A contribution to the knowledge of the genera Fissocantharis Pic, 1921 and Micropodabrus Pic, 1920 from Taiwan (Coleoptera, Cantharidae)

1 Email: 695163750@qq.com 2 Email: 2940331171@qq.com * Co-corresponding authors: 3 yxyang@hbu.edu.cn, 4 musoku@city.kurashiki.okayama.jp 1 urn:lsid:zoobank.org:author:EF6832A3-76B9-495E-B552-D304D3CAF494 2 urn:lsid:zoobank.org:author:2D7E44B4-1472-43C0-A669-B86433EEA6DE 3 urn:lsid:zoobank.org:author:CA819BAF-F411-4917-9205-4F1C36AC4DC9 4 urn:lsid:zoobank.org:author:BB590861-E5AE-438B-AC0E-A8724F8FAAB5


Supplementary description
Aedeagus (Fig. 1A-C).Ventral process of each paramere wide, slightly narrowed apically, moderately hooked at apex; conjoint dorsal plate of parameres moderately developed, about half length of ventral process and as wide as half of the distance between outer margins of ventral processes, moderately narrowed apically, with straight apical margin.

Remarks
In the original publication (Wittmer 1995), this species was only illustrated with male antennae.Here, the aedeagus is illustrated and described for the first time.

Distribution
Taiwan.

Material examined
Same data as specimens examined in Yang et al. (2012).

Supplementary description
Aedeagus (Fig. 1D-F).Ventral process of each paramere wide, slightly narrowed apically, moderately hooked at apex; conjoint dorsal plate of parameres moderately developed, about half length of ventral process and as wide as one-third of the distance between outer margins of ventral processes, distinctly narrowed apically, with straight apical margin.
Abdominal sternite VIII of female (Fig. 5A).Slightly and roundly emarginate in middle and on both sides of posterior margin.
Internal organ of reproductive system (Fig. 2A).Vagina stout, abruptly thinned and extended into a short duct at ventroapical portion; diverticulum and spermathecal duct arising from the end of the duct of vagina; diverticulum moderately long, spiral and distinctly thinned apically; spermathecal duct slightly shorter than diverticulum and slightly thinner than basal portion of diverticulum; spermatheca  (Wittmer, 1995).D-F.F. denominata (Wittmer, 1997).G-I.F. fenchihuensis (Wittmer, 1982).Scale bars: 1.0 mm.composed of a spiral tube which is slightly longer than diverticulum, provided with a thin accessory gland which is much shorter than the spiral tube of spermatheca; median oviduct situated in middle of vagina.

Remarks
In the original publication (Wittmer 1954), this species was only illustrated with male antennae.Here the genitalia of both sexes and abdominal sternite VIII of female are illustrated and described for the first time.

Distribution
Taiwan.

Supplementary description
Aedeagus (Fig. 1G-I).Ventral process of each paramere wide, distinctly narrowed apically, rounded at apex; conjoint dorsal plate of parameres developed, about two-thirds length of ventral process and as wide as half of the distance between outer margins of ventral processes, slightly widened apically, with straight apical margin.
Abdominal sternite VIII of female (Fig. 5B).Distinctly protuberant in middle of posterior margin, the middle protuberance slightly and triangularly emarginate at apex, forming a pair of median lobe, which are slightly rounded at apices.
Internal organ of reproductive system (Fig. 2B).Vagina stout, abruptly thinned and extended into a slightly long duct at ventroapical portion; diverticulum and spermathecal duct arising from the end of the duct of vagina; diverticulum moderately long, spiral and slightly thinned apically; spermathecal duct distinctly shorter than and thicker than diverticulum; spermatheca composed of a spiral tube which is distinctly longer than diverticulum, provided with a thin accessory gland which is slightly shorter than the spiral tube of spermatheca; median oviduct situated in middle of vagina.

Remarks
In the original publication (Wittmer 1982), the species was illustrated with male antenna and aedeagus.
Here, the aedeagus is illustrated again and described in detail, also the characters of abdominal sternite VIII and genitalia of female are provided for the first time.

