taxonID	type	description	language	source
03AD2179AE59FFC4FF78FB36FD3BF875.taxon	materials_examined	Type species. Fontidessus toboganensis Miller, new species by current designation.	en	Miller, Kelly B., Spangler, Paul J. (2008): Fontidessus Miller and Spangler, a new genus of Bidessini from Venezuela (Coleoptera: Dytiscidae: Hydroporinae) with three new species. Zootaxa 1827: 45-52, DOI: 10.5281/zenodo.183126
03AD2179AE59FFC4FF78FB36FD3BF875.taxon	diagnosis	Diagnosis and description. Fontidessus is clearly a member of the tribe Bidessini based on the bi-segmented lateral lobes of the aedeagus (Figs 10, 13, 16) (Biström, 1988) and the presence of a distinct spermathecal spine (Fig. 17) (Miller, 2001). Fontidessus differs from other genera in the tribe by the combination of: 1) transverse occipital line absent (Figs 1 – 3), 2) basal pronotal striae present (Figs 1 – 3), 3) basal elytral stria absent (Figs 1 – 3), 4) elytral sutural stria faintly present in some specimens (e. g. Fig. 1), 5) anterior clypeal margin unmodified, 6) elytron without longitudinal carinae, 7) epipleuron without transverse carina at humeral angle, 8) lateral lobes of aedeagus two-segmented (Figs 10, 13, 16), 9) habitus elongate, oval, with lateral pronotal and elytral margins nearly continuously and shallowly curved (Figs 1 – 3), and 10) metatrochanter extremely large relative to metafemur, approximately 0.6 × length of metafemur (Fig. 4). This genus is most similar to the genera Uvarus Guignot and Bidessodes Régimbart (to which it keys in Biström’s 1988 key to bidessine genera). From Bidessodes it differs by 1) dorsum not iridescent, and 2) metatibiae without natatory setae and metatarsi with few natatory setae. Bidessodes appears to be a monophyletic group. Although some of its members have dramatic modifications to the male legs or genitalia, they are superficially quite similar and have similar iridescence and color pattern on the dorsal surface. Fontidessus specimens lack these features. Despite the absence of a cladistic phylogenetic analysis, Fontidessus appears to be clearly outside Bidessodes. From Uvarus, Fontidessus genus differs in lacking basal elytra striae and natatory setae. It is, however, quite similar to Uvarus genus in most characters and general features such as shape, size and coloration. The problem with these observations is that Uvarus is a heterogeneous group with a worldwide distribution. Within Bidessini it is the “ trashcan ” group that includes those taxa without an occipital line but also without any unifying synapomorphy. Placing these new species in Uvarus would require an expansion of an already problematic group to include a new character combination. It seems more desirable to place these species in a new genus until the classification of the Bidessini can be more thoroughly tested (see Discussion below).	en	Miller, Kelly B., Spangler, Paul J. (2008): Fontidessus Miller and Spangler, a new genus of Bidessini from Venezuela (Coleoptera: Dytiscidae: Hydroporinae) with three new species. Zootaxa 1827: 45-52, DOI: 10.5281/zenodo.183126
03AD2179AE59FFC4FF78FB36FD3BF875.taxon	etymology	Etymology. This genus is named Fontidessus from the Latin word fontis, meaning “ spring, ” referring to the seepage-like habitat of the type locality of the type species, and dessus, a common root for genus names in the tribe Bidessini	en	Miller, Kelly B., Spangler, Paul J. (2008): Fontidessus Miller and Spangler, a new genus of Bidessini from Venezuela (Coleoptera: Dytiscidae: Hydroporinae) with three new species. Zootaxa 1827: 45-52, DOI: 10.5281/zenodo.183126
