taxonID	type	description	language	source
03FB8786FFE7FFCF498EFAC6B0DBFA83.taxon	type_taxon	Type species. Jubogaster towai sp. nov. here designated.	en	Parker, Joseph, Maruyama, Munetoshi (2013): Jubogaster towai, a new Neotropical genus and species of Trogastrini (Coleoptera: Staphylinidae: Pselaphinae) exhibiting myrmecophily and extreme body enlargement. Zootaxa 3630 (2): 369-378, DOI: 10.11646/zootaxa.3630.2.11
03FB8786FFE7FFCF498EFAC6B0DBFA83.taxon	diagnosis	Diagnosis. Euplectite pselaphines with the following characters: transverse head articulating ventrally under pronotum when deflexed, typical of Trogastrini. Head with vertex excavate, lacking vertexal fovea; ventral head without the typical dense “ genal beard ” of setae common to most Trogastrini. Eleven-segmented moderately geniculate antennae, received by large excavations anterior to eyes. Pronotum jubine-like: broad and obcordate, constricted at basal quarter, with weakly-evident medial longitudinal sulcus and distinct antebasal sulcus. Pronotum afoveate, with margins smooth, devoid of crenulations or small spines found in other Trogastrini. Elytra lacking any stereotypical fovea. Legs with contiguous, projecting coxae, broad femora, basally flattened tibia, and three tarsal segments; segment 1 very small, 2 and 3 elongate and subequal in length. Tarsi with two equally-sized claws, unlike other Trogastrini which have unequally-sized claws. Bifid anterior protarsal claw in the type species J. towai.	en	Parker, Joseph, Maruyama, Munetoshi (2013): Jubogaster towai, a new Neotropical genus and species of Trogastrini (Coleoptera: Staphylinidae: Pselaphinae) exhibiting myrmecophily and extreme body enlargement. Zootaxa 3630 (2): 369-378, DOI: 10.11646/zootaxa.3630.2.11
03FB8786FFE7FFCF498EFAC6B0DBFA83.taxon	etymology	Etymology. A combination of Jubus Schaufuss and Trogaster Sharp, the type genera of Jubini and Trogastrini, respectively. The name is chosen on account of the membership of the new genus in Trogastrini, despite apparent Jubini-like characters such as the shape of the pronotum and the presence of two equally-sized tarsal claws (a character state found in Jubini, but not other Euplectitae). The bifid anterior protarsal claw of the type species J. towai is also shared with Phamisus myrmophilus Silvestri of Jubini.	en	Parker, Joseph, Maruyama, Munetoshi (2013): Jubogaster towai, a new Neotropical genus and species of Trogastrini (Coleoptera: Staphylinidae: Pselaphinae) exhibiting myrmecophily and extreme body enlargement. Zootaxa 3630 (2): 369-378, DOI: 10.11646/zootaxa.3630.2.11
03FB8786FFE7FFCF498EFAC6B0DBFA83.taxon	description	Description. Body large (5.1 mm for the type species J. towai shown in Figure 1). Body shape approximately ovoid and strongly convex. Head: Approximately triangular, strongly transverse (Fig 4). Vertex afoveate, concave, with moderately deep excavation extending from basal quarter to anterior margin of frons; excavation narrowing apically along midline as it passes between raised frontolateral margins. Frontolateral margins carinate, extending roughly arcuately from posterior of antennal insertion cavity to clypeus (Fig 5). Frontolateral face accommodating large circular cavity anterior to eyes to receive antennae (Fig 5). Eye flanked directly anteriorly by additional large excavation for retracted maxillary palpus (Fig 5). Ventral surface of head bisected by longitudinal gular carina situated within broader gular sulcus terminating in gular fovea. Gular sulcus widens basally, spanning the head-neck interface. Maxillary palpi apparently 4 - segmented, with segment II long and curved at apex, III short and triangular, wider than II, IV oval, approximately 2 x longer than wide (Fig 5). Antennae (Fig 3) with 11 antennomeres, moderately geniculate with enlarged segment I. Antennomeres IX – XI strongly elongate (XI longest). Thorax: Pronotum (Fig 