Distribution
Taiwan.
Internal organ of reproductive system (Fig. 2C).Vagina stout, abruptly thinned and extended into a slightly long duct at ventroapical portion; diverticulum and spermathecal duct arising from the end of the duct of vagina; diverticulum moderately long, spiral and moderately thinned apically; spermathecal duct slightly shorter than diverticulum and nearly as thick as basal portion of diverticulum; spermatheca composed of a spiral tube which is slightly longer than diverticulum, provided with a thin accessory gland which is slightly longer than the spiral tube of spermatheca; median oviduct situated in middle of vagina.

Remarks
In the present study, the abdominal sternite VIII and genitalia of female are photographed and described for the first time.

Distribution
Taiwan.
Abdominal sternite VIII of female (Fig. 5D).Slightly protuberant in middle of posterior margin, the middle protuberance widely and shallowly emarginate at apex, forming a pair of median lobes, which are slightly arcuate at apices.
Internal organ of reproductive system (Fig. 2D).Vagina stout, abruptly thinned and extended into a slightly long duct at ventroapical portion; diverticulum and spermathecal duct arising from the end of the duct of vagina; diverticulum moderately long, spiral and distinctly thinned apically; spermathecal duct distinctly shorter than diverticulum and narrower than basal portion of diverticulum; spermatheca composed of a spiral tube which is nearly as long as diverticulum, provided with a thin accessory gland which is slightly shorter than the spiral tube of spermatheca; median oviduct situated in middle of vagina.

Remarks
The aedeagus of this species was illustrated in ventral and dorsal views by Wittmer (1982), here it is illustrated again and described in detail.Also, the characters of abdominal sternite VIII and genitalia of female are provided for the first time.

Distribution
Taiwan.

Supplementary description
Aedeagus (Fig. 3D-F).Ventral process of each paramere wide, slightly narrowed apically, slightly hooked at apex; conjoint dorsal plate of parameres slightly reduced, about one fourth length of ventral process and as wide as one-fourth of the distance between outer margins of ventral processes, slightly narrowed apically, with slightly rounded apical margin.
Abdominal sternite VIII of female (Fig. 5E).Distinctly protuberant in middle of posterior margin, the middle protuberance distinctly and roundly emarginate at apex, forming a pair of median lobes, which are narrowly rounded at apices.
Internal organ of reproductive system (Fig. 4A).Vagina stout, abruptly thinned and extended into a slightly long duct at ventroapical portion; diverticulum and spermathecal duct arising from the end of the duct of vagina; diverticulum moderately long, spiral and distinctly thinned apically; spermathecal duct slightly longer than diverticulum and distinctly thinner than basal portion of diverticulum; spermatheca composed of a spiral tube which is nearly as long as diverticulum, provided with a thin accessory gland which is slightly shorter than the spiral tube of spermatheca; median oviduct situated in middle of vagina.

Remarks
In the original publication (Wittmer 1954), this species was only illustrated with male antennae.Here the genitalia of both sexes and abdominal sternite VIII of female are illustrated and described for the first time.

Distribution
Taiwan.

Supplementary description
Aedeagus (Fig. 3G-I).Ventral process of each paramere slightly wide, slightly narrowed apically, slightly hooked at apex; conjoint dorsal plate of parameres moderately developed, about half length of ventral process and as wide as half of the distance between outer margins of ventral processes, moderately narrowed apically, with straight apical margin.
Abdominal sternite VIII of female (Fig. 5F).Distinctly protuberant in middle of posterior margin, the middle protuberance moderately and triangularly emarginate at apex, forming a pair of median lobes, which are widely rounded at apices.
Internal organ of reproductive system (Fig. 4B).Vagina stout, abruptly thinned and extended into a slightly long duct at ventroapical portion; diverticulum and spermathecal duct arising from the end of the duct of vagina; diverticulum moderately long, spiral and moderately thinned apically; spermathecal duct distinctly shorter than diverticulum and thinner than basal portion of diverticulum; spermatheca composed of a spiral tube which is slightly longer than diverticulum, provided with a thin accessory gland which is distinctly shorter than the spiral tube of spermatheca; median oviduct situated in middle of vagina.

