Cyphopisthes dohertyi ( Paulian, 1942 )

Cyphopisthes dohertyi ( Paulian, 1942) View in CoL

Figs. 1–5 View FIGURE 1 View FIGURE 2 View FIGURE 3 View FIGURE 4 View FIGURE 5

Philharmostes Dohertyi Paulian, 1942: 71 View in CoL (incorrect original spelling, description)

Cyphopisthes dohertyi View in CoL : Paulian 1978: 512 (new combination, redescription, key); Ocampo & Ballerio 2006: 185 (catalogue)

Type material examined. Holotype: type / Doherty / India Or. Manipur / Fry Coll. 1905-100 / Cyphopisthes Dohertyi sp. nov. det. R. Paulian 1931 ( BMNH) . Paratype: Doherty / India Or. Manipur / Fry Coll. 1905-100 / Cotype / Cyphopisthes Dohertyi Paul. (MNHN) .

Additional material examined. 1♂, 2♀, CHINA: Guangxi, Baise, Napo County, Mt. Meilin [ DZüƜ], 1145 m, leg. Jiaheng Chen ( ZCLC) ; 4♂, 3♀, CHINA: Guangxi, Baise, Napo County, Pocong Village [ 坡丛ñ], 1229 m, leg. Lu Qiu ( LQC) ; 1♂, 1♀, CHINA: Guangxi, Baise, Napo County, Pocong Village [ 坡丛ñ], 1229 m, leg. Lu Qiu ( ABCB) .

Redescription. Small-sized Ceratocanthinae : HL: 0.8 mm, HW: 1.3 mm, PL: 1.0 mm, PW: 2.0 mm, EL: 2.4 mm, EW: 1.2 mm, BL: 4.2 mm. Body brown to black, without metallic sheen ( Figs. 1 View FIGURE 1 , 2A, B View FIGURE 2 ). Glabrous (in 20x magnification). Volant.

Head wider than long (W/L ratio: 1.33), subpentagonal, with short transverse dense comma-shaped punctures around frons; frons smooth, with small shallow sparse simple punctures. Clypeus triangular, with apex sharp; distally with several distinct transversal lines. Genae protruding outwards, genal canthus almost complete, shortly interrupted before reaching occipital area, interocular distance about seven times the maximum width of dorsal ocular area, ventral ocular area larger than dorsal ocular area.Antennae ( Fig. 2C View FIGURE 2 ) short, with ten antennomeres; scape long and elongate, about half the length of total antennae length, slightly curved and apically clavate; pedicel short and plump; antennal club distinctly longer than funicle (antennomeres 2 to 7). Labrum ( Figs. 2D, E View FIGURE 2 ) with anterior margin bisinuate, distally truncate by a slight carina bearing a transverse row of long fine erect setae. Mandibles ( Fig. 2G View FIGURE 2 ) with mesal brush with long setae. Maxilla ( Fig. 2F View FIGURE 2 ) with lacinia oval, with dense setae; maxillary palpus four segmented.

Pronotum transverse (W/L ratio: 2.05), with maximum width at middle. Anterior angles triangular. Dorsal surface covered with relatively large ocellate dense punctures on disc (sometimes absent on a smooth longitudinal area on disc), becoming mixed to horseshoe-shaped punctures with opening outwards at pronotal sides, each puncture with a small setigerous pore at middle. Interpunctural distance inferior than punctural diameter (sometimes equal to puncture diameter only on disc). A symmetrical slight depression present basally at each side of disc.

Scutellum wider than long, triangular. Covered with dense punctures, punctures at middle ocellate or ocellate mixed to horseshoe-shaped.

Legs. Protibiae ( Fig. 2H View FIGURE 2 ) slightly swollen at middle. Outer margin smooth, distally with two teeth in female and with one tooth in male. Inner margin with setae, apical spur short. Mesotibiae ( Fig. 2I View FIGURE 2 ) slightly narrower at distal third. Outer margin with short setae. Inner margin with few setae, seta at middle longer and stronger than the others, two apical spurs present, straight in females and with the inner one bent inwards in males. Metatibiae ( Fig. 2J View FIGURE 2 ) subtriangular, inner margin slightly sinuate; both outer and inner margins with setae, setae at inner margin stronger than those at outer margin; two apical spurs present.

