taxonID	type	description	language	source
081787E4FF8BFF82FF4EFF53FCAB58B0.taxon	diagnosis	Diagnosis. Length 3 – 4 mm; elongate; legs stout; head recumbent; eyes small, distance from subgenal ridge to bottom of eye a little more than diameter of eye, subgenal ridge without buttonhole; antenna without enlarged scape, antennomeres 2 & 3 not reduced; labial palpi straight (sensu Watts & Zwick (2019 )); mandible with one strong tooth, molar without small spines; pronotum transverse, anterolateral angles rounded; pronotal process broad at apex, corresponding notch in mesoventrite well marked; mesoventral process about twice as wide as long; mesepisternum smooth. Penis short; trigonium short, broad, anterior angles each with a short hook; tegmen without styli (Fig. 65 in Watts 2011). Female micropterous, with disc of pronotum deeply excavated each side of midline, excavation open behind; ventrite 4 without row of modified setae; gonocoxites strongly sclerotised, short, shovel-like; gonostyli minute, subapical; prehensor well developed (Fig. 32 in Watts 2011). Larvae not known. Included species. Chameloscyphon huonensis Watts.	en	Watts, C. H. S., Bradford, T. M., Cooper, S. J. B. (2021): A new genus, Perplexacara, and new generic placements of species of Australian marsh beetles (Coleoptera: Scirtidae) based on morphology and molecular genetic data. Zootaxa 4927 (4): 539-548, DOI: 10.11646/zootaxa.4927.4.4
081787E4FF8BFF82FF4EFD40FD905E8D.taxon	diagnosis	Diagnosis. Length 6 – 11 mm. Eyes small, distance from subgenal ridge to bottom of eye a little more than diameter of eye, subantennal groove deep, with supraantennal ridge separating groove from eye, subantennal groove deep, with supraantennal ridge separating groove from eye, subgenal ridge without buttonhole; antenna without enlarged scape, antennomeres 2 & 3 not reduced; labial palpi straight (sensu Watts & Zwick 2019); mandible with one strong tooth, molar without small spines; pronotum transverse, anterolateral angles strongly produced forward; pronotal process broad at apex, corresponding notch in mesoventrite well marked; mesoventral process about twice as wide as long; mesepisternum rugose. Penis moderately elongate, trigonium bilobed, tegmen without styli (Figs 60 – 62 in Watts 2011). Female micropterous; ventrite 4 with a row of long setae near posterior edge; gonocoxites elongate, moderately sclerotized, gonostyli moderately long, apical; prehensor well developed (Fig. 33 in Watts 2011, Ruta 2020). Larvae not known. Included species. Daploeuros lamingtonensis Watts, 2011; D. reichertae Ruta, 2020; D. spencei (Armstrong, 1953); D. hadrostiktos Watts, 2011.	en	Watts, C. H. S., Bradford, T. M., Cooper, S. J. B. (2021): A new genus, Perplexacara, and new generic placements of species of Australian marsh beetles (Coleoptera: Scirtidae) based on morphology and molecular genetic data. Zootaxa 4927 (4): 539-548, DOI: 10.11646/zootaxa.4927.4.4
081787E4FF8BFF82FF4EFA80FBA45CE1.taxon	diagnosis	Diagnosis. Length 3 – 5 mm. Legs stout. Eyes small, head recumbent, distance from subgenal ridge to bottom of eye a little more than diameter of eye, subantennal groove deep, with supraantennal ridge separating groove from eye, antenna without enlarged scape, antennomeres 2 and 3 not reduced; labial palpi straight (sensu Watts & Zwick (2019 )) or nearly so (in Dasyscyphon tasmanicus); mandible with one strong tooth, molar without small spines; pronotum transverse, anterolateral angles weakly projecting forward; elytral punctures much larger than those on pronotum; pronotal process broad at apex, corresponding notch in mesoventrite well marked; mesoventral process about twice as wide as long; mesepisternum rugose / punctate. Penis stout, simple; trigonium finger-like, as long as parameroids; tegmen without styli (Figs 63, 64 in Watts 2011). Female winged, ventrite 4 without transverse row of long setae; gonocoxites thin, moderately sclerotised, gonostyli thin, very short, apical; prehensor well developed (Figs 43, 44 in Watts (2011). Larvae not known. Included species. Dasyscyphon hadrostiktos Watts; D. victoriaensis Watts.	en	Watts, C. H. S., Bradford, T. M., Cooper, S. J. B. (2021): A new genus, Perplexacara, and new generic placements of species of Australian marsh beetles (Coleoptera: Scirtidae) based on morphology and molecular genetic data. Zootaxa 4927 (4): 539-548, DOI: 10.11646/zootaxa.4927.4.4
