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Two new genera and species of the termite symbiont lineage Termitohospitini (Coleoptera, Staphylinidae, Aleocharinae) from Bolivia and peninsular Malaysia

Abstract Coptotermocola clavicornis gen. & sp. n. and Neotermitosocius bolivianus gen. & sp. n. of the termite inquilinous tribe Termitohospitini are described from peninsular Malaysia and Bolivia, respectively. The Termitohospitini are most readily diagnosable by the distally migrated anterior tentorial pits that are no longer associated with the antennal fossae, and by the enlarged vertex which obscures the antennal fossae dorsally. Additionally, the Termitohospitini are hypothesized to share a recent common ancestor with the Masuriini and Myllaenini due to shared derived morphologies of the lacinia distal teeth with lateral cuticular processes, presence of a unique maxillary palpomere III sensilla, and anterolateral angles of mentum produced. Habitus photographs and illustrations of diagnostic features are provided for the two new genera in order to facilitate future work.


introduction
Eusocial insect colonies are often accompanied by a unique faunal assembly that takes advantage of available colony resources (e.g. Hölldobler andWilson 1990, Wheeler 1928). Symbionts of termites are commonly referred to as termitophiles (Kistner 1969). A large number of termitophiles have evolved within the beetle family Staphylinidae, particularly in lineages of the subfamily Aleocharinae. Among the recognized 58 tribes (Bouchard et al. 2011), 1,151 genera and 12,851 species (Thayer 2005) of Aleocharinae, approximately 650 species and 190 genera among 17 tribes have recorded associations with termites, of which eleven tribes are comprised exclusively of termitophilous species (Kanao unpublished data).
Termitophilous Aleocharinae that exhibit relatively complex symbiosis with their host termites have repeatedly converged on physogastric and limuloid body plans . Physogastry is the result of abdominal or thoracic inflation and is hypothesized to serve a mimetic function (Kistner 1969). Physogastric species are generally well integrated into the termite social system, actively interacting with their hosts and slow moving. On the other hand, limuloid taxa are teardrop-shaped and the body plan is hypothesized to serve a defensive function (Kistner 1979). Limuloid taxa often also exhibit extremely compact appendages and cavities to receive structures more vulnerable to host aggression (e.g. distal podomeres). In contrast to physogastric species, limuloid species are faster moving and usually do not actively interact with their hosts.
Most species of Termitohospitini ancestrally share a limuloid body form but a few taxa have secondarily evolved physogastric forms (Kanao et al. in preparation). Notably, the species of Coptoxenus Kistner, 1976 exhibits physogastry of the thorax, a peculiar condition among aleocharines. With a diversity of body forms, the tribe offers a unique opportunity to study the independent evolution of adaptive morphologies as it correlates with integration and behavior with host termite societies. However, as with most termitophiles, basic understanding of Termitohospitini biodiversity lags behind that of their free-living relatives. Inquilinous groups are difficult to collect and relatively few specimens are available for comprehensive understanding of their biodiversity. Here we describe two new genera and species of Termitohospitini as part of our ongoing research on the evolution of termitophily in termitohospitines. We provide habitus photographs, illustrations of diagnostic features, a new tribal diagnosis as an update to previous diagnoses (Seevers 1941(Seevers , 1957, and putative synapomorphies that support a close relationship of Termitohospitini with Masuriini and Myllaenini.

Materials and methods
The first author conducted fieldwork at Ulu Gombak, which is approximately 25 km northwest of Kuala Lumpur, Malaysia in 2010, 2011 and 2012. Specimens of a new genus collected at the site for this study were initially preserved in 2.0 ml vials of 80% ethanol. The other undescribed genus was discovered in the Snow Entomological Museum Collection at the University of Kansas.
The technical procedures used for this study generally follow those described in Maruyama (2006). When a permanent mount was made, cleared specimens or parts were dehydrated by successively transferring specimens to higher ethanol (EtOH) concentrations until finally reaching 100% EtOH, and then placed into Euparal. When appropriate, dissections were made within Euparal on a glass mounting substrate. Full body dissections were made and preserved on glass microscope slides. Dissected body parts were preserved and mounted on halved glass cover slips, subsequently glued onto a halved paper glue board, and mounted under the respective specimen (see Maruyama 2004, for details).