4) obcordate, 2 x wider than head, almost as wide as elytra. Afoveate, with smooth margins. Anterior margin broadly arcuate, largely obscuring the deflexed head when viewed from above, and constricted sharply in basal quarter, so that pronotum is laterally expanded into broad, blunt spurs with curved edges. Medial longitudinal sulcus present on pronotal disc, weakly impressed, thin, terminating at deeper, bowshaped antebasal sulcus that extends between bases of mediolateral spurs. Prosternum afoveate, with lateral longitudinal carinae extending from sides of neck to base. Mesosternum with median mesosternal foveae (MMSF), lateral mesosternal fovea (LMSF) and lateral mesocoxal fovea (LMCF) present. Other typical pselaphine mesosternal fovea listed in Chandler (2001) lacking. Metasternum strongly convex. Lateral metathoracic foveae (LMTF) present; median metathoracic fovea (MMTF) absent. Abdomen: Abdomen with tergites strongly convex. Visible segments 1 – 3 (IV – VI) subequal in length (Fig 1), segment 4 (VII) longest (not apparent in Fig 1 due to curvature of abdomen). Tergites 1 – 3 (IV – VI) with pronounced paratergites, 4 (VII) with small tapering paratergites. Tergite 6 (IX) internalised. Six sternites visible (Fig 2), sternite 7 (IX) internalised. First visible sternite (III) with median process projecting between metacoxae. Sternites 2 and 3 (IV and V) subequal in length, 4 (VI) slightly shorter (Fig 2). Elytra: Strongly convex, slightly broader than pronotum (Fig 1), approximately 0.3 x total body length. At their longest, equal in length to visible abdominal segments 2 + 3 (V + VI) combined. Elytral humeri broadly rounded to almost half elytral length. Sutural stria entire; lacking sutural fovea and other stereotypical antebasal fovea. Lacking subhumeral fovea and subhumeral stria. One weakly defined, somewhat discal depression is evident halfway between suture and humerus; its homology to canonical pselaphine antebasal elytral fovea is unclear. Legs: All coxae contiguous (Fig 2), procoxae conical, projecting ventrally, mesocoxae globose, metacoxae (Fig 7) transverse-conical, spanning from ventral midline to metasternal margin, and projecting posteriorly. Femora somewhat thickened, and excavated apically to receive retracted tibia. Tibia flattened basally. Femora and tibia lacking spines or processes. Tarsi elongate, 3 - segmented, first segment extremely short, 0.2 x the length of segment 2. Segment 2 and 3 subequal in length. All tarsi with 2 well-developed, equally-sized tarsal claws. Bifid anterior protarsal claw in type species J. towai (Fig 8).	en	Parker, Joseph, Maruyama, Munetoshi (2013): Jubogaster towai, a new Neotropical genus and species of Trogastrini (Coleoptera: Staphylinidae: Pselaphinae) exhibiting myrmecophily and extreme body enlargement. Zootaxa 3630 (2): 369-378, DOI: 10.11646/zootaxa.3630.2.11
03FB8786FFE6FFCA498EFA2EB0FDF9B5.taxon	materials_examined	type material. Holotype, male, Peru, Satipo, Santa Ana, 25 I 2012, with Pheidole xanthogaster Wilson. Specimen deposited in Field Museum of Natural History, Chicago, IL, USA, with catalog number: FMNH-INS 0000 093 389.	en	Parker, Joseph, Maruyama, Munetoshi (2013): Jubogaster towai, a new Neotropical genus and species of Trogastrini (Coleoptera: Staphylinidae: Pselaphinae) exhibiting myrmecophily and extreme body enlargement. Zootaxa 3630 (2): 369-378, DOI: 10.11646/zootaxa.3630.2.11
03FB8786FFE6FFCA498EFA2EB0FDF9B5.taxon	diagnosis	Diagnosis. The sole known species of Jubogaster is highly distinctive among Trogastrini due to its greatly enlarged and robust body covered in thick, blunt golden setae, the unique concave head that lacks vertexal fovea, the large, circular excavations on the frons that receive the long, geniculate antennae, the absence of a genal beard, an afoveate pronotum and elytra, the two equally-sized tarsal claws, and the bifid anterior protarsal claw.	en	Parker, Joseph, Maruyama, Munetoshi (2013): Jubogaster towai, a new Neotropical genus and species of Trogastrini (Coleoptera: Staphylinidae: Pselaphinae) exhibiting myrmecophily and extreme body enlargement. Zootaxa 3630 (2): 369-378, DOI: 10.11646/zootaxa.3630.2.11
