Revised systematic position of Nasutitermes brevipilus Emerson, 1925 (Isoptera: Termitidae: Nasutitermitinae) and the designation of Hyleotermes gen. nov.

A new monotypic nasute termite genus, Hyleotermes gen. nov., is proposed for Nasutitermes brevipilus Emerson, 1925. Hyleotermes brevipilus, comb. nov., is redescribed and illustrated based on the morphology of the imago, soldier, and worker castes. It is expanded into Amazonia. The soldier of Hyleotermes differs from that of Nasutitermes Dudley, 1890 in that the former has a long and cylindrical nasus and the head capsule lacks long setae and is covered with microscopic setae. Unlike the worker of Nasutitermes, the Hyleotermes worker has a short mixed segment and an enteric valve is adorned with narrow spines on conical bases. The phylogenetic position of H. brevipilus comb nov., is reconstructed based on a dataset with two mitochondrial markers (COI and 16SrRNA) for 36 terminals, under maximum likelihood and Bayesian inference. Results corroborate that this species is unrelated to Nasutitermes and should be excluded from the genus.

First record of Ereymatermes rotundiceps (Termitidae: Nasutitermitinae) as inquiline of Embiratermes neotenicus (Termitidae: Syntermitinae) in the Colombian Amazon / Primer registro de Ereymatermes rotundiceps (Termitidae: Nasutitermitinae) como inquilino de Embiratermes neotenicus (Termitidae: Syntermitinae) en la Amazonia Colombiana

We report, for the first time, the presence of the termite Ereymatermes rotundiceps Constantino, 1991 as an inquiline of the termite Embiratermes neotenicus (Holmgren, 1906) in the Colombian Amazon, and provide new morphometric information on the species. Soldiers of E. rotundiceps were collected in epigeal nests built by E. neotenicus in pastures during the transition from the dry to the rainy season. The association between host and inquiline termites does not seem to be antagonistic, but the relationship between these two species still needs to be clarified.(AU)

Reportamos por primera vez la presencia de la termita Ereymatermes rotundiceps Constantino, 1991 como inquilino de Embiratermes neotenicus (Holmgren, 1906) en la Amazonia Colombiana y aportamos nueva información morfométrica para las especies. Soldados de E. rotundiceps fueron recolectados en nidos epigeos construidos por E. neotenicus en pasturas durante la transición de la estación seca a la lluviosa. La asociación entre las termitas hospederas e inquilinas parece no ser antagónica, sin embargo, las relaciones entre estas dos especies están aún por esclarecer.(AU)

Osteophagia by Nasutitermes guayanae (Blattodea:Isoptera: Termitidae) on human bone remains in theAndean Amazon, Caquetá, Colombia / Osteofagia por Nasutitermes guayanae (Blattodea, Isoptera, Termitidae)sobre restos óseos humanos en la Amazonia Andina, Caquetá, Colombia

We present the first report of Nasutitermes guayanae feeding on human bone remains found in an urban area of the municipality of Florencia, Caquetá, Colombia, in the Colombian Amazon piedmont. The record indicates an expansion in the diet of these termites. The observation suggests that the association of N. guayanae with decomposing bodies may be a possible tool for the estimation of postmortem intervals.

Presentamos el primer reporte de Nasutitermes guayanae alimentándose de restos óseos humanos encontrados en un área urbana del município de Florencia, Caquetá, Colombia, en el Piedemonte Amazónico colombiano. El registro indica una expasión en la dieta de esas térmitas. La observación sugiere que la asociación de N. guayanae con cuerpos en descomposición puede ser una posible herramienta para la estimación de intervalos postmortem.

Osteophagia by Nasutitermes guayanae (Blattodea:Isoptera: Termitidae) on human bone remains in theAndean Amazon, Caquetá, Colombia / Osteofagia por Nasutitermes guayanae (Blattodea, Isoptera, Termitidae)sobre restos óseos humanos en la Amazonia Andina, Caquetá, Colombia

We present the first report of Nasutitermes guayanae feeding on human bone remains found in an urban area of the municipality of Florencia, Caquetá, Colombia, in the Colombian Amazon piedmont. The record indicates an expansion in the diet of these termites. The observation suggests that the association of N. guayanae with decomposing bodies may be a possible tool for the estimation of postmortem intervals.(AU)

Presentamos el primer reporte de Nasutitermes guayanae alimentándose de restos óseos humanos encontrados en un área urbana del município de Florencia, Caquetá, Colombia, en el Piedemonte Amazónico colombiano. El registro indica una expasión en la dieta de esas térmitas. La observación sugiere que la asociación de N. guayanae con cuerpos en descomposición puede ser una posible herramienta para la estimación de intervalos postmortem.(AU)