03AD2179AE59FFC4FF78FB36FD3BF875.taxon	distribution	Distribution and habitat. The genus is known from three sympatric species from only a single site in Venezuela (see type localities below). In each case, however, the species do not appear to occupy typical open water. Instead they appear to occur in seeps or shallow water at stream margins.	en	Miller, Kelly B., Spangler, Paul J. (2008): Fontidessus Miller and Spangler, a new genus of Bidessini from Venezuela (Coleoptera: Dytiscidae: Hydroporinae) with three new species. Zootaxa 1827: 45-52, DOI: 10.5281/zenodo.183126
03AD2179AE59FFC4FF78FB36FD3BF875.taxon	discussion	Discussion. The genera of Bidessini are currently defined based on the combination of several characters such as the presence or absent of an occipital line, basal sutural and pronotal striae, epipleural carinae, elytra carinae and more subjective characters such as general shape. Because of the nature of the various combinations it is clear that homoplasy, or loss, is exhibited in many of the characters throughout the tribe. There has been only limited cladistic testing of these characters (e. g. Miller et al., 2006). When new species are discovered with a new combination of characters, such as those described here, it is not easy to decide whether to erect a new genus or expand the definition of existing genera (which may require synonymy with other genera). It seems best, in the absence of cladistic testing of relationships and putative homologies, to place anomalous species in new genera to draw attention to their unique character combination until such time as the genera can be phylogenetically revised. In further support of the placement of this taxon outside both Uvarus and Bidessodes, a recent molecular phylogenetic analysis of Dytiscidae by Ribera et al. (2008) included at least one member of this new genus. According to M. Balke (personal communication), “ unknown Bidessini a ” in that analysis (MB 1171, Ribera et al., 2008) is the species described here as F. toboganensis. In their analysis F. toboganensis is resolved with two other unidentified Bidessini and well removed from either Uvarus or Bidessodes (Ribera et al., 2008).	en	Miller, Kelly B., Spangler, Paul J. (2008): Fontidessus Miller and Spangler, a new genus of Bidessini from Venezuela (Coleoptera: Dytiscidae: Hydroporinae) with three new species. Zootaxa 1827: 45-52, DOI: 10.5281/zenodo.183126
03AD2179AE5BFFC2FF78FD19FD68F8AC.taxon	materials_examined	Type locality. Venezuela, Territorio Federal Amazonas, 40 km S Puerto Ayacucho, at El Tobogán, Coromoto.	en	Miller, Kelly B., Spangler, Paul J. (2008): Fontidessus Miller and Spangler, a new genus of Bidessini from Venezuela (Coleoptera: Dytiscidae: Hydroporinae) with three new species. Zootaxa 1827: 45-52, DOI: 10.5281/zenodo.183126
03AD2179AE5BFFC2FF78FD19FD68F8AC.taxon	diagnosis	Diagnosis. This is the largest known species in the genus (TL> 1.4 mm). The prosternal process is moderately broad and parallel-sided and has the apex broadly truncate (Fig. 5). The male genitalia are distinctive with the median lobe elongate, slender and broadly recurved in lateral aspect (Fig. 9). There is an elongate ventral sclerite that fits into the ventral groove of the median lobe (Fig. 9).	en	Miller, Kelly B., Spangler, Paul J. (2008): Fontidessus Miller and Spangler, a new genus of Bidessini from Venezuela (Coleoptera: Dytiscidae: Hydroporinae) with three new species. Zootaxa 1827: 45-52, DOI: 10.5281/zenodo.183126
03AD2179AE5BFFC2FF78FD19FD68F8AC.taxon	description	Description. Measurements. TL = 1.5 – 1.6 mm, GW = 0.8 – 0.9 mm, PW = 0.6 – 0.7 mm, HW = 0.4 – 0.5 mm, EW = 0.3 – 0.4 mm, TL / GW = 1.8 – 1.7, HW / EW = 1.6 – 1.7. Body (Fig. 1) oval, elongate; lateral outline nearly continuous between pronotum and elytron; lateral margins of pronotum gently curved; lateral margins of elytron evenly and gently curved. Coloration. Head yellow brown, pronotum yellow brown, brown medially along posterior margin (Fig. 1); elytron brown with the following yellow maculae: 1) a basal macula in a band extending from lateral to sutural margins, posterior