03FB8786FFE6FFCA498EFA2EB0FDF9B5.taxon	etymology	Etymology. The new species is named in honour of Towa Shimada, son of Taku Shimada, a remarkable collector of myrmecophiles.	en	Parker, Joseph, Maruyama, Munetoshi (2013): Jubogaster towai, a new Neotropical genus and species of Trogastrini (Coleoptera: Staphylinidae: Pselaphinae) exhibiting myrmecophily and extreme body enlargement. Zootaxa 3630 (2): 369-378, DOI: 10.11646/zootaxa.3630.2.11
03FB8786FFE6FFCA498EFA2EB0FDF9B5.taxon	description	Description. Body length 5.1 mm (Fig 1). Colour dark reddish brown, antennae and legs somewhat lighter. Integument shiny. Most dorsal regions covered by conspicuous thick, erect, blunt golden setae, arranged at regularly spaced intervals and orientated posteriorly (depicted most clearly in figures 4 – 6). These are interspersed with shorter pubescence of the same colour, also orientated posteriorly. Ventral surfaces covered with fine, mostly aciculate golden pubescence (Fig 2). Head: Eyes moderate in size. Transverse, triangular head (Fig 4) widest just behind eyes. Vertex and occiput uniformly setiferous with thick, blunt, erect golden setae, and shorter, sharper pubescence. Margins of large lateral cavities for antennal insertion and maxillary palp retraction carinate; interiors of cavities glabrous (Fig 5). Neck approximately 0.4 x head width between eyes, shining and with sparse, very short dark setae on dorsal side; glabrous and with finger-print-like (“ strigate ”) microsculpturation on ventral side. Fourth segments of maxillary palpi (Fig 5) densely covered in short golden setae. Antennae (Fig 3) pigmented similarly to tibia and very elongate, approximately 2 x pronotum length. Antennomere I conical, widest in basal half and equal in length to III + IV combined. Laterodorsal margin with short arcuate carina extending to one-quarter antennomere length. Lateral face glabrous in basal half. With circular apical notch to permit geniculation with II; II small and cylindrical, approximately 0.6 x the length of III; III cylindrical-obconical, equal in length to IV; IV cylindrical, twice as long as wide; V slightly shorter and narrower than IV; V – VIII cylindrical-obconical, becoming gradually shorter apically, so that segment V is almost 2 x longer than wide and VIII is as only slightly longer than wide; IX – X much larger, IX almost as long as VI – VIII combined; X slightly longer than VII + VIII combined. Both IX and X cylindrical-obconical, apical margins somewhat sloping interiorly due to slight lengthening of exterior antennomere face relative to interior face. Antennomere XI longest, twice the length of X; exterior face shallowly and evenly convex, interior face weakly concave before bending obtusely to form conical apex. Several long sensory setae on apical third (Fig 3). Thorax: Pronotum (Fig 4) obcordate, margins smooth, lacking crenulations or other protuberances. Uniformly setiferous and pubescent. Prosternum with lateral longitudinal carinae abruptly bending dorsally on reaching base (Fig 2). Region between carinae with reduced setation and pubescence relative to lateral regions, and with small transverse projection situated medially, anterior to procoxae. Region between carinae projecting sharply in basolateral portions to receive procoxae. Mesosternum with setiferous median mesosternal foveae (MMSF) and lateral mesocoxal fovea (LMCF). Region anterior to mesocoxae formed into a shield-shaped plate with carinate margins. Anterior margin of metasternum carinate around mesocoxae. Convex metasternum with median declivity adjacent to metacoxal projections. Metasternum