First description of the worker of Planicapritermes longilabrum with notes on the nest of Planicapritermes planiceps (Blattaria: Isoptera: Termitidae) / Primeira descrição dos operários de Planicapritermes longilabrum, com notas sobre o ninho de Planicapritermes planiceps (Blattaria: Isoptera: Termitidae)

Planicapritermes Emerson, 1949 is a South American termite genus with two nominal species: Planicapritermes planiceps (Emerson, 1925) and Planicapritermes longilabrum Constantino, 1998. The soldiers of this genus are characterized by strongly asymmetrical mandibles. Planicapritermes planiceps was described from soldiers and workers, and P. longilabrum only from soldiers. Here we describe and illustrate workers of P. longilabrum based on specimens collected in Rondônia state, Brazil. We also present a comparison between workers and soldiers of P. planiceps and P. longilabrum, and describe the nest of P. planiceps.(AU)

Planicapritermes Emerson, 1949 é um gênero de cupins da América do Sul com duas espécies nominais: Planicapritermes planiceps (Emerson, 1925) e Planicapritermes longilabrum Constantino, 1998. Os soldados deste gênero são caracterizados por ter mandíbulas fortemente assimétricas. Planicapritermes planiceps foi descrita com base em soldados e operários, e P. longilabrum somente em soldados. Aqui nós descrevemos e ilustramos os operários de P. longilabrum com base em espécimes coletados em Rondônia, Brasil. Também comparamos operários e soldados de P. planiceps e P. longilabrum, e descrevemos o ninho de P. planiceps.(AU)

Comparative anatomy of the gut of the South American species of Amitermes, with description of two new species and an identification key based on soldiers and workers.

In this contribution we present updates on the taxonomy and morphology of the South American species of Amitermes. Two new species are described: Amitermes bandeirai, sp. n., from Brazil, and Amitermes lilloi, sp. n., from Argentina. Amitermes nordestinus is a junior synonym of Amitermes aporema. The imago of A. aporema is described for the first time. Detailed comparative gut anatomy of the eight species is presented for the first time. The geographic distribution of Amitermes in South America is expanded and the distribution patterns of some species are discussed.

Development and comparative morphology of the reproductive system in different aged males of the drywood termite Cryptotermes brevis (Blattaria, Isoptera, Kalotermitidae).

Termites are eusocial cockroaches, which have received great attention due to their diversity of reproductive strategies. Although these novelties allow new interpretations concerning the mating biology of these insects, studies highlighting the structure of the reproductive system are limited to some termite lineages. Here we provide the first comparative analysis of the reproductive system of a drywood termite, using different aged males of Cryptotermes brevis as models. This species represents an important structural pest in tropical regions, and most aspects of its reproductive biology remain unknown, especially on males. The reproductive apparatus of C. brevis is equipped with paired testes, composed of seven testicular lobes, in which developing spermatozoa are located. The basal portion of the lobes connects to the vasa deferentia and transport spermatozoa to a pair of enlarged chambers, the seminal vesicles. These structures join in a median ejaculatory duct, which opens to the external region through a retractile penis. Spermatozoa were observed in all C. brevis males, exhibiting elongated morphology and measuring about 10 µm in length/4 µm in width. Compared with last-instar nymphs and alates, functional kings showed enlarged testes and seminal vesicles, as well as an intense secretory activity towards the lumen of the latter structures. Histochemical tests evidenced strongly PAS and xylidine Ponceau positive reactions of the secretion only in functional kings, indicating the occurrence of glycoproteins. Thus, we suggest that morphophysiological changes establish during the maturation of the reproductive system in C. brevis.

Drywood Pest Termite Cryptotermes brevis (Blattaria: Isoptera: Kalotermitidae): a Detailed Morphological Study of Pseudergates.