Elytra slightly longer than wide (W/L ratio: 0.94), subsquared in dorsal view, slightly convex. Humeral callus indistinct, without longitudinal lines. Elytral punctation dense (interpunctural distance inferior than punctation diameter); proximal third covered by dense horseshoe-shaped punctures with opening backwards; median third covered by relatively elongate dense horseshoe-shaped punctures often mixed to relatively elongate ocellate punctures, punctation along elytral suture made of elongate comma-shaped punctures or identical to the one of the remaining surface of elytra; distal third covered by dense ocellate punctures. Peseudoepipleura medially with elongate transverse horseshoe-shaped punctures, with a narrow opening upwards, mixed to transverse elongate ocellate punctures, distally with only transverse elongate ocellate punctures. Marginal area (area between striated articular area and inferior sutural stria) present ( Fig. 2B View FIGURE 2 ), relatively narrow, starting at about humeral callus, about one-third of elytral length. Striated articular area present ( Fig. 2B View FIGURE 2 ), relatively narrow and short.

Male genitalia ( Guangxi specimens only) ( Figs. 3A–C, F View FIGURE 3 ). Aedeagus overall weakly sclerotized, with only parameres well sclerotized: parameres sharp and distinctly curved (lateral view); ventral extensions of the parameres present, sclerotized and curved ventrad. Phallobase membranous; internal sac with a flagellum-like curled narrow long temone present. Spiculum gastrale ( Fig. 3D View FIGURE 3 ) elongate and oblique. Female genitalia ( Guangxi specimens only): C-shaped spermatheca, with two constrictions proximally.

Sexual dimorphism. Protibiae: female with two outer apical teeth, male with only one outer apical tooth. Mesotibiae: inner apical spur of males curved inwards, in female inner apical spur is straight.

Distribution and habitat. Northeast India ( Manipur) ( Paulian 1942), new record for China ( Guangxi). Paulian (1942) wrongly indicated “ Assam, Manikpur” and corrected the mistake in 1978 into “ Assam, Manipur ”. It must be noticed that the current Indian Manipur State in 1942 was part of the Assam Province. The locality labels of the holotype and of the single paratype clearly indicate “ India Or. Manipur ” ( Fig. 5D View FIGURE 5 ). The Guangxi localities fall within the Northern Indochina Subtropical Forests ecoregion, while the State of Manipur falls within the Mizoram- Manipur-Kachin Rain Forests ecoregion ( Wikramanayake et al. 2002). The Guangxi specimens were found in a secondary montane broadleaf evergreen forest characterized by the high diversity of understory vegetation, and the wet and rainy climate.

Biology. Representatives of the genus Cyphopisthes are known to occur in termite nests, in leaf litter or are caught through flight intercept traps ( Ballerio 2013; Wang 2025). However, in Guangxi, on Mount Meilin, individuals of this species were collected standing on green leaves of alive plants, along a road in the undergrowth of a secondary montane broadleaf forest during night ( Fig. 4C View FIGURE 4 ).

Remarks. Cyphopisthes Gestro, 1898 is a genus currently comprising 13 species ( Paulian 1978; Ballerio 2013; Ballerio & Grebennikov 2016; Wang 2025). The genus occurs in the Oriental and Australasian Regions, with a wide geographic range spanning from Northeast India to Southern Queensland, with a doubtful record from New Caledonia ( Paulian 1978; Ballerio 2013; Ballerio & Grebennikov 2016) ( Fig. 7 View FIGURE 7 ). The phylogenetic analysis based on morphological data by Ballerio & Grebennikov (2016) supported a clade formed by Cyphopisthes , Ebbrittoniella Martínez, 1962 , and Paulianostes Ballerio, 2000 , confirming the hypothesis proposed by Ballerio (2000b). This group of genera shares the truncate labrum, with a distinct plate-like structure (see Fig. 2D and E View FIGURE 2 of this paper and also Fig. 4b View FIGURE 4 in Ballerio 2000a). Within this group of three genera, the adults of Cyphopisthes are characterized by the combination of a) pronotum regularly convex (not saddle-shaped in lateral view), b) fore angles of pronotum triangular, c) humeral callus not marked by tubercles or cariniform processes, d) antennal club with antennomeres longer than funicle (excluding scape) and e) W/L ratio of mesotibiae equal to or inferior than 0.2 (Ballerio 2000; Ballerio & Grebennikov 2016; Jiang et al. 2020). The genus Cyphopisthes was erected by Gestro (1898). In 1942 Paulian ascribed all Gestro’s species to the genus Philharmostes Kolbe, 1895 , and described Philharmostes dohertyi from India. Later, again Paulian (1978) came back to Gestro’s classification, designated Synarmostes amphicyllis Sharp, 1875 as the type species, and provided a key to the ten species of Cyphopisthes known till then. Subsequently, he described two more species, i.e., C. krikkeni Paulian, 1980 from Sumatra and C. inexpectatus Paulian, 1981 from the Philippines. Ballerio (2000a) transferred C. gestroi to the genus Ebbrittoniella , shortly after ( Ballerio 2000b) he redefined the genus Cyphopisthes , transferred C. acromialis (Pascoe, 1860) and C. georyssoides Gestro, 1898 to Paulianostes and suggested that C. inexpectatus Paulian, 1981 was not a true Cyphopisthes . Subsequently, Ballerio (2013) described other two new species from Queensland, Australia. Currently the two main centers of diversity of the genus are the Sundaic Region, with four species, and the Australasian region with five species ( Fig. 7 View FIGURE 7 ), but our understanding of the diversity of the genus is affected by the lack of a complete revision, made difficult by the shortage of available material and by the very homogeneous morphology of the genus, with the identification of species mostly relying on fine details of dorsal punctation and sculpturing, often difficult to assess.

