basilan correspond to those of the holotype from Luzon. Its setose 9th tergite is noteworthy. Specimens from other localities should be re-examined in this regard.
A–F. Examnes kawakamii Yokoi, Makihara & Noerdjito. Male from Basilan, Philippines. (New record). Genitalia. A. Median lobe with endophallus, lateral view; B. Ditto, ventral; C. Endophallus, apical process, dorsal view; D. Ditto, lateral; E. Ditto, ventral; F. 8th sternite and tergite, ventral view. Scale bars: 0.5 mm (C–E); 1 mm (A, B, F).
A–L. Examnes philippensis Newman. Male from Basilan, Philippines (New record). Genitalia. A. Median lobe with endophallus, dorsal view; B. Ditto, lateral; C. Ditto, ventral; D. Endophallus, apical process, dorsal view; E. Ditto, lateral; F. Ditto, ventral; G. Tegmen, dorsal view; H. Ditto, lateral; I. Ditto, ventral; J. 8th sternite, ventral view; K. Ditto, with 8th tergite in the background; L. 9th sternite (spicum gastrale) and tergite, ventral view. Scale bars: 0.5 mm (D–F); 1 mm (A–C, G–L).
28A, B. Ceresium elongatipenne sp. nov. Paratype female. 28A. 8th sternite, ventral view; 28B. 8th tergite, dorsal view. 29A, B. Ceresium quinquemaculatum sp. nov. Holotype female. 29A. 8th sternite, ventral view; 29B. 8th tergite, dorsal view. 30A–C. Ceresium striatocolle Holzschuh. Female from Borneo. 30A. 8th sternite, ventral view; 30B. 8th tergite, dorsal view; 30C. Spermatheca. Scale bars: 1 mm (28A, B; 29A, B; 30A–C).
Including the previously examined species (
Endophallus–Apical process:
– The apical process of C. jimcopei sp. nov. is exceptional. An apical process in baculi-form has not been observed before, even in the tribe Callidiopini as a whole. It is also extraordinarily large in relation to the median lobe, the largest ever measured. It is noteworthy that the endophallus of C. jimcopei is so peculiar in morphology, while many of its external characteristics are comparable to those of known species.
The endophallus of C. huedepohli and its subspecies is no less remarkable. Its apical process is very narrow, thin, sharply curved, fundamentally differing from those of other species. In contrast, the external characteristics of this species are at least partly comparable to those of the sympatric C. balkei Yokoi, 2019, or of C. gracilipipenne Yokoi, 2021, from the Solomon Islands. The observation can be interpreted either as an additional evidence of the extreme diversity of Ceresium species in terms of the endophallus, or as an indication of the unique status of this species in taxonomy. Examinations of additional species are necessary.
– In general, the apical process of the endophallus is distinctive and characteristic for each Ceresium species studied. In a few rare cases, however, affinity in morphology can be identified. Resemblance between C. holzschuhi sp. nov. and C. sugiartoi is such an example. The processes of both species are in turn comparable to those of Examnes species. Further, the apical process of C. aethiops from the Philippines is analogous in architecture to that of C. ikuoyokoii Yokoi, 2019, from Sumatra. (See corresponding Comparative notes). Nevertheless, the compared species differ in other aspects as external characteristics or 8th sternite.
– So far as the hitherto examined Ceresium species are concerned, morphology of the endophallus is not correlated with morphologies of other essential characteristics. As described above, external characteristics of two species can be comparable in one way or another, while their endophalli are fundamentally different, and vice versa. It is possible that the endophalli were independently diversified in Ceresium, decoupled from the evolution of other essential characteristics. Further investigation is necessary to answer this question. Only 1/6 of the 180 Ceresium species have been so far examined regarding genitalia; the endophallus in particular.
Surface of endophallic tube
Endophallic tubes of a few Ceresium species from the Philippines are scaly on the surface. The scales are diminutive or obtuse in most cases. Those of C. jimcopei sp. nov. are, however, distinct, large, dense, even partly sub-reticulate. This type of endophallic tube was observed in a few Austro-Asian species as Ceresium pulekerai Yokoi or Ceresium furutaraui Yokoi from the Solomon Islands (
Sternite and tergite
New types of 8th sternites and tergites have been observed. In particular, 8th sternite of C. elongatipenne sp. nov. is finely structured with a short, multi-emarginated blade, while the tergite is extraordinarily elongated. No less noteworthy is the distinctly setose apical elevation on the 8th sternite of C. holzschuhi sp. nov., and the unusually setose 8th sternite and tergite of C. nigricolle sp. nov. In turn, peduncle of the 9th tergite is disproportionately elongated in C. lingafelteri.
So far, no correlation of these external genital parts was observed, either to the diverse endophalli or to other essential characteristics.
Median lobe and tegmen
The median lobe and tegmen of C. lingafelteri are atypical (See the Description and Comparative notes). Above all, both clusters of bushy lateral setae on the lobe of parameres are peculiar, a new observation in the Callidiopini.
Male genitalia of one new subspecies and 2 known species from the Philippines are described above. Altogether, five Examnes species have now been examined as to male genitalia (
Although genitalia of Examnes are relatively homogenous, they do include a few interesting deviations. The disproportionately large appendage of the apical process of the endophallus as well as the adjacent perforation, observed in E. kawakamii, is such an example. Likewise noteworthy are the apical setae on the peculiar structure of the 9th tergite of E. philippensis. The function of these setae and their taxonomical significance should further be investigated.
There are a few ambiguities as to the classification of Examnes species. The distinction between Examnes and Ceresium, as originally described (
In any case, types of endophalli should be duly considered as an additional indicator for classification of species in question. In fact, as indicated in the corresponding comparative notes above, Ceresium longicorne shares not only a few essential external characters but also a similar type of male genitalia with Examnes species. The genitalia of Ceresium immite have not yet been examined but at least its external characters are likewise comparable. In view of these affinities, both C. longicorne and C. immite could be better placed in Examnes than in Ceresium. In this publication, however, and until additional studies are performed, both C. longicorne and C. immite are treated as members of Ceresium. The above question as to the taxonomic position of the genera Ceresium and Examnes should be first clarified before any generic synonymies are published.
The female genitalia of five species were examined. Deviations were observed in the structure of the 8th sternites and tergites. They differ in the form of the blades, the relative length of the peduncle (struts) and the apical setae. The 8th sternite and tergite of Ceresium crassum sp. nov., apically thickened, is noteworthy.
The author is indebted to Dr. Damir Kovac and Mrs. Andrea Hastenpflug-Vesmanis, both of Senckenberg Museum Frankfurt, Mr. Jim Cope of San Jose, California, USA, Mr. Carolus Holzschuh of Villach, Austria, Prof emer. Ohbayashi of Kanagawa, Japan, Dr. Tatsuya Niisato, Tokyo, Japan, and Dr. Hiroshi Makihara of Tsukuba, Japan, for providing specimens which formed the material base of this publication. The author would like to thank Mr. Max Barclay, Senior Curator, of the Natural History Museum in London, for enabling the comparative observation of holotypes, and Mr. Keita Matsumoto, Curatorial Assistant, for providing the photographs of C. immite which are illustrated in this publication. He thanks Dr. Klaus-Dieter Klass, Senckenberg Museum Dresden, for useful advice. Last but not least, the author thanks his wife, Karin de Wit-Yokoi, for proof reading and other editorial assistance.