Termitohospitini Seevers, 1941
Type genus: Termitohospes Seevers, 1941. Diagnosis. Members of the tribe Termitohospitini most closely resemble the tribe Myllaenini and Masuriini (sensu Ahn and Ashe 2004), and are hypothesized to be closely related based on the following combination of putative synapomorphies: 1) subapical marginal setae of maxillary lacinia with basal, paired cuticular processes (spinose scales of Ahn and Ashe 2004); 2) presence of a unique sensory patch on lateral surface of maxillary palpomere III; 3) anterolateral angles of mentum produced.
Species of Termitohospitini are diagnosable from all other Aleocharinae, including Myllaenini and Masuriini, by the following putative synapomorphies: 1) anterior tentorial arms of the head migrated anteriorly and disassociated from antennal fossae (Seevers 1941); 2) antennal fossae dorsally obscured by enlarged vertex (Seevers 1941); 3) maxillary lacinia with basal paired cuticular processes reduced in size; 4) processes of anterolateral angles of mentum reduced in size; 5) ligula broad and reduced in length. Diagnosis. This monotypic genus is distinguishable from other Termitohospitini by its slender parallel-sided body, longer legs and apically truncate pronotum, which dorsally exposes much of the head (Fig. 1).

Neotermitosocius bolivianus
All type specimens are deposited in the Snow Entomological Museum Collection (SEMC).
Diagnosis. This species is diagnosable based on the generic diagnosis above. Description. Body (Figs 1-2) approximately 2 mm in length (1.84-2.00 mm, N = 2), almost uniformly orange brown, but head slightly darker. Dorsal surface of head (Fig. 3) sparsely covered with setae, a pair of long setae present at anterior margin of clypeus. Eyes with uniform cover of inter-ommatidial setae. Antennae (Fig. 4) sparsely covered with setae; antennomere I with a long macroseta at middle of internal lateral surface; antennomere II with 4-5 macrosetae around apical margin, one macroseta conspicuously longer; antennomere III with 4 thick, very long macrosetae at apical margin; antennomeres IV-X with 3-4 macrosetae around apical margin; antennomere XI with several macrosetae near middle and apex. Labrum (Fig. 5) dorsal surface with 2 pairs of setae at anterolateral corners, 3 pairs of longer and 3 pairs of shorter setae at mesal area of disc; epipharynx ( Fig. 5) with two pairs of lateral setulae. Mandible (Figs 6-7) dorsum with 2 pairs of scrobal setae near middle of aboral margin, six setae around base of disc. Maxillary (Fig. 8) lacinia with 2 pores at middle; galea with 3 pores apicomedially; maxillary palpal article I with a long seta at lateral margin and a medial pore, article II sparsely covered with setae, article III sparsely covered with setae and 2 stronger setae medially. Pronotum (Fig. 11) transverse (pronotum length = 0.30-0.40 mm, pronotum width = 0.54-0.72 mm, N = 7), densely covered with setae, with a pair of long macrosetae at anterolateral corners. Elytra (Fig. 12) longer than wide (elytra length = 0.42-0.44 mm, elytra width = 0.30-0.34 mm, N = 5), densely covered with setae, setae of lateral margins longer. Mesoventrite (Fig. 13) sparsely covered with minute setae at central and posterolateral areas. Metaventrite (Fig.13) posterior one third sparsely covered with setae. Fore leg (Fig. 14) uniformly covered with setae; coxa with a macroseta near dorsomedial of anterior surface and 5-6 long setae at apical margin; femur with longer setae near anterior inner margin; tibia with apical setae stronger, 2 spurs present at apex. Mid leg (Fig. 15) uniformly covered with setae except coxa; coxa with several long setae around anterior margin; tibia with long macroseta at middle on dorsal surface, 3 spurs present at apex. Hind leg (Fig. 16) uniformly covered with setae, coxa medially nude; coxa with 3 longer setae along cavity; trochanter and femur with a macroseta near middle of anterior surface; tibia with one longer and shorter macroseta on anterior surface, 3 apical spurs present.
Female. Spermatheca (Fig. 23) cuticle at apex with wrinkle-like sculpture. Etymology. The specific epithet is derived from its type locality of Bolivia Latinized. The gender is masculine. Distribution. Cochabamba, Bolivia. Ecology. All known specimens of Neotermitosocius bolivianus were collected with a passive sample method (flight intercept trap) and nothing is known about its biology. It is hypothesized to be a termite symbiont based on phylogenetic relationships but the host is unknown.
Diagnosis. This monotypic genus is distinguishable from all other Termitohospitini most distinctly by the strongly carinate mesocoxal process (Figs 27,38), and additionally by the compact antennae (Fig. 29) and short maxillary palpi (Fig. 33).