081787E4FF8BFF85FF4EF8A4FDA359DB.taxon	description	Modified description. Length 4.5 – 11.0 mm. Eye normal size; without well-marked subantennal carina, without subantennal groove; antenna with scape greatly enlarged, widely oval, compressed, upper edge sharp, antennomere 2 small, inserted at end of scape, antennomere 3 smaller, together equal to or shorter than antennomere 4; subgenal ridge without buttonhole; with one strong mandibular tooth and a smaller one and / or a roughened area behind it; labial palpi forked (sensu Watts & Zwick 2019) or straight; anterolateral angles of pronotum square or projected forward, anterolateral angles of clypeus not projecting forward; mesoventral notch relatively long, narrowly ‘ V’ - shaped; pilosity on ventrites uniform except in M. insolitus which has a small area of larger punctures / setae on each side of ventrite. Male genitalia relatively simple, without endophallus (Fig. 66 in Watts 2011). Females winged, gonocoxites elongate, never strongly sclerotized. Notes. Macrodascillus was first proposed by Carter (1935) to accommodate a large scirtid from Barrington Tops New South Wales which he named M. denticornis. This species was subsequently shown by Watts (2011) to be conspecific with Helodes scalaris Lea, 1895 leading to the placement of H. scalaris into Macrodascillus. The species is recognized by its large size, enlarged scape, very small antennomeres 2 and 3, serrated antennae particularly in the male, strong mandibles each with a tooth, the anterolateral angles of the pronotum projected forward, male genitalia simple and female ovipositor simple with well-developed prehensor (Watts 2011, Watts & Zwick 2019). A second relatively similar species, M. pannuceus Watts, was added by Watts (2011). This paper adds two additional species, M. insolitus (Watts 2010) and M. lamingtonensis (Watts 2010). When describing these two species Watts 2010 drew attention to their similarity to M. scalaris, particularly the form of the scape and antennomeres 2 and 3. Phylogenetic analysis of the sequence data in this paper places both of these species as congeneric with M. scalaris and we herein transfer them from Prionocyphon Redenbacher to Macrodascillus Carter. The larvae of three of the species now in Macrodascillus are known (Watts 2014). Those of M. insolitus and M. scalaris are saproxylic and those of M. lamingtonensis live in small rock pools beside creeks and rivers. These larvae are morphologically distinct from each other and do not support a close relationship between these three species.	en	Watts, C. H. S., Bradford, T. M., Cooper, S. J. B. (2021): A new genus, Perplexacara, and new generic placements of species of Australian marsh beetles (Coleoptera: Scirtidae) based on morphology and molecular genetic data. Zootaxa 4927 (4): 539-548, DOI: 10.11646/zootaxa.4927.4.4
081787E4FF8BFF85FF4EF8A4FDA359DB.taxon	materials_examined	The genus as now constituted is found in wet Eucalypt forest from southern Queensland to southern Tasmania with one species (M. pannuceus) restricted to higher areas of the Atherton Tablelands.	en	Watts, C. H. S., Bradford, T. M., Cooper, S. J. B. (2021): A new genus, Perplexacara, and new generic placements of species of Australian marsh beetles (Coleoptera: Scirtidae) based on morphology and molecular genetic data. Zootaxa 4927 (4): 539-548, DOI: 10.11646/zootaxa.4927.4.4
081787E4FF8BFF85FF4EF8A4FDA359DB.taxon	discussion	Included species. Macrodascillus insolitus (Watts, 2010), M. lamingtonensis (Watts, 2010), M. scalaris (Lea, 1895), M. pannuceus Watts, 2011.	en	Watts, C. H. S., Bradford, T. M., Cooper, S. J. B. (2021): A new genus, Perplexacara, and new generic placements of species of Australian marsh beetles (Coleoptera: Scirtidae) based on morphology and molecular genetic data. Zootaxa 4927 (4): 539-548, DOI: 10.11646/zootaxa.4927.4.4