uniformly covered with yellow pubescence of intermediate length. Median declivity somewhat less pubescent. Lateral metasternal fovea small and setiferous. Posterior margin of metasternum bow shaped (Fig 2); projection between metacoxae with small medial indentation. Abdomen: Tergites strongly convex with visible tergite 1 (IV) sloping to abdomen base. Visible tergite 2 (V) broadest, slightly broader than 1 (IV), with segments 3 – 5 (VI – VIII) evenly narrowing. Tergite 4 (VII) longest, approximately 1.5 x tergite 3 (VI). Broad paratergites on segments 1 – 3 (IV – VI); paratergite margins of segment 4 converging apically to create a triangular paratergite. Anterior edges of abdominal tergites with narrow band of black pigment. Remainder of tergite reddish brown; intertergital membrane yellow with latticed microsculpture. Base of visible tergite 1 (IV) with short, blunt tubercles, situated halfway between midline and lateral margin. First visible sternite (IV) medially acuminate in anterior half, projecting ventrally between metacoxae. Last visible segment (6; VIII) with median declivity. Sternite 7 (XI) internalised. Pubescence on sternites 2 – 6 (IV – VIII) becoming finer and sparser toward midline (Fig 2). Elytra: Blunt setae on elytra somewhat longer and slightly thicker than on other parts of body. Apicolateral angles of elytra with margins weakly excised (Fig 6). Legs: Coxae contiguous. Mesocoxae with proximodistal carina along posterior face. Metacoxae glabrous except for pubescence on internal face and around projection to trochanter. Apices of metacoxae produced into flat, blunt spines that extend posteriorly to half trochanter length (Fig 7). Metatrochanters with small rhomboid-shaped tubercles on exterior face (Fig 7). Femora brownish red, covered with thick, blunt golden setae. Tibia somewhat flattened in proximal half, carinate along anterior edge to half-tibia length. Tibia lighter in colour than femora, yellowish brown, with similar but shorter setae. Tarsi narrow and longer than half tibial length; also yellowish brown, with fine, sharp, aciculate setae. Protarsus with bifid anterior claw (Fig 8). Aedeagus: Asymmetric, broad, lacking parameres (Fig 9 and 10). Wide, globose basal bulb with asymmetric left and right apical projections. An asymmetric, apically-truncate sclerotised stylus emerges from within the basal bulb.	en	Parker, Joseph, Maruyama, Munetoshi (2013): Jubogaster towai, a new Neotropical genus and species of Trogastrini (Coleoptera: Staphylinidae: Pselaphinae) exhibiting myrmecophily and extreme body enlargement. Zootaxa 3630 (2): 369-378, DOI: 10.11646/zootaxa.3630.2.11
03FB8786FFE6FFCA498EFA2EB0FDF9B5.taxon	biology_ecology	Biological and collection information. The single specimen of J. towai was collected in a colony of Pheidole xanthogaster Wilson under a log. The ants were apparently unaware of the beetle, which was was found deep inside the nest. This suggests that J. towai is a true myrmecophile, but information about its biology and behaviour are unavailable at this time. It has no clear myrmecophilous adaptions, so may belong to the “ obligate synoekete ” class.	en	Parker, Joseph, Maruyama, Munetoshi (2013): Jubogaster towai, a new Neotropical genus and species of Trogastrini (Coleoptera: Staphylinidae: Pselaphinae) exhibiting myrmecophily and extreme body enlargement. Zootaxa 3630 (2): 369-378, DOI: 10.11646/zootaxa.3630.2.11