margin of macula irregular, 2) a lateral longitudinal macula at humeral angle, 3) a narrow longitudinal macula along lateral margin, and 4) a subtriangular subapical macula (Fig. 1). Ventral surfaces of thorax and abdomen brown except prosternum, prosternal process, propleuron and pronotal epipleuron yellow-brown; appendages yellow to yellow-brown. Sculpture and structure. Head with very fine, inconspicuous, irregular punctation, surface between punctures shiny with indistinct microsculpture in the form of small cells; eyes medium in size (Fig. 1, HW / EW = 1.6 – 1.7). Pronotal surface similar to that of head; with posterior angles obtuse; lateral bead narrow, of even width throughout; pronotal striae finely incised, extending nearly 1 / 2 distance across pronotum (Fig. 1). Elytron with anterolateral angle obtuse, not extended anteriorly (Fig. 1); surface similar to pronotum. Prosternal process broad, lateral margins subparallel, apex of process broadly rounded (Fig. 5); metacoxal process with lateral lobe minute. Pro- and mesotarsi relatively narrow in both male and female, but slightly broader in male. Male genitalia. Median lobe in ventral aspect extremely slender and long, apex slender and sharply pointed (Fig. 8); in lateral aspect narrow, extremely slender and long in apical half, abruptly curved medially to sharply pointed, recurved apex (Fig. 9); with elongate, slender dorsal sclerite (Fig. 9). Lateral lobe in lateral aspect broad basally, bent medially with apical segment elongate and parallel sided to narrowly rounded apex (Fig 10). Female genitalia (Fig. 17). Bursa copulatrix elongate and narrow; spermathecal duct long, slender, somewhat coiled, expanded prior to receptacle; receptacle small, about half size of spermatheca, intermediate duct between receptacle and spermatheca broad; spermatheca globular, spermathecal spine prominent, but not elongate; fertilization duct elongate, tightly curved in spiral, attaching to large fertilization sac; vagina elongate, moderately narrow; gonocoxae elongate, slender with abrupt, small, triangular expansion at apex (Fig. 17).	en	Miller, Kelly B., Spangler, Paul J. (2008): Fontidessus Miller and Spangler, a new genus of Bidessini from Venezuela (Coleoptera: Dytiscidae: Hydroporinae) with three new species. Zootaxa 1827: 45-52, DOI: 10.5281/zenodo.183126
03AD2179AE5BFFC2FF78FD19FD68F8AC.taxon	etymology	Etymology. The species name is derived from the name of the type locality, El Tobogán.	en	Miller, Kelly B., Spangler, Paul J. (2008): Fontidessus Miller and Spangler, a new genus of Bidessini from Venezuela (Coleoptera: Dytiscidae: Hydroporinae) with three new species. Zootaxa 1827: 45-52, DOI: 10.5281/zenodo.183126
03AD2179AE5BFFC2FF78FD19FD68F8AC.taxon	distribution	Distribution and habitat. According to notes discovered in P. J. Spangler’s office: “ The series of [Fontidessus toboganensis] was collected from seepage areas alongside a cascade. The specimens were obtained by pulling matted plant roots from granitic bedrock and using an aspirator to suck up the water that accumulated in rock depressions as runoff from the roots. The dirty concentrated residue was then filtered through fine-meshed, nylon cloth and the very small adults and larvae were again aspirated as they crawled out of the residue. ” Label data indicate the collection of specimens of the species from the following habitats: “ seep, ” “ sunlit stream ”, “ bare rock streambed in sunlight; beneath wet leaves at margin, ” “ sunlit stream; slow water over sand and gravel, ” and “ leaves from stream. ”	en	Miller, Kelly B., Spangler, Paul J. (2008): Fontidessus Miller and Spangler, a new genus of Bidessini from Venezuela (Coleoptera: Dytiscidae: Hydroporinae) with three new species. Zootaxa 1827: 45-52, DOI: 10.5281/zenodo.183126