Remarks
In the original publication (Wittmer 1982), the aedeagus was illustrated in dorsal view, here it is illustrated in ventral, dorsal and lateral views and described in detail.Its abdominal sternite VIII and genitalia of female are illustrated and described for the first time.

Distribution
Taiwan.

Supplementary description
Abdominal sternite VIII of female (Fig. 5G).Distinctly protuberant in middle of posterior margin, the middle protuberance widely subtriangular at apex.
Internal organ of reproductive system (Fig. 4C).Vagina stout and nearly global, abruptly narrowed and extended into a moderately long duct near middle of ventral side, the duct distinctly narrowed apically; diverticulum and spermathecal duct arising from the end of the duct of vagina; diverticulum short, spiral and thin; spermathecal duct nearly as long as and slightly thicker than diverticulum; spermatheca composed of a spiral tube which is distinctly longer than diverticulum, basal portion extended into a short tube, where the accessory gland opening; accessory gland thin and very long, much longer than the spiral tube of spermatheca; median oviduct situated at base of vagina.

Remarks
In the present study, the abdominal sternite VIII and genitalia of female are photographed and described for the first time.

Distribution
Taiwan.
Abdominal sternite VIII of female (Fig. 5H).Distinctly protuberant in middle of posterior margin, the middle protuberance moderately and triangularly emarginate at apex, forming a pair of median lobes, which are widely subrounded at apices.
Internal organ of reproductive system (Fig. 8A).Vagina stout, abruptly thinned and extended into a slightly long duct at ventroapical portion; diverticulum and spermathecal duct arising from the end of the duct of vagina; diverticulum moderately long, spiral and distinctly thinned apically; spermathecal duct distinctly shorter than diverticulum and thinner than basal portion of diverticulum; spermatheca composed of a spiral tube which is distinctly longer than diverticulum, provided with a thin accessory gland which is slightly shorter than the spiral tube of spermatheca; median oviduct situated in middle of vagina.

Remarks
In the original publication (Wittmer 1982), the aedeagus was illustrated in dorsal view, here it is illustrated in ventral, dorsal and lateral views and described in detail.Its abdominal sternite VIII and genitalia of female are illustrated and described for the first time.

Distribution
Taiwan.

Supplementary description
Aedeagus (Fig. 6D-F).Ventral process of each paramere wide, slightly narrowed apically, moderately hooked at apex; conjoint dorsal plate of parameres moderately developed, about half length of ventral processes and as wide as half of the distance between outer margins of ventral processes, distinctly narrowed apically, with straight apical margin.
Abdominal sternite VIII of female (Fig. 11A).Distinctly protuberant in middle of posterior margin, the middle protuberance moderately and triangularly emarginate at apex, forming a pair of median lobes, which are widely triangular at apices.
Internal organ of reproductive system (Fig. 8B).Vagina stout, abruptly thinned and extended into a slightly long duct at ventroapical portion; diverticulum and spermathecal duct arising from the end of the duct of vagina; diverticulum moderately long, spiral and distinctly thinned apically; spermathecal duct distinctly shorter than diverticulum and thinner than basal portion of diverticulum; spermatheca composed of a spiral tube which is slightly longer than diverticulum, provided with a thin accessory gland which is slightly shorter than the spiral tube of spermatheca; median oviduct situated in middle of vagina.

Remarks
In the original publication (Wittmer 1995), this species was only illustrated with male antennae.Here the genitalia of both sexes and abdominal sternite VIII of female are illustrated and described for the first time.

Distribution
Taiwan.