The Kalotermitidae Cryptotermes brevis (Walker) presents colonies that lack a true worker caste. They have totipotent worker-like individuals named pseudergates. Few studies have characterized the morphology of immature instars, including pseudergates. In order to identify these instars and characterize the pseudergates, we conducted a comparison between morphometric and morphological variations among immature individuals of C. brevis colonies. Juvenile hormone analog (JHA) was used in the first instar nymphs to induce regressive molts and compare morphological differences between nymphs and pseudergates. Results showed the existence of three larval instars and four nymphal instars. These immatures were morphologically characterized. Individuals classified as third instar larvae presented white body, 10 to 12 antennal articles, absent or small non-pigmented compound eyes, and absence of wing buds. Pseudergates presented pigmented abdomen and sclerotized cuticle, 10 to 12 antennal articles, and absent or small compound eyes, and few specimens had large pigmented compound eyes and absence of wing buds. First instar nymphs had pigmented abdomen and sclerotized cuticle, 10 to 12 antennal articles, both large non-pigmented and pigmented compound eyes, the presence of wing buds. Bioassays using JHA on first instar nymphs resulted in a large percentage of nymph-soldier intercastes. We concluded that abdomen pigmentation and sclerotized cuticle are good characters to differentiate pseudergates from larvae and the absence of wing buds is a good character to differentiate pseudergates from nymphs. Our findings not only contribute to the basic biological and morphological information of this species but also help to identify correctly pseudergates in further studies that involve applied bioassays.

A new termite species of the genus Dihoplotermes Araújo (Blattaria, Isoptera, Termitidae) from the Brazilian Amazonian rainforest / Uma nova espécie de térmita do gênero Dihoplotermes Araujo (Blattaria, Isoptera, Termitidae) da floresta Amazônica Brasileira

Dihoplotermes, a previously monotypic genus, is now composed of two species. Dihoplotermes taurus sp. nov., a new Termitinae termite species, is illustrated and described based on morphological characters from soldiers, workers, and imago. The new species is distinguished from Dihoplotermes inusitatus Araujo by having a spine-like protuberance with blunt apex on the postmentum, the presence of a spine on the gizzard and the absence of dimorphic soldiers. Furthermore, the internal morphological characteristics of a Dihoplotermes worker are described for the first time. Both internal and external morphological characters were used to improve the characterization of the species.(AU)

Dihoplotermes, que era um gênero monotípico, agora é composto por duas espécies. Dihoplotermes taurus sp. nov., uma nova espécie de cupim Termitinae, é descrita e ilustrada com base nos caracteres morfológicos dos soldados, operários e imago. A nova espécie se diferencia de Dihoplotermes inusitatus Araujo por apresentar uma protuberância em forma de espinho não pontiagudo no posmento, presença de um espinho na moela e ausência de soldados dimórficos. Além disso, são descritas pela primeira vez as características morfológicas internas de um operário de Dihoplotermes e utilizamos ambos caracteres morfológicos internos e externos para melhor caracterizar as espécies.(AU)

Hypopharynx in termites: Morphological and functional aspects.

The hypopharynx morphology in workers, pseudergates, and soldiers was described in six families of Isoptera using scanning electronic microscopy. This is the first comparative study to detail the hypopharynx among different species of termites, and included 4 one-piece type (OP) and 9 multiple-pieces type termites (MP). The results showed different arrangements of the hypopharynx trichomes, with fewer of these structures in the frontal region of the hypopharynx in pseudergates and soldiers of OP termites. These characteristics did not apply to Serritermes serrifer, since its pseudergates displayed a large quantity of trichomes on the frontal surface. This species may not resemble other OP termites because it does not live inside a piece of wood, but rather are inquilines inside Cornitermes spp. nests. Similar morphology of this mouthpart was observed in the different species of MP termites, with workers exhibiting many unidirectional trichomes on the frontal surface and soldiers presenting fewer of these trichomes. The morphology of the hypopharynx seems to reflect the lifestyle of these species, because MP termites are able to exploit external water sources. Since different arrangements of hypopharynx trichomes are used for water intake via capillary action, the present data reinforce the soldiers' dependence on workers to obtain water and food through the process of trophallaxis. Three different types of sensilla were observed on the lateral region of the hypopharynx: chaetica, basiconica and campaniformia. Chaetica act as chemoreceptors, while basiconica and campaniformia are mechanoreceptors, used in feeding and environmental perception. Better understanding of hypopharynx morphology permits the inference of ecological habits among OP and MP termites, and reflects the physiological differences between workers and soldiers.

Muelleritermes: A new termite genus with two species from the Brazilian Atlantic Forest (Isoptera: Termitidae: Nasutitermitinae).

We present the description of Muelleritermes, new genus, and two new species: M. fritzi, sp. n. and M. globiceps, sp. n. Both species were found only in the Atlantic Forest of southern Brazil. All castes are described and illustrated, and a distribution map is provided. These species seem to be closely related to the genera Velocitermes and Diversitermes, sharing traits such as the presence of three types of soldiers and workers and a short mixed segment. This genus differs from Velocitermes and Diversitermes in the presence of a few ommatids on soldier's head, behind the antennae. It also differs from Velocitermes in the lack of a constriction on the head of major soldiers, and from Diversitermes in the presence of short hairs on top of the soldier's head, instead of microscopic ones.