03AD2179AE5BFFC2FF78FD19FD68F8AC.taxon	materials_examined	Material examined. HOLOTYPE in USNM: male labeled, “ VENEZUELA: T. F. Amazonas Puerto Ayacucho (40 km. S) El Tobogán, Cano Coromoto 26 January 1989 / seep at upper shelter collected by PJSPangler RAFaitoute & CBBarr / HOLOTYPE: Fontidessus toboganensis Miller and Spangler, 2008 [red label with double black line border]. ” PARATYPES, 250 total, all with same locality data as holotype, 11 with same collecting date as holotype; 1 with 22 February 1986; 21 with 25 February 1986; 24 with 21 January 1985; 1 with 16 November 1987; 192 with 23 January 1989.	en	Miller, Kelly B., Spangler, Paul J. (2008): Fontidessus Miller and Spangler, a new genus of Bidessini from Venezuela (Coleoptera: Dytiscidae: Hydroporinae) with three new species. Zootaxa 1827: 45-52, DOI: 10.5281/zenodo.183126
03AD2179AE5DFFC3FF78FF4CFC9FF8C6.taxon	materials_examined	Type locality. Venezuela, Territorio Federal Amazonas, 40 km S Puerto Ayacucho, at El Tobogán, Coromoto.	en	Miller, Kelly B., Spangler, Paul J. (2008): Fontidessus Miller and Spangler, a new genus of Bidessini from Venezuela (Coleoptera: Dytiscidae: Hydroporinae) with three new species. Zootaxa 1827: 45-52, DOI: 10.5281/zenodo.183126
03AD2179AE5DFFC3FF78FF4CFC9FF8C6.taxon	diagnosis	Diagnosis. This species is smaller than F. toboganensis (TL <1.3 mm), but is similar in size (though slightly smaller) and external appearance to F. wheeleri. Fontidessus ornatus has the prosternal process moderately elongate with the lateral margins in the apical portion straight and evenly tapered to the pointed apex (Fig. 6). The pale regions on the elytron are diffuse and poorly demarcated and are yellow-brown unlike F. wheeleri which have the elytral maculae brighter yellow and more clearly demarcated. The male genitalia are diagnostic with the median lobe slender and elongate with a subapical brush of setae on the ventral margin (Fig. 12).	en	Miller, Kelly B., Spangler, Paul J. (2008): Fontidessus Miller and Spangler, a new genus of Bidessini from Venezuela (Coleoptera: Dytiscidae: Hydroporinae) with three new species. Zootaxa 1827: 45-52, DOI: 10.5281/zenodo.183126
03AD2179AE5DFFC3FF78FF4CFC9FF8C6.taxon	description	Description. Measurements. TL = 1.1 – 1.2 mm, GW = 0.6 – 0.7 mm, PW = 0.6 – 0.7 mm, HW = 0.4 – 0.5 mm, EW = 0.1 – 0.2 mm, TL / GW = 1.8 – 1.9, HW / EW = 1.5 – 1.6. Body (Fig. 2) oval, elongate; lateral outline slightly continuous between pronotum and elytron; lateral margins of pronotum gently curved; lateral margins of elytron evenly and gently curved. Coloration. Head yellow brown, darker in region near medial margins of eyes; pronotum yellow brown, diffusely brown medially along posterior margin and in narrow band along anterior margin; elytron brown with diffuse yellow pattern, yellow basally, laterally along about ¾ length of elytra and in subtriangular subapical area (Fig. 2). Ventral surfaces of thorax and abdomen brown except prosternum, prosternal process, propleuron and pronotal epipleuron yellow-brown; appendages yellow to yellow-brown. Sculpture and structure. Head with very fine, inconspicuous, irregular punctation, surface between punctures shiny with indistinct microsculpture in the form of small cells; eyes medium in size (Fig. 2, HW / EW = 1.5 – 1.6). Pronotal surface similar to that of head; with posterior angles obtuse; lateral bead slender, of even width throughout; pronotal striae finely incised, extending nearly 1 / 2 distance across pronotum (Fig. 2). Elytron with anterolateral angle obtuse, not extended anteriorly (Fig. 2); surface similar to pronotum but with microsculpture more indistinct. Prosternal process relatively narrow, lateral margins tapered to narrowly rounded apex (Fig. 6); metacoxal process with lateral lobe minute. Pro- and mesotarsi relatively narrow in both male and female. Male genitalia. Median lobe in ventral aspect slender, elongate and with lateral margins parallel to pointed apex (Fig. 11); in lateral aspect moderately broad at base, curved, with apical portion evenly tapered to pointed apex, subapically with small brush of setae (Fig. 12). Lateral lobe elongate, expanded apically, broadly rounded at apex, with series of long setae along dorsal margin (Fig 13).	en	Miller, Kelly B., Spangler, Paul J. (2008): Fontidessus Miller and Spangler, a new genus of Bidessini from Venezuela (Coleoptera: Dytiscidae: Hydroporinae) with three new species. Zootaxa 1827: 45-52, DOI: 10.5281/zenodo.183126
