Tauritermes bandeirai: A new drywood termite (Isoptera, Kalotermitidae) from the Caatinga and Atlantic Forest of Brazil

Abstract The imago and soldier castes of a new Tauritermes Krishna, 1961 species, Tauritermes bandeiraisp. nov. are described. It is the fourth species of Tauritermes and occurs from the Caatinga and Atlantic Forest of Brazil. Unlike its congeners, the soldier of T. bandeirai has prominent frontal horns.

Tauritermes, as other kalotermitid genera, is best characterized by venation of the winged imago and soldier head capsule morphology. For Kalotermitidae, most diagnostic characters at the intrageneric level are attributed to head capsule characters. Noirot (1995) reported some consistent differences in the gut anatomy of kalotermitids compared to other lower termite families. Furthermore, Gonçalvez (1979), Myles (1997), and Godoy (2004) described parts of the gut of Rugitermes, Marginitermes, and Tauritermes, respectively. They found that differentiation of the gut morphology of kalotermitids remains elusive at both the generic and specific levels.
Herein, we report on a new Tauritermes species, T. bandeirai sp. nov. from samples collected by Vasconcellos et al. (2005) and additional specimens from the Brazilian Caatinga and Atlantic Forest. This is the fourth species of Tauritermes to be described based on external imago and soldier characters.

Material and methods
Photomicrographs were taken as multi-layer montages using a Leica M205C stereomicroscope controlled by the Leica Application Suite version 3 software. Preserved specimens were taken from 85% ethanol and suspended in a pool of Purell Hand Sanitizer to position the specimens on a transparent Petri dish background. All University of Florida Termite Collection (UFTC) records are available online (Scheffrahn 2019b).