Head capsule (Fig. 28) transverse, widest behind eyes. Antennal fossae deep, as large as eyes and dorsally obscured by vertex. Eyes large and produced anterolaterally. Antennae (Fig. 29) compact with 11 articles; antennomere I longer than II-X and partially hidden within antennal fossae; antennomere II dilated at apex to receive antennomere III; antennomeres III-X strongly transverse and more than twice as wide as long, length increasing distally; antennomere XI fusiform; pedicles of antenomeres IV-XI obscured by apex of preceding article. Labrum (Fig. 30) transverse and semicircular, anterior margin broadly concave; disc basomedially sparsely covered with pores; epipharynx (Fig. 30) glabrous. Mandibles (Figs 31-32) asymmetrical, dorsally covered with numerous pores around middle. Right mandible (Fig. 32) with a small subapical tooth; anterior one fifth strongly curved adorally. Left mandible (Fig. 31) without a subapical tooth. Maxillary (Fig. 33) lacinia elongate and strongly recurved adorally, adoral margin with 11 setae; apical two setae without basal cuticular processes (spinose scale of Ahn and Ashe 2004), third apical seta with a proximal basal cuticular process, central five setae with paired basal cuticular processes, basal three setae without basal cuticular processes; galea tapered apically, apex densely furnished with long trichae; palpus with 4 articles; article I triangular; article II trapezoidal and dilated toward apex; article III oviform, more than three times longer than wide and sparsely covered with long setae; article IV slightly narrowed toward apex with inconspicuous pseudosegment, pseudopores present apicomedially. Mentum (Fig. 34) trapezoidal, disc covered with long setae and several pores near middle and lateral areas; anterolateral corners slightly produced and with a pair of long macrosetae. Labial (Fig 35) palpus with 3 articles, first article a fusion of I + II; article I + II with a mediobasal seta on ventral surface (probably homologous to seta c of Sawada [1972]), twin and medial pores present; article III almost half as long as article I + II with 2 pores at apex; ligula triangular with medial subtriangular area sclerotized; prementum wider than long, with 2 pairs of both real and pseudopores, and a pair of setal pores present; apodemes broad and longer than disc, dilated posteriorly with apices recurved internally and almost touching. Pronotum (Fig. 36) transverse, widest at posterolateral angles; anterior margin concave with anterolateral angles slightly produced anteriorly; posterior margin rounded and slightly produced medially; marginal cuticle thin and somewhat translucent (Fig.  26). Prosternum reduced in length. Elytra (Fig. 37) subquadrate, slightly wider than long. Wings fully developed. Mesoventrite (Fig. 38) slightly shorter than metaventrite; mesoventral process produced as an extremely large structure widest medially and tapered at ends, microsculpture composed of longitudinal wavy lines and pores (Fig.  39), structure ventrally partially obscuring basal podomeres; mesocoxal cavity marginal bead complete, narrowly separated. Legs (Figs 40-42) stout and laterally flattened; femora subrectangular; apical tarsomeres longest, tarsal formula 4-4-5. Fore leg ( Fig.  40) with coxa approximately as long as femur; trochanter subtriangular; tibia thin. Mid leg (Fig. 41) with coxa globular; trochanter very thin; tibia thin. Hind leg (Fig.  42) subrectangular and almost as long as trochanter and femur combined; trochanter globular-triangular; femur slightly dilated apically; tibia thin.
Etymology. The generic name is derived from a combination of the generic name of the host termite, Coptotermes Wasmann, 1896 and the Latin noun cola meaning "dweller". The gender is feminine. Paratypes: 6??, MALAYSIA: same data as the holotype, one specimen is preserved in 99.5% EtOH; ♀, same locality data as the holotype, differing data reads "XXI May 2010,/ T. Kanao leg. KT-33", fully disarticulated; 3??, same locality data as the holotype, differing data reads "XXIX May 2012,/ T. Kanao leg. KT-312", one specimen is preserved in 99.5% EtOH.
All type specimens are deposited in the Kyushu University Museum.
Female. Spermatheca (Fig. 49) apical bulb surface with transverse wrinkle-like sculpture; stalk basal to membranous area three times as long as apical bulb.
Etymology. The specific epithet is derived from a combination of the Latin noun clava meaning "club" and Latin adjective cornis meaning "to be horned", in reference to the diagnostic robust antennae of the species. The gender is feminine.
Ecology. Specimens acquired during the KT261 collecting event were collected from the galleries of the host termites within a rotting log. The galleries were large and arranged in a complex manner. Another specimen (KT33) was collected from a trail of the hosts that occupied the exterior of a large log. KT312 specimens were collected from a rotting log occupied by the host termites. All Coptotermes colonies that yielded C. clavicornis were located near rivers where the habitat in general was comparatively more moist compared to its surroundings.
All C. clavicornis specimens moved faster than their host termites. They did not avoid contact with hosts but instead recurved their abdomens over their bodies when they came into contact. The inquilines wedged themselves under their hosts on several occasions, but the host termites regaurdless never attacked the beetles.