03AD2179AE5DFFC3FF78FF4CFC9FF8C6.taxon	etymology	Etymology. The species name is derived from the Greek root orno, meaning “ adorned, ” for the attractive coloration characterizing this species.	en	Miller, Kelly B., Spangler, Paul J. (2008): Fontidessus Miller and Spangler, a new genus of Bidessini from Venezuela (Coleoptera: Dytiscidae: Hydroporinae) with three new species. Zootaxa 1827: 45-52, DOI: 10.5281/zenodo.183126
03AD2179AE5DFFC3FF78FF4CFC9FF8C6.taxon	distribution	Distribution and habitat. The species is known only from the type locality. Label data indicated the following habitats; “ seep, ” and “ bare rock streambed. ”	en	Miller, Kelly B., Spangler, Paul J. (2008): Fontidessus Miller and Spangler, a new genus of Bidessini from Venezuela (Coleoptera: Dytiscidae: Hydroporinae) with three new species. Zootaxa 1827: 45-52, DOI: 10.5281/zenodo.183126
03AD2179AE5DFFC3FF78FF4CFC9FF8C6.taxon	materials_examined	Material examined. HOLOTYPE in USNM: male labeled, “ VENEZUELA: T. F. Amazonas Puerto Ayacucho (40 km. S) at Tobogán, 22 Feb 1986 P. J. Spangler, Colln. # 6 / Bare rock streambed in sunlight; beneath wet leaves at margin / HOLOTYPE: Fontidessus ornatus Miller, 2008 [red label with double black line bor- der]. ” PARATYPES, 14 total; 3 with same label data as holotype; 9, Venezuela, T. F. Amazonas, Puerto Ayacucho, 40 km S El Tobogán, Caño Coromoto, 23 January 1989, seep at upper shelter, P. J. Spangler, R. A. Faitoute, and C. B. Barr; 2, same except 26 January 1989.	en	Miller, Kelly B., Spangler, Paul J. (2008): Fontidessus Miller and Spangler, a new genus of Bidessini from Venezuela (Coleoptera: Dytiscidae: Hydroporinae) with three new species. Zootaxa 1827: 45-52, DOI: 10.5281/zenodo.183126
03AD2179AE5DFFC0FF78F8BCFE46F96B.taxon	materials_examined	Type locality. Venezuela, Territorio Federal Amazonas, 40 km S Puerto Ayacucho, at El Tobogán, Coromoto.	en	Miller, Kelly B., Spangler, Paul J. (2008): Fontidessus Miller and Spangler, a new genus of Bidessini from Venezuela (Coleoptera: Dytiscidae: Hydroporinae) with three new species. Zootaxa 1827: 45-52, DOI: 10.5281/zenodo.183126
03AD2179AE5DFFC0FF78F8BCFE46F96B.taxon	diagnosis	Diagnosis. This species is smaller than F. toboganensis (TL <1.4 mm), but is similar in size (though slightly larger) and external appearance to F. o r n a t u s. Fontidessus wheeleri has the prosternal process elongate with the lateral margins slightly concave and tapered to sharply pointed apex (Fig. 7). The pale regions on the elytron are brighter yellow and more clearly demarcated in F. w h e e l e r i than in F. ornatus which have the elytral maculae more pale and diffuse. The male genitalia are diagnostic with the median lobe broad and deeply bifid in ventral aspect (Fig. 14).	en	Miller, Kelly B., Spangler, Paul J. (2008): Fontidessus Miller and Spangler, a new genus of Bidessini from Venezuela (Coleoptera: Dytiscidae: Hydroporinae) with three new species. Zootaxa 1827: 45-52, DOI: 10.5281/zenodo.183126