There are some slight differences between the Guangxi specimens of C. dohertyi and the holotype and paratype from Manipur (to our knowledge the only two known specimens of C. dohertyi before the discovery of the Guangxi population, Figs. 5A–C View FIGURE 5 ). The two populations are set apart by more than 1,500 km but, despite the distance, we were unable to find reliable and constant external morphology characters to separate them as distinct species (the holotype and paratype are too old and fragile for allowing dissection therefore their genitalia were not checked). It must be stressed that in the concerned area there are other species of Ceratocanthinae which display wide distributional ranges, e.g., spanning from NE India to the whole Indochinese peninsula (e.g., Madrasostes feae (Gestro, 1898) ( Ballerio & Bezděk 2016) and, to a lesser extent, M. tonkinensis (Paulian, 1945) ( Ballerio & Maruyama 2025)) . The Guangxi specimens show slightly denser punctation on pronotal disc, more rounded horseshoe-shaped punctures on elytra, the presence of some ocellate punctures mixed to horseshoe-shaped punctures on elytra, and a larger number of ocellate punctures mixed to horseshoe-shaped punctures on pronotum, but we deem all those differences too weak to warrant a separate species status and therefore prudently avoid to describe the Guangxi specimens as a new species. Cyphopisthes dohertyi can be differentiated from the other known species of the genus as follows: C. crux (Sharp 1875) has a much sparser, smaller and shallower dorsal pronotal punctation and a different elytral color pattern, with a distinctive cross-like dark spot, C. szentivanyi Paulian, 1973 has three longitudinal rows of dense recumbent setae on elytra, C. minutus Paulian, 1978 has a very small dorsal ocular area, C. luzonicus Paulian, 1978 has the proximal third of elytral parasutural area with short transverse comma-shaped punctures and pronotal punctation more spaced-out, all the remaining species have one or two longitudinal lines departing from elytral humeral callus and shortly extending backwards and which are absent in C. dohertyi . Currently the species most similar to it is C. erlangshen (see below under C. erlangshen for differential characters).

The aedeagus of Cyphopisthes species is characterized by the presence of small sharp parameres and is weakly sclerotized, especially the phallobasis, hence its observation is often difficult, in particular when extracted from dry specimens or specimens preserved in ethanol for a long time (ethanol hardens soft structures, making their observation difficult). In the case of the Chinese specimens of Cyphopisthes dohertyi the aedeagus has a distinctive structure which was not previously recorded for Ceratocanthinae : all over the length of the internal sac there exists a flagellum-like sclerotized structure, which we tentatively interpret as a modified temone ( Figs. 3A–C, F View FIGURE 3 ). The discovery of this structure by the first author led the second author to dissect other species of the genus in his collection, and a species from Borneo ( Cyphopisthes cf. wallacei ) had a similar structure, although more pronounced ( Fig. 6 View FIGURE 6 ), which looks like a distally sclerotized temone-like structure. The availability of fresh material representing more species could hopefully shed more light on the exact nature of this structure.