081787E4FF8CFF87FF4EFBB2FE83582F.taxon	description	Description. Length 3.0 – 5.5 mm. Eye normal size (small in P. latusmandibulara); without well-marked subantennal groove or carina. Antenna with scape greatly enlarged, widely oval, compressed, upper edge sharp; antennomere 2 small, inserted at end of scape, antennomere 3 smaller, together equal to or shorter than antennomere 4; subgenal ridge without buttonhole; mandible with one small tooth near apex; anterolateral angles of pronotum square or projected forward, anterolateral angles of clypeus not projecting forward (except in P. latusmandibulara); labial palpi forked (sensu Watts & Zwick 2019); pronotal process narrow between procoxae, expanding to a narrow diamondshape at apex; mesoventral notch absent or small and U-shaped; pilosity on ventrites uniform. Females winged, gonocoxites elongate, never strongly sclerotized. Larvae not known.	en	Watts, C. H. S., Bradford, T. M., Cooper, S. J. B. (2021): A new genus, Perplexacara, and new generic placements of species of Australian marsh beetles (Coleoptera: Scirtidae) based on morphology and molecular genetic data. Zootaxa 4927 (4): 539-548, DOI: 10.11646/zootaxa.4927.4.4
081787E4FF8CFF87FF4EFBB2FE83582F.taxon	etymology	Etymology. Latin perplexa – enigmatic, a reference to the enigmatic morphology of the three included species. Cara – a common Scirtidae genus ending for Australian Scirtidae. Gender feminine.	en	Watts, C. H. S., Bradford, T. M., Cooper, S. J. B. (2021): A new genus, Perplexacara, and new generic placements of species of Australian marsh beetles (Coleoptera: Scirtidae) based on morphology and molecular genetic data. Zootaxa 4927 (4): 539-548, DOI: 10.11646/zootaxa.4927.4.4
081787E4FF8CFF87FF4EFBB2FE83582F.taxon	type_taxon	Type species. Prionocyphon macroflavidus Watts. Notes. A group of three species previously placed in the genus Prionocyphon principally on their enlarged scape and small antennomeres 2 and 3. All three of these species were considered to be atypical Prionocyphon by Watts (2010) when describing them although all in different ways. Perplexacara latusmandibulara in particular has a large number of distinctive apomorphic character states (Watts 2010). Phylogenetic analysis of sequence data in Cooper et al. (2014) placed the three species now grouped together under Perplexacara as distinct from the other Australian Prionocyphon as well as from P. serricornis Müller, the type species of the genus. The current study strongly suggests that, despite their considerable morphological differences, they form a close phylogenetic relationship. These three species can be separated from the other Australian species currently placed in Prionocyphon by the lack of a buttonhole on the subgenal ridge, the homogeneity of the abdominal pilosity, and the absence of an endophallus, all characters found in many of the Australian species currently placed in Prionocyphon. Clearly the relationship of European Prionocyphon serricornis to Perplexacara and the Australian species remaining in Prionocyphon, together with the other Prionocyphon- like species from Southeast Asia requires additional study (Zwick 2016). Although well separated genetically (Fig. 1) species of Perplexacara are morphologically close to Macrodascillus but can be separated from this genus by their smaller size, presence of only one small tooth on the mandible rather than one strong one with a roughened area behind it and a small, U’ - shaped mesoventral notch. Larvae unknown. Notes. The genus is found in wet Eucalypt forest in Southeast Australia from northern New South Wales to southern Tasmania. Included species. Perplexacara caementum (Watts, 2010), P. latusmandibulara (Watts, 2010), P. macroflavida (Watts, 2010).	en	Watts, C. H. S., Bradford, T. M., Cooper, S. J. B. (2021): A new genus, Perplexacara, and new generic placements of species of Australian marsh beetles (Coleoptera: Scirtidae) based on morphology and molecular genetic data. Zootaxa 4927 (4): 539-548, DOI: 10.11646/zootaxa.4927.4.4
