Zootaxa 4221 (5): 562–574 ISSN 1175-5326 (print edition) http://www.mapress.com/j/zt/ Article ZOOTAXA Copyright © 2017 Magnolia Press ISSN 1175-5334 (online edition) https://doi.org/10.11646/zootaxa.4221.5.5 http://zoobank.org/urn:lsid:zoobank.org:pub:2779DDB4-83EB-49C1-8247-3222D1FE3322 Sandsitermes gen. nov., a new nasute genus from South America (Isoptera, , )

CAROLINA CUEZZO1, ELIANA M. CANCELLO1 & TIAGO F. CARRIJO1,2 1Museu de de Zoologia da Universidade de São Paulo, Av. Nazaré, 481, CEP 04263-000, São Paulo, SP, Brazil. E-mail: [email protected] 2Centro de Ciências Naturais e Humanas, Universidade Federal do ABC, São Bernardo do Campo, SP, Brazil

Abstract

We establish herein a new genus of Neotropical of the subfamily Nasutitermitinae, Sandsitermes gen. nov.. The new genus accommodates a previously described species, Nasutitermes robustus (Holmgren), and is diagnosed by worker characters, including the mandibles, the gut-coiling configuration in situ, and the pattern of cushions and spines of the enteric valve. We characterized and illustrated the imago for the first time and redescribe the soldier and worker castes of Sandsitermes robustus gen. et comb. nov. from syntypes and other samples from eastern Peru. We support our taxonomic decision, presenting morphological differences between the gut pattern and enteric valve of S. robustus and 13 neotropical Nasutitermes species studied for this report, and discuss possible relationships with other neotropical nasute termites.

Key words: gut coiling, enteric valve, Nasutitermes, , termites

Introduction

The termite subfamily Nasutitermitinae comprises 78 genera and 597 species worldwide, including 28 genera and 161 species in the Neotropical region (Krishna et al. 2013; Constantino 2016). Nasutitermitinae is well corroborated as a monophyletic clade (Inward et al. 2007; Legendre et al. 2015), but discerning the relationships among lineages within the subfamily remains a daunting systematic challenge. The type genus of the subfamily, Nasutitermes Dudley, is characterized by a soldier caste with a rather conical nasus and reduced mandibles with conspicuous points (Mathews 1977; Sands 1957, 1965). With more than 250 living species described worldwide and 78 in the Neotropical region, Nasutitermes is the most speciose of all termite genera. Many species build arboreal nests and most of them, probably all, feed on wood. They are extremely abundant in some environments (Cancello et al. 2014; Constantino & Acioli 2006), playing key roles in many ecological processes (Vasconcellos & Moura 2010), and also becoming important urban and agricultural pests (Boulogne et al. 2016; Constantino 2002). Most Neotropical Nasutitermes species were described by Silvestri (1901, 1903), Holmgren (1906, 1910), Emerson (1925) and Snyder (1926). Mathews (1977) also made important contributions to the taxonomy of the genus. However, these descriptions were based mostly on soldier morphology and only seldom on imago morphology, since imagoes are rarely available in collections to help to accurately identify the species. Although Holmgren (1909) provided some insights on termite gut anatomy, internal morphological characters were not incorporated into formal descriptions until the 1950s, when the taxonomic importance of the gut coiling and anatomy was perceived, especially for the family Termitidae (Grassé & Noirot 1955; Kovoor 1969; Johnson 1979; Fontes 1987a; Sands 1998; Noirot 2001). Nowadays, worker gut morphology is commonly included in termite taxonomic descriptions. During a study of the syntypes of Nasutitermes robustus (Holmgren 1906), morphological differences with Nasutitermes s.s. were recognized, such as in the mandibles, gut-coiling pattern, and enteric-val