03AD2179AE5DFFC0FF78F8BCFE46F96B.taxon	description	Description. Measurements. TL = 1.2 – 1.3 mm, GW = 0.6 – 0.7 mm, PW = 0.5 – 0.6 mm, HW = 0.4 – 0.5 mm, EW = 0.2 – 0.3 mm, TL / GW = 1.7 – 1.8, HW / EW = 1.6 – 1.7. Body (Fig. 3) oval, elongate; lateral outline slightly continuous between pronotum and elytron; lateral margins of pronotum gently curved; lateral margins of elytron evenly and gently curved. Coloration. Head yellow brown, brown along posterior margins and medial margins of eyes; pronotum broadly brown along posterior margin, narrowly dark brown along anterior margin; elytron brown with the following yellow maculae: 1) a basal macula in an irregular band extending posteriorly along suture at medial margin, 2) a lateral longitudinal macula posterior to humeral angle, broader anteriorly and extending posteriorly in narrow band to near apex, 3) a subtriangular subapical macula (Fig. 3). Ventral surfaces of thorax and abdomen dark brown except prosternum, prosternal process, propleuron and pronotal epipleuron yellowbrown; appendages yellow to yellow-brown. Sculpture and structure. Head with very fine, inconspicuous, irregular punctation, surface between punctures shiny, without distinct microsculpture; eyes medium in size (Fig. 3, HW / EW = 1.6 – 1.7). Pronotal surface similar to that of head, with posterior surface with fine microsculpture; with posterior angles obtuse; lateral bead slender, of even width throughout; pronotal striae finely incised, extending nearly 1 / 2 distance across pronotum (Fig. 3). Elytron with anterolateral angle obtuse, not extended anteriorly (Fig. 3); surface similar to pronotum, with microsculpture indistinct. Prosternal process elongate and slender, lateral margins slightly concave, apex of process rounded (Fig. 7); metacoxal process with lateral lobe small. Pro- and mesotarsi relatively narrow in both male and female, but slightly broader in male. Male genitalia. Median lobe in ventral aspect very broad, deeply bifid, with branches broad, subapically strongly constricted on lateral margins, apices sharply pointed (Fig. 14); in lateral broad at base, irregularly narrowed to sharply pointed apex (Fig. 15). Lateral lobe narrow at base, medially expanded, apical portion parallel-sided to slightly expanded, broadly rounded apex, with small series of elongate setae along dorsal margin (Fig 16).	en	Miller, Kelly B., Spangler, Paul J. (2008): Fontidessus Miller and Spangler, a new genus of Bidessini from Venezuela (Coleoptera: Dytiscidae: Hydroporinae) with three new species. Zootaxa 1827: 45-52, DOI: 10.5281/zenodo.183126
03AD2179AE5DFFC0FF78F8BCFE46F96B.taxon	etymology	Etymology. This species is named in honor of my good friend and fellow coleopterist, Dr. Quentin D. Wheeler, Arizona State University.	en	Miller, Kelly B., Spangler, Paul J. (2008): Fontidessus Miller and Spangler, a new genus of Bidessini from Venezuela (Coleoptera: Dytiscidae: Hydroporinae) with three new species. Zootaxa 1827: 45-52, DOI: 10.5281/zenodo.183126
03AD2179AE5DFFC0FF78F8BCFE46F96B.taxon	distribution	Distribution and habitat. The species is known only from the type locality. Label data indicate the following habitats; “ seep, ” and “ bare rock streambed. ”	en	Miller, Kelly B., Spangler, Paul J. (2008): Fontidessus Miller and Spangler, a new genus of Bidessini from Venezuela (Coleoptera: Dytiscidae: Hydroporinae) with three new species. Zootaxa 1827: 45-52, DOI: 10.5281/zenodo.183126
03AD2179AE5DFFC0FF78F8BCFE46F96B.taxon	materials_examined	Material examined. HOLOTYPE in USNM: male labeled, “ VENEZUELA: T. F. Amazonas Puerto Ayacucho (40 km. S) at Tobogán, 22 Feb 1986 P. J. Spangler, Colln. # 6 / Bare rock streambed in sunlight; beneath wet leaves at margin / HOLOTYPE: Fontidessus wheeleri Miller, 2008 [red label with double black line bor- der]. ” PARATYPES, 1, same as holotype except 23 January 1989, seep at upper shelter, P. J. Spangler, R. A. Faitoute and C. B. Barr, colrs.	en	Miller, Kelly B., Spangler, Paul J. (2008): Fontidessus Miller and Spangler, a new genus of Bidessini from Venezuela (Coleoptera: Dytiscidae: Hydroporinae) with three new species. Zootaxa 1827: 45-52, DOI: 10.5281/zenodo.183126