A taxonomic revision of the neotropical termite genus Ruptitermes (Isoptera, Termitidae, Apicotermitinae).

The taxonomy of Ruptitermes Mathews 1977 is revised. Nine new species are described: R. araujoi, sp. n. (from Brazil), R. atyra, sp. n. (from Brazil, Peru, and Panama), R. bandeirai, sp. n. (from Brazil), R. cangua, sp. n. (from Brazil), R. kaapora, sp. n. (from Paraguay and Brazil), R. krishnai, sp. n. (from Trinidad), R. maraca, sp. n. (from Brazil), R. piliceps, sp. n. (from Brazil), and R. pitan, sp. n. (from Brazil). Ruptitermes franciscoi (Snyder 1959), new combination, is transferred from Anoplotermes Fr. Müller to Ruptitermes. Ruptitermes proratus Emerson 1949 is placed under the synonymy of R. reconditus (Silvestri, 1901). The genus is redescribed to accommodate all the species included. Ruptitermes arboreus, R. reconditus and R. xanthochiton are redescribed based on the examination of large series. A key to the 13 species of the genus based on the external morphology of workers is provided. The enteric valve armature is described and illustrated for all species except R. kaapora. Distribution maps are presented for all species.

Compositermes bani sp.n. (Isoptera, Termitidae, Apicotermitinae), a new species of soldierless termite from Bolivia.

Species of neotropical Apicotermitinae (Termitidae) are soldierless, restricting species identification in this group to workers or seasonally present winged imagos. All neotropical Apicotermitinae were placed in the genus Anoplotermes, until Mathews (1977) described two new genera, Grigiotermes and Ruptitermes. Fontes (1986) described two more genera, Aparatermes and Tetimatermes. Twenty years passed without any taxonomic advances in this group until Scheffrahn et al. 2006 described two new Anoplotermes species from the West Indies and Bourguignon et al. 2010 described Longustitermes manni, (=Anoplotermes manni Snyder), Anoplotermes janus, and placed six other species into synonymy. Finally, Scheffrahn 2013 described a new genus and species, Compositermes vindai, characterized by a ring of sclerotized paddles at the junction of the enteric valve seating (EVS) and paunch (P3).

Incisitermes nishimurai, a new drywood termite species (Isoptera: Kalotermitidae) from the highlands of Central America.

Incisitermes nishimurai is described from soldiers and winged imagos collected in the highlands of Honduras. It is distinguished from all other Incisitermes species by the dark rugosity of the soldier frons and by the rather large, uniformly very dark, imago.

A taxonomic revision of the soil-feeding termite genus Anhangatermes (Isoptera: Termitidae: Nasutitermitinae).

The taxonomy of Anhangatermes Constantino 1990 is revised. Four new species are described: Anhangatermes anhanguera, sp. n., Anhangatermes eurycephalus, sp. n., Anhangatermes juruena, sp. n., and Anhangatermes pilosus, sp. n. The imago of Anhangatermes is described for the first time (A. pilosus). A key to the species of the genus based on soldiers and workers is provided. The diagnosis of the genus is updated to accommodate the new species. The geographic range of the genus is expanded to the Cerrado region of Central Brazil. The gizzard armature of the worker of Anhangatermes is described in detail for the first time.

Delineating species boundaries using an iterative taxonomic approach: the case of soldierless termites (Isoptera, Termitidae, Apicotermitinae).

Species boundaries are traditionally inferred using morphological characters, although morphology sometimes fails to correctly delineate species. To overcome this limitation, researchers have widely taken advantage of alternative methods such as DNA barcoding or analysis of cuticular hydrocarbons (CHs) profiles, but rarely use them simultaneously in an iterative taxonomic approach. Here, we follow such an approach using morphology, DNA barcoding and CHs profiles to precisely discriminate species of soldierless termites, a diversified clade constituting about one-third of the Neotropical termite species richness, but poorly resolved taxonomically due to the paucity of useful characters. We sampled soldierless termites in various forest types of the Nouragues Nature Reserve, French Guiana. Our results show that morphological species determination generally matches DNA barcoding, which only suggests the existence of three cryptic species in the 31 morphological species. Among them, Longustitermes manni is the only species whose splitting is corroborated by ecological data, other widely distributed species being supported by DNA barcoding. On the contrary, although CHs profiles provide a certain taxonomic signal, they often suggest inconsistent groupings which are not supported by other methods. Overall, our data support DNA barcoding and morphology as two efficient methods to distinguish soldierless termite species.