03FB8786FFE6FFCA498EFA2EB0FDF9B5.taxon	discussion	Comments. J. towai has distinctly internalised terminal tergites and sternites. The plesiomorphic condition in Pselaphinae is for visible sternite 7 (IX) to form a penial plate, flanked by left and right sclerites formed from visible tergite 6 (which is also segment IX). However, in J. towai, segment IX is internalized; in figure 2 the internal sclerites can be seen enclosed within the genital aperture. This arrangement makes segment VIII (visible sternite 6 and visible tergite 5) the posterior-most external segment. Don Chandler (University of New Hampshire) has kindly surveyed the character state of segment 7 (XI) and found internalisation to be relatively common in Trogastrini: in many genera the segment is not externally visibile (Rhexinia Raffray, Xerhius Raffray, Aporhexius Raffray, Rhexius LeConte, Eurhexius Sharp, Platomesus Chandler, Austrorhexius Chandler) while in others it is clearly present (Conoplectus Brendel, Fletcherhexius Park, Adrogaster Raffray). In still others, the segment is visible but thin (Anarmodius Raffray, Rhexidius Casey). Two genera, Oropus and Mesoplatus, are polymorphic, with species falling into two or all of the three classes. Furthermore, in most Jubini and some Goniaceritae, segment 7 is not externally apparent (D. S. Chandler, pers comm.). Hence, a recurring trend is for internalization of the terminal segment, as exemplified by J. towai.	en	Parker, Joseph, Maruyama, Munetoshi (2013): Jubogaster towai, a new Neotropical genus and species of Trogastrini (Coleoptera: Staphylinidae: Pselaphinae) exhibiting myrmecophily and extreme body enlargement. Zootaxa 3630 (2): 369-378, DOI: 10.11646/zootaxa.3630.2.11
03FB8786FFE3FFCB498EF94AB35AFD14.taxon	discussion	While the head of Jubogaster is suggestive of Trogastrini, other characters typical of this tribe are absent. There is no dense genal beard of setae, the tarsal claws are both well-developed and equally-sized, rather than being unequal in size or consisting of one well-developed and one setiferous claw. A strongly-impressed median longitudinal sulcus on the pronotal disc is also lacking; although the sulcus is present, it is barely evident and only weaklydefined. Furthermore, the obcordate form of the pronotum, sharply constricted at the base, is far more similar to members of Jubini, another euplectite tribe. Jubogaster ’ s curious suite of features confounds a definitive assignment of the new taxon to tribe using morphology alone. To reliably place Jubogaster phylogenetically, we employed a non-destructive procedure to extract DNA from the type specimen, without damaging or altering its external morphology (Gilbert et al. 2007). Using this approach, we successfully amplified a fragment of 28 s rDNA from J. towai. This region is particularly informative for higher-level pselaphine systematics, resolving many relationships at the supertribal, tribal and subtribal levels (J. Parker and A. Vogler, unpublished data). We also sequenced this gene region from a range of other Pselaphinae including 5 genera of Trogastrini, 3 Jubini genera, and representatives of all other euplectite tribes except Metopiasini, Dimerini and Mayetiini. A variety of non-euplectite pselaphines from different supertribes were included, totalling 24 ingroup taxa. Three other staphylinids (from the subfamilies Proteininae, Euaesthetinae and Scydmaeninae) and the silphid Silpha obscura were used as outgroups. Both MP (Fig 11) and Bayesian (Fig 12) analyses placed Jubogaster within the Trogastrini with maximum support, forming a clade with Platomesus and Rhexius. This placement was recovered across all alignment parameter combinations tested with consistently high support. Although the taxonomic coverage is limited, Jubogaster emerges in the same position in a far more comprehensive analysis of> 150 ingroup taxa, with extensive taxon sampling from almost all pselaphine tribes and subtribes (data not shown; J. Parker and A. Vogler, unpublished data). Hence, we conclude with a high degree of certainty that Jubogaster is indeed a member of Trogastrini.	en	Parker, Joseph, Maruyama, Munetoshi (2013): Jubogaster towai, a new Neotropical genus and species of Trogastrini (Coleoptera: Staphylinidae: Pselaphinae) exhibiting myrmecophily and extreme body enlargement. Zootaxa 3630 (2): 369-378, DOI: 10.11646/zootaxa.3630.2.11
