Hidden Brazilian Amazon diversity: four new species, redescription and notes on natural history of Termitozophilus Silvestri, 1901 (Coleoptera: Staphylinidae: Aleocharinae) Author Silva, Ruan Felipe Da 0000-0002-7096-226X Programa de Pós-graduação em Biodiversidade e Evolução. & Museu Paraense Emílio Goeldi. Coordenação de Zoologia. Av. Perimetral, 1901, 66077 - 830, Belém, Pará, Brazil. silvaruanbio@gmail.com Author Zilberman, Bruno 0000-0003-2613-4827 Museu de Zoologia da Universidade de São Paulo, Caixa Postal 42494; 04218 - 970, São Paulo, Brazil. & Programa de Pós-graduação em Sistemática, Taxonomia Animal e Biodiversidade. brunozilberman@usp.br Author Carvalho-Filho, Fernando Da Silva 0000-0002-2480-3874 Museu Paraense Emílio Goeldi. Coordenação de Zoologia. Av. Perimetral, 1901, 66077 - 830, Belém, Pará, Brazil. fernandofilho@museu-goeldi.br text Zootaxa 2022 2022-11-21 5209 5 501 534 journal article 193695 10.11646/zootaxa.5209.5.1 80fa12f4-e62b-487d-91b1-b1ec37da5ba4 1175-5326 7469353 360B047E-5396-48AE-B26F-00B34BCB835A Termitozophilus Silvestri, 1901 Termitozophilus Silvestri, 1901: 7 (description, type species: Termitozophilus laetus Silvestri, 1901 ); Fenyes 1920: 348 (redescription); Seevers, 1957: 65 (redescription); Campbell, 1973: 92 (redescription and key); Jacobson et al. , 1986: 18 (redescription); Zilberman, 2019: 96 (redescription). Corymbogaster Mann, 1923: 346 (description, type species: Corymbogaster miranda Mann, 1923 ) [synonymized by Borgmeier, 1950: 638 , 640]. FIGURE 1. A) Cornitermes pilosus Holmgren nest, Peixe-Boi, Pará, Brazil. B) Cornitermes pilosus nest and collector (1.80m). C) Opened nest of C. pilosus . D) Fragment of the nest with Termitozophilus together with termites (arrow showing Termitozophilus ). E) Material utilized in nest sorting. F) Cornitermes and Termitozophilus specimens under observation in a petri dish (arrow showing Termitozophilus ). Termitozophilus has been recently redescribed by Zilberman (2019). However, because the four new species revealed in this study more than doubled the number of known species in the genus, a redescription and additional discussion are required. Head elongate to subquadrate; gula subtriangular, widest at base, fused to mentum; foramen magnum short with vertex bearing four to six bristles and with a strong V-shaped impression in front, between the eye. Antenna with 11 antennomeres, antennomeres shape varying among species, interarticular membrane between antennomeres II and III largely exposed. Labrum more than twice wider than long; five main bristles on each lateral half, distributed in two pairs in diagonal rows and one with variable position. Mandible symmetrical, internal margin with a mesal broad tooth; with pores on surface of mesal region. Maxilla with stipe almost as wide as long, a long bristle at apex, close to first palpomere, and four short bristles close to galea; lacina subquadrate to elongate, covered with bristles of different length at margins; galea narrow, about as long as lacina; surface covered with some moderately long bristles; maxillary palpus densely setose and with four palpomeres with different degree of sclerotization: palpomere I short and subquadrate, palpomeres II–IV elongate; only palpomeres II and III covered with long bristles; palpomeres I–III porose and with a scaled surface, gradually increasing from I to III; palpomere IV not scaled and with sensilla. Labium with epipharynx covered with pores and with short bristles; lateral lobes of epipharynx developed, with a row of bristles on each side; labial palpus setose, 3-articulated; palpomeres I–II subquadrate; last palpomere elongate and narrow; ligula narrowed at base, with rounded or bifid apex, almost variating in shape among species; mentum and submentum distinct or not; preapical and proximal bristles always present and apical bristle present or not. Abdomen. Inner and outer paratergites fused or not to a single sclerite by secondary sclerotization; paratergites fused or not to tergites or sternites; each inner and outer paratergite with two rows of three to six bristles; tergite VII from subquadrate to about twice as wide as long, with or without bristles; when present, distributed in two rows of long bristles, medial region with two bristles and posterior region with six bristles, short bristles covering entire surface, glandular structure present, continuous along its length (not separated). Aedeagus. Paramere welldeveloped, with two to four bristles on apical lobe and velar sac well developed extending to apical lobe; median lobe with rounded bulbus and with subquadrate tubus apex, with sclerites of the internal sac well developed, variable among species. Spermatheca. Capsule large and sclerotized; chamber and duct fused, curved, with duct placed in posterior region, weakly sclerotized. Oviparity. Knowledge about the reproductive biology of the species in the tribe Corotocini or even in other termitophilous taxa of Staphylinidae is poorly known. Emerson (1935) described a curious case of a Nasutitermes guyanae (Holmgren) nest with four species of Corotocini : Thyreoxenus major Mann , Eburniola leucogaster Mann (both Corotocina), Termitophya amica Mann (Abrotelina) and Xenopelta cornuta Mann (Termitogastrina) , where two kinds of larvae were found, as well as three different kinds of eggs, often found among termite eggs. One of the egg kinds was elongate (supposedly belonging to T . major ), and the other two were less elongate, with one kind being even smaller than a termite egg, which Emerson (1935) supposedly attributed to E. leucogaster specimens. Eight spherical eggs were found in a dissected female abdomen of Termitozophillus laetus housed at MZSUP ( Figs. 2A–D ). Although no Termitozophilus species eggs or larvae have been found in termite nests, we presume this genus is oviparous based on the number and size of eggs reported in T . laetus . The most complete reproductive biology data about a Corotocini species is now available for viviparous species of the genus Corotoca Schiødte , in which females carry three very large eggs in the abdomen with embryos that grow asynchronously ( Zilberman et al. 2019 ). Unlike Termitozophillus , spermatheca of Corotoca is reduced, though the relationship between spermatheca loss or reduction and viviparity, if any, is unclear ( Oliveira et al. 2018 ; Pisno et al. 2019; Zilberman et al. 2019 ). Although Corotocini biology is little understood, oviparity has been the most documented strategy in genera of this tribe ( Emerson 1935 , Seevers 1957 , Zilberman et al. , 2019 ). Geographic distribution and host relationship. The genus Termitozophilus is found only in the Neotropical Region, from the northernmost areas of South America ( Guyana ) to Argentina and Paraguay (Zilberman 2019). This distribution is directly related to the distribution of termite hosts ( Cornitermes ). The genus Cornitermes contains 14 valid species, although only six of them have been shown to harbor Termitozophilus species. As a result, new Termitozophilus species are expected as nests of Cornitermes species that have not been screened for termitophilous beetles are studied. Termitozophilus laetus is the most widely distributed species, appearing in a variety of biomes, including the Amazon Rainforest, where it is associated with C . incisus Emerson , and the Atlantic Forest, Cerrado (savanna-like vegetation), and southern grasslands, where it is associated with C . cumulans (Kollar) , C . snyderi Emerson , C . bequaerti Emerson and Cornitermes “near villosus ”. The polyxenic condition of T. laetus is likely one of the key reasons for its widespread dispersion. Only the type series collected in the Amazon of Guyana , Termitozophilus mirandus which is associated with C. pugnax and a widely distributed species in the Amazon region, is known. In this sense, as it occurs with its termite host, T. mirandus may be widely distributed in the Amazon. FIGURE 2. Termitozophilus laetus Silvestri, 1901 , female. A) Habitus , dorsal view. B) Abdomen with removed tergites, dorsal view. C) Egg. D) Detail of egg surface. Scales: A–B = 400 µm; C = 80 µm; D = 30 µm. Termitozophilus belleae is only known from the type series collected in the Brazilian Cerrado (savanna-like vegetation) in the state of Mato Grosso , in C. silvestri nests. It is, however, a widely distributed host, found across the Cerrado biome and sections of the Brazilian Amazon Rainforest ( Constantino, 2002 ; Zilberman, 2019). Hence, there is a chance that T . belleae distribution will extend, as additional collecting efforts are made in previously unsampled areas. The four species described in this study were found in Cornitermes nests in pastures in the Brazilian Amazon’s northernmost state of Pará . Remarks. The description of four new Termitozophilus species in this study, as well as a review of existing data from descriptions, photographs, and mounted slides of previously reported species, allowed us to expand the generic diagnosis and redescription. Scanning electron microscope images also aided in the advancement of the morphological research by revealing new features. All Termitozophilus species have articular membranes that are largely exposed between antennomeres II-III, as well as pores across the major sclerotized region of sternites. The previously sparse descriptions of the aedeagus and spermatheca are now expanded to include information regarding chaetotaxy of the paramere, velar sac, median lobe, inner sac sclerites (all aspects of male genitalia), and the morphology of the chamber and duct of the spermatheca. In the internal view, the anteromedial border of the elytra is scaled, a condition initially identified in some Termitomorpha Wasmann species ( Aleocharinae : Corotocini : Termitogastrina) ( Caron et al. 2018 ). The discovery of the same structure in a separate subtribe emphasizes the need for more inquiry as well as a potential hypothesis of primary homology in a larger sense, maybe extending to other Corotocini tribes. The condition of the paratergites, which can be united to sternites or not, demonstrates significant plasticity in the evolution of abdominal sclerites. The shape and arrangement of abdominal sclerites identified in stenogastric specimens may aid in understanding the evolution of abdominal sclerites. Furthermore, the arrangement of paratergites is also useful for species identification.