Taxonomy
Tauritermes bandeirai sp. nov. http://zoobank.org/3C739327-4FB8-4B77-8AD0-862A82237292 Figures 1, 2 Diagnosis. The soldier of T. bandeirai differs from soldiers of the other three Tauritermes species by having a distinct and robust frontal horn and a roundly protruding dorsal horn (Fig. 2C). The dorsal and frontal horns of T. triceromegas (Silvestri, 1901) are more angular but much smaller, barely elevated above the frons in oblique view (Fig. 3C). In T. vitulus Araujo & Fontes, 1979, the dorsal horn is similar to that of T. bandeirai sp. nov., while the fronal horn is absent (Araujo and Fontes 1979, fig. 10). In T. taurocephalus (Silvestri, 1901), the dorsal horn is more elevated and angular, while the frontal horn is also absent (Fig. 4C).
The T. bandeirai soldier differs from its congeners in that its frons and horns are more rugose, the basal mandible humps are broader and more angular, and the third antennal article is more club-shaped. Only T. taurocephalus has a similarly shaped postmentum (Figs 2D, 4D). In T. triceromegas, the postmentum is posteriorly elongated (Fig. 3D) and in T. vitulus it is posteriorly widened (Araujo and Fontes 1979, fig. 15).
The imago head and pronotum of T. bandeirai are mostly unremarkable, except for a relatively large ocellus in comparison with the rather small compound eye. Among kalotermitid genera, the forewing venation is closest to Incisitermes Krishna with one diagnostic exception. In Incisitermes, the median vein is not sclerotized and its terminus does not closely approach the radial sector (Scheffrahn 2014, fig. 5). In T. bandeirai, the distal third of the median vein is sclerotized and closely paralells the radial sector (Fig. 1C). The wing venation of T. vitulus is similar to that of T. bandeirai, but no sclerotization is reported (Araujo and Fontes 1979, fig. 1).
Description. Imago ( Fig. 1A-C, Table 1). Head capsule and pronotum light brown. Compound eye obtusely triangular; ocellus a shade lighter than vertex, very large, and roundly ellipsoid; nearly touching eye margin. Vertex covered with about one dozen short setae. Pronotum about as wide as head capsule; anterior margin weakly emarginate in middle. Pronotum covered with a few dozen setae in middle, lateral margins with about one dozen setae each. Antennae with 15 articles, basal article relative lengths 1>2<3>4. Fore wing with subcosta joining costal margin at about oneeighth of wing length from suture. Radius joining costal margin at one-third wing length; radial sector with about four anterior branches. Median vein becoming lightly sclerotized at distal third as it encroaches near the radial sector. Wing membrane and first 4-5 branches of cubitus lightly pigmented, concolorous with apical radial sector and median veins. Arolium present.
Soldier ( Fig. 2A-D, Table 2). Head capsule, in dorsal view ( Fig. 2A), dark castaneous brown from postclypeus to antennal carinae, grading to orange at occiput. Head capsule narrowing toward antennal carinae; antennal carinae rugose and visible from above. Frontal ridge V-shaped with deep median cleft. About a dozen fine setae on vertex and genae. Eye spots hyaline, narrowly elliptical. In lateral view (Fig. 2B), dorsal horns (or protuberances) forming a rounded shelf near right angle; frontal horns projecting above base of mandibles. Genal horns slightly posterior to frontal horns, anterior to dorsal horns. Pronotum shield-shaped, wider than head; anterior margin rugose, incised in middle with rounded anterior lobes. In oblique view (Fig. 2C), dorsal and frontal horns rising prominently from frons; horns and frons coarsely rugose. Postclypeus (Fig. 2D) forming elongated, nearly symmetrical hexagon. Third antennal article slightly club-shaped, relative article length 2<3>4=5. Mandibles two-fifths length of head capsule; basal humps robust, rugose; lateral margin of humps parallel. Outside margin of blade nearly straight from above, then curving at right angle at onefifth from apex; apical tooth thick, marginal dentition weak. Mandibles curve evenly by about 15° in lateral view.   Distribution. (Fig. 5) Northeastern Brazil, Caatinga and Atlantic Forest biomes. Tauritermes localities taken from the literature are given in Table 3.
Etymology. Named for Dr. Adelmar Gomes Bandeira, the graduate and postgraduate advisor of AV who died in 2019. Dr. Bandeira was one of the first termitologists to work on termite ecology in the New World.
Biology. The colonies of T. bandeirai were collected inside dry trunks in the beginning stages of decomposition (diameter > 3cm) and in dead terminal branches still attached to the trunks, both in areas of Caatinga and Atlantic Forest. In the Caatinga, colonies of T. bandeirai were relatively easy to extract from dead terminal branches of Commiphora leptophloeos (Mart.) J.B. Gillett (Burseraceae). This tree is also a "hot spot" for collecting other kalotermitids, such as Cryptotermes, Neotermes, and Rugitermes.
Using light traps over a year (December 2017 to November 2018) in a Caatinga area located in the municipality of São José dos Cordeiros, Paraíba-Brazil, the alates of T. bandeirai were collected five times; once in December, thrice in January, and once in February. This period represents a transition between the dry and rainy season in the area. For the Atlantic Forest, alates were recorded in wood in March, June, and December.    Araujo and Fontes (1979) There are no records of Tauritermes bandeirai infestations in buildings, either in urban or agricultural environments. Other Tauritermes species infest sound, dry wood (RHS, unpubl.) and are even structural pests (Araujo and Fontes 1979).

Discussion
The Caatinga and the Atlantic Forest are neighboring domains (Fig. 5), but drastically different in terms of age and environmental conditions. The Caatinga is a semiarid region of northeastern Brazil and is part of the "Seasonally Dry Tropical Forests" (Silva et al. 2017). On the other hand, the Atlantic Forest is distributed along the east coast of South America and is part of the "Tropical Rain Forests" (Morellato and Haddad 2000). Even with such different physiognomies and ecological dynamics, several species of termites, in addition to T. bandeirai sp. nov, are found in both domains such as Heterotermes longiceps (Snyder, 1924), Ruptitermes reconditus (Silvestri, 1901), Nasutitermes macrocephalus (Silvestri, 1903), Microcerotemes indistinctus Mathews, 1977, among others.