MRT letter: A novel tegumental gland in female imagoes of the neotropical termite Cornitermes cumulans (Isoptera, termitidae, syntermitinae).

In general, the exocrine glands of social insects are structures involved in the chemical communication associated with social life. Here, we report the discovery of an unknown tegumental gland that is present in the female imagoes of Cornitermes cumulans and occurs next to the well-developed tergal glands that have previously been described. The tegumental glands release their secretion in the intersegmental membrane and are composed of bicellular units, a secretory cell and a canal cell, that are closely located to the epidermal cells in the inferior part of the eighth and ninth tergites. The ultrastructure of the glandular cells showed abundant smooth endoplasmic reticulum, suggesting that the secretion may be pheromonal, although its function is still unknown. These exocrine structures are facing the tergal glands, and we hypothesized that they act synergistically with the tergal glands to generate short-range attraction during tandem behavior.

Pheromones and exocrine glands in Isoptera.

Termites are eusocial insects that have a peculiar and intriguing system of communication using pheromones. The termite pheromones are composed of a blend of chemical substances and they coordinate different social interactions or activities, including foraging, building, mating, defense, and nestmate recognition. Some of these sociochemicals are volatile, spreading in the air, and others are contact pheromones, which are transmitted by trophallaxis and grooming. Among the termite semiochemicals, the most known are alarm, trail, sex pheromones, and hydrocarbons responsible for the recognition of nestmates. The sources of the pheromones are exocrine glands located all over the termite body. The principal exocrine structures considered pheromone-producing glands in Isoptera are the frontal, mandibular, salivary or labial, sternal, and tergal glands. The frontal gland is the source of alarm pheromone and defensive chemicals, but the mandibular secretions have been little studied and their function is not well established in Isoptera. The secretion of salivary glands involves numerous chemical compounds, some of them without pheromonal function. The worker saliva contains a phagostimulating pheromone and probably a building pheromone, while the salivary reservoir of some soldiers contains defensive chemicals. The sternal gland is the only source of trail-following pheromone, whereas sex pheromones are secreted by two glandular sources, the sternal and tergal glands. To date, the termite semiochemicals have indicated that few molecules are involved in their chemical communication, that is, the same compound may be secreted by different glands, different castes and species, and for different functions, depending on the concentration. In addition to the pheromonal parsimony, recent studies also indicate the occurrence of a synergic effect among the compounds involved in the chemical communication of Isoptera.

The frontal gland in workers of Neotropical soldierless termites.

The presence of the frontal gland is well established in termite soldiers of Rhinotermitidae, Serritermitidae, and Termitidae. It is one of their main defensive adaptations or even an exclusive weapon. The gland was also occasionally reported in alate imagoes, but never in the worker caste. Here, we report the first observation of a frontal gland in workers of several Neotropical and one African species of Apicotermitinae. The ultrastructure of Aparatermes cingulatus and Anoplotermes nr. subterraneus is described in detail. In these two species, the gland is well-developed, functional and consists of class 1 secretory cells. The presence of envelope cells, wrapping the gland, is an unusual feature, as well as the presence of several zonulae adherens, connecting neighbouring glandular cells. The frontal gland of workers is homologous to this organ in soldiers and imagoes, as evidenced by the same position in the head and its connection to the same muscle. However, the defensive role of the frontal gland in workers remains to be confirmed.

Anatomy of the frontal gland and ultramorphology of the frontal tube in the soldier caste of species of Nasutitermitinae (Isoptera, Termitidae).

All termite species (Isoptera) are eusocial. These insects live in societies where the division of labor is a predominant characteristic, conditioned by the presence of castes with different morphology, ontogeny, and development. The soldier caste is unique among social insects and it is responsible for colony defense. Soldiers belonging to the Nasutitermitinae subfamily are very peculiar, since they may be polymorphic and present a nasus in addition to either developed or vestigial mandibles. The defensive secretions of soldiers of the neotropical Nasutitermitinae have been the aim of several chemical studies, but few data exist concerning the anatomy and histology of the exocrine glands. This article presents a comparative study on the anatomy of the frontal gland of soldiers of several Nasutitermitinae species: Syntermes dirus (Burmeister), Syntermes nanus (Constantino), Constrictotermes cyphergaster (Silvestri), Nasutitermes corniger (Motschulsky) and Velocitermes heteropterus (Silvestri), with emphasis on the ultramorphology and ultrastructure of the frontal tube.