Diagnosis
This species is similar to F. formosana (Pic, 1910), but can be differentiated from the latter by the black head, eyes strongly protruding, pronotum distinctly widened posteriorly, aedeagus: conjoint dorsal plate of parameres with round apical margin.It also resembles F. minor (Wittmer, 1997) in the body coloration, but differs in the following characters: eyes strongly protruding, pronotum slightly longer than wide, aedeagus: conjoint dorsal plate of parameres about half length of ventral process, while in the latter species, eyes moderately protruding, pronotum nearly as long as wide, aedeagus: conjoint dorsal plate of parameres as long as ventral process.

Etymology
The specific epithet is derived from the Latin niger (black) and ceps (head), referring to its black head.
Body.Black, prothorax and legs orange, tarsi black.Body densely covered with recumbent gray pubescence.
Head.Subquadrate, temples distinctly narrowed posteriorly, surface densely and finely punctate; eyes strongly protruding, head width across eyes distinctly wider than anterior margin of pronotum; terminal maxillary palpomeres subtriangular, widest nearly in middle, acurate on apical parts of inner margins, acute at apices; antennae filiform and simple, extending to basal two-length of elytra, antennomeres II slightly longer than wide, III about twice as long as II, IV slightly longer than III, XI slightly longer than X, pointed at apices.Pronotum.Subquadrate, slightly longer than wide, distinctly widened posteriorly, anterior margin rounded, lateral margins slightly sinuate, posterior margin narrowly bordered and arcuate, anterior angles rounded, posterior angles nearly rectangular; disc distinctly convex on posterolateral parts, surface semilustrous, sparsely and finely punctate.
Legs.With all tarsal claws bifid, upper claws nearly as long as lower ones.
Aedeagus (Fig. 6G-I).Ventral process of each paramere wide, slightly narrowed apically, moderately hooked at apex; conjoint dorsal plate of parameres moderately developed, about half length of ventral process and as wide as half of the distance between outer margins of ventral processes, distinctly narrowed apically, with round apical margin.

Female
Unknown.

Supplementary description
Aedeagus (Fig. 7A-C).Ventral process of each paramere wide, distinctly narrowed apically, moderately hooked at apex; conjoint dorsal plate of parameres slightly developed, about one-fourth length of ventral process and as wide as half of the distance between outer margins of ventral processes, moderately narrowed apically, with slightly rounded apical margin.
Abdominal sternite VIII of female (Fig. 11B).Moderately protuberant in middle of posterior margin, the middle protuberance moderately and widely triangularly emarginate at apex, forming a pair of median lobes, which are subtriangular at apices.
Internal organ of reproductive system (Fig. 8C).Vagina stout, slightly thinned and extended into a slightly long duct at ventroapical portion; diverticulum and spermathecal duct arising from the end of the duct of vagina; diverticulum moderately long, spiral and abruptly narrowed apically; spermathecal duct distinctly shorter than diverticulum and thinner than basal portion of diverticulum; spermatheca composed of a spiral tube which is slightly longer than diverticulum, provided with a thin accessory gland which is slightly shorter than the spiral tube of spermatheca; median oviduct situated in middle of vagina.

Distribution
Taiwan.

Supplementary description
Aedeagus (Fig. 7D-F).Ventral process of each paramere wide, slightly narrowed apically, moderately hooked at apex; conjoint dorsal plate of parameres strongly reduced, slightly triangularly protuberant in middle of apical margin.
Abdominal sternite VIII of female (Fig. 11C).Distinctly and triangularly emarginate in middle of posterior margin.
Internal organ of reproductive system (Fig. 8D).Vagina stout, abruptly thinned and extended into a slightly long duct at ventroapical portion; diverticulum and spermathecal duct arising from the end of the duct of vagina; diverticulum moderately long, spiral and moderately thinned apically; spermathecal duct slightly shorter than diverticulum and thinner than basal portion of diverticulum; spermatheca composed of a spiral tube which is slightly longer than diverticulum, provided with a thin accessory gland which is nearly as long as the spiral tube of spermatheca; median oviduct situated in middle of vagina.

Remarks
In the original publication (Wittmer 1982), the aedeagus of species was illustrated in ventral and dorsal views, here it is illustrated again and also in lateral view.Here the genitalia and abdominal sternite VIII of female are illustrated and described for the first time.

Distribution
Taiwan.
composed of a spiral tube which is nearly as long as diverticulum, provided with a thin accessory gland, which is distinctly shorter than the spiral tube of spermatheca; median oviduct situated in middle of vagina.

Distribution
Taiwan.

Supplementary description
Aedeagus (Fig. 9D-F).Ventral process of each paramere wide, slightly narrowed apically, moderately hooked at apex; conjoint dorsal plate of parameres moderately developed, about half length of ventral process and as wide as half of the distance between outer margins of ventral processes, slightly narrowed apically, with nearly straight apical margin.
Abdominal sternite VIII of female (Fig. 11F).Moderately protuberant middle of posterior margin, the middle protuberance moderately and triangularly emarginate at apex, forming a pair of median lobes, which are rounded at apices.
Internal organ of reproductive system (Fig. 10C).Vagina stout, abruptly thinned and extended into a slightly long duct at ventroapical portion; diverticulum and spermathecal duct arising from the end of the duct of vagina; diverticulum slightly long and spiral, moderately thinned apically; spermathecal duct distinctly shorter than diverticulum and slightly thinner than basal portion of diverticulum; spermatheca composed of a spiral tube which is slightly longer than diverticulum, provided with a thin accessory gland which is nearly as long as the spiral tube of spermatheca; median oviduct situated in middle of vagina.

Remarks
In the original publication (Wittmer 1982), the aedeagus of this species was illustrated only in dorsal view, here it is illustrated again and also in ventral and lateral views.Here the genitalia and abdominal sternite VIII of female are illustrated and described for the first time.

Distribution
Taiwan.

Diagnosis
This genus is very similar to Fissocantharis, but differentiated from the latter by the aedeagus presenting with laterophyses.The structures of female genitalia are variable, most species are similar to those of Fissocantharis, but some with vagina abruptly and strongly expanded at apical part, spermathecal duct absent, median oviduct situated at base of vagina, such as Micropodabrus multicostata (Wittmer, 1982) and M. nodicornis (Wittmer, 1982).

Supplementary description
Aedeagus (Fig. 13A-C).Ventral process of each paramere wide, abruptly narrowed at apical portion and pointed at apex; conjoint dorsal plate of parameres well-developed, slightly longer than the length of ventral process and as wide as nearly half of the distance between outer margins of ventral processes, nearly parallel-sided, with apical margin deeply emarginate in middle, the deepness of the middle emargination about half length of conjoint dorsal plate; laterophyses thin, straight and pointed at apices, separated but leaning against each other and sin middle of dorsal side of conjoint dorsal plate.
Abdominal sternite VIII of female (Fig. 16A).Distinctly protuberant in middle of posterior margin, the middle protuberance widely rounded at apex.
Internal organ of reproductive system (Fig. 15A).Vagina stout, abruptly thinned and extended into a long duct at apex; diverticulum and spermathecal duct arising from the end of the duct of vagina; diverticulum very long, thin and spiral; spermathecal duct distinctly shorter and thicker than diverticulum; spermatheca composed of a spiral tube which is nearly as long as diverticulum, provided with a thin accessory gland, which is much shorter than the spiral tube of spermatheca; median oviduct situated at apex of vagina.

Remarks
In the original publication (Wittmer 1972), this species was only illustrated with antenna of the male.
Here the genitalia of both sexes and abdominal sternite VIII of female are illustrated and described for the first time.

Distribution
Taiwan.

Remarks
In the original publication (Wittmer 1982), the aedeagus of species was illustrated in ventral and dorsal views, here it is illustrated again and also in lateral view.Here the genitalia and abdominal sternite VIII of female are illustrated and described for the first time.

Distribution
Taiwan.

Supplementary description
Aedeagus (Fig. 14D-F).Ventral process of each paramere distinctly narrowed apically, slightly hooked at apex; conjoint dorsal plate of parameres well-developed, distinctly longer than the length of ventral process and wider than the distance between outer margins of ventral processes, slightly narrowed apically, with nearly straight apical margin; laterophyses separated, thin, nearly straight and pointed at apices, situated in middle of dorsal side of median lobe, distinctly shorter than conjoint dorsal plate.
Abdominal sternite VIII of female (Fig. 16D).Distinctly and widely emarginate in middle of posterior margin, the middle emargination moderately and triangularly protuberant in middle of bottom margin, lateroapical angles narrowly rounded.
Internal organ of reproductive system (Fig. 15D).Vagina stout, abruptly thinned and extended into a short and thick duct at apex; diverticulum and spermathecal duct arising from the end of the duct of vagina; diverticulum moderately long, spiral and abruptly thinned apically; spermathecal duct very short, distinctly thinner than basal portion of diverticulum; spermatheca composed of a spiral tube, which is slightly shorter than diverticulum, provided with a thin accessory gland, which is distinctly shorter than the spiral tube of spermatheca; median oviduct situated at apex of vagina.

Remarks
The aedeagus was illustrated in ventral view by Wittmer (1982), here it is illustrated again and also in dorsal and lateral views.Besides, the genitalia and abdominal sternite VIII of female are illustrated and described for the first time.

Distribution
Taiwan.

Diagnosis
This species could be easily differentiated from other species of Micropodabrus by its brown elytra and the structures of aedaegus: ventral process of each paramere wide, laterophyses strongly sclerotized and bent dorsally at apices.

Etymology
The specific epithet is derived from the Latin brunneus (brown) and penna (wing), referring to its brown elytra.
Body.Black and shiny, elytra brown.Body densely covered with recumbent gray pubescence, pronotum and elytra with brown pubescence, which is slightly longer on the latter.
Head.Rounded, temples slightly narrowed posteriorly, surface densely and coarsely punctate; eyes moderately protruding, head width across eyes slightly wider than anterior margin of pronotum; terminal maxillary palpomeres subtriangular, widest nearly in middle, acurate on apical parts of inner margins, acute at apices; antennae filiform, slightly thickened, extending to elytral midlength, antennomeres II nearly as long as wide, III about twice as long as II, IV about one-third longer than III, XI slightly longer than X, pointed at apices.Pronotum.Subquadrate, slightly wider than long, slightly widened posteriorly, anterior margin arcuate, lateral margins sinuate, posterior margin narrowly bordered and bisinuate, anterior angles rounded, posterior angles nearly rectangular; disc distinctly convex on posterolateral parts, surface lustrous, sparsely and finely punctate.
Elytra.About 3.0 times as long as humeral width, 4.5 times longer than pronotum, nearly parallelsided, disc densely and coarsely punctate.
Legs.With all tarsal claws bifid.
Aedeagus (Fig. 14G-I).Ventral process of each paramere wide, slightly sinuate at outer margins, moderately hooked at apex; conjoint dorsal plate of parameres well-developed, slightly shorter than the length of ventral processes and distinctly narrower than distance between outer margins of ventral processes, slightly sinuate in middle of lateral margins, with nearly straight apical margin; laterophyses separated, strongly sclerotized and bent dorsally at apices, situated in middle of dorsal side of median lobe, as long as conjoint dorsal plate.

Female
Unknown.

Distribution
Taiwan.

Discussion
Apart from the characters provided in traditional description and comparisons, Fissocantharis laticornis differs from other species of Fissocantharis by the female genitalia: vagina is nearly global, abruptly narrowed and extended into a moderately long duct near middle of ventral side; median oviduct is situated at base of vagina.Other species of this genus share the characters summarized in the generic diagnosis, but can be easily differentiated by the shapes of antennae in males, see the keys by Wittmer (1982Wittmer ( , 1995)).
The species of Micropodabrus can be distinguished using the key provided by Yang et al. (2009).The shapes of laterophyses of the aedeagus are variable among the species within this genus.This variation can be also observed well in the genus Themus Motschulsky, 1860.Accordingly, the female genitalia are also variable, particularly the shape of the vagina and the position of median oviduct.