Diminishing the taxonomic gap in the neotropical soldierless termites: descriptions of four new genera and a new Anoplotermes species (Isoptera, Termitidae, Apicotermitinae).

The neotropical Apicotermitinae is a common and widespread clade of mostly soil-feeding soldierless termites. With few exceptions, species of this group were originally assigned to the genus Anoplotermes Müller, 1873. The application of internal worker morphology coupled with genetic sequencing has recently shed light on the true diversity of this subfamily. Herein, Anoplotermessusanae Scheffrahn, Carrijo & Castro, sp. nov. and four new species in four new genera are described: Hirsutitermeskanzakii Scheffrahn, Carrijo & Castro, gen. nov. et sp. nov., Krecekitermesdaironi Scheffrahn, Carrijo & Castro, gen. nov. et sp. nov., Mangolditermescurveileum Scheffrahn, Carrijo & Castro, gen. nov. et sp. nov., and Ourissotermesgiblinorum Scheffrahn, Carrijo & Castro, gen. nov. et sp. nov. Worker descriptions are based mainly on worker gut morphology, including the enteric valve, while imagoes were described based on external characters. A Bayesian phylogenetic tree of New World Apicotermitinae was constructed using the complete mitogenome to infer genera relationships and corroborate the taxonomic decisions. Distribution maps and a dichotomic key to the known Neotropical Apicotermitinae genera are provided.

First record of the major termite pest species, Reticulitermes flavipes (Isoptera: Rhinotermitidae), in Argentina.

Reticulitermes flavipes is the most invasive species in its genus and is responsible for causing significant damage to human structures in areas where it has been introduced. Although it has already become established in Chile and Uruguay, it had not previously been reported in Argentina. In this study, we report the first detection of this species in Buenos Aires, the capital of Argentina. The colony was already producing alates, and species identification was confirmed through both morphology and mitochondrial gene 16S rRNA analysis. Our results, while not conclusive, suggest that this introduction was independent from the one that occurred in Chile and Uruguay, and potentially originated from the United States. The detection of R. flavipes in Argentina is significant because it highlights the potential for this species to establish itself in new regions and underscores the need for future research on and control of R. flavipes in this country.

Chasitermespax, a new genus and species of soldierless termite (Termitidae, Apicotermitinae) from the island of Trinidad.

Chasitermespax Scheffrahn & Carrijo gen. et sp. nov. is described from workers collected from a single colony in the Northern Range of Trinidad. The shape and texture of the unsclerotized enteric valve, tubular shape of the enteric valve seating, and prominent spherical mesenteric tongue of C.pax are the diagnostic characters for both the genus and species. A Bayesian phylogenetic analysis using the COI gene and including all neotropical Apicotermitinae genera described to date supports the new genus as a distinct terminal.

Can Rubber Crop Systems Recover Termite Diversity in Previously Degraded Pastures in the Colombian Amazon Region?

Livestock production extension in Amazon has caused deforestation and soil degradation, with negative consequences on diversity and environmental services. Recently, rubber crops have been established in deteriorated soils of the Colombian Amazon as an option to restore hectares of unproductive degraded pastures. Bioindicator insects, such as termites, have been used to assess soil quality and fertility restoration. This study evaluated differences in termite abundance, species richness, and community composition in three different rubber crop systems as an indirect way of evaluating soil diversity restoring. Three rubber crop systems were sampled: clonal fields (rubber monocultures with different rubber clones), traditional commercial rubber plantations (rubber monocultures with just one rubber clone), and mixed plantations (rubber fields intercropped with copoazú fruit trees). Additionally, pastures in use for livestock production and natural forest relicts were compared to rubber crop systems, to serve as reference habitats. Termites were sampled using a 105-m transect method. Alpha diversity and beta diversity were estimated and compared between rubber crops and reference habitats. A total of 80 termite species belonging to two families were collected. Mixed plantations and pastures presented the lowest diversity rates. Species richness in rubber crop systems was 39% higher than that in pastures and included 72% of the termite species found in natural forests. Indicator species analysis associated soil-feeding termites with less diverse habitats and wood-feeding termites with high diverse habitats. Our results demonstrate that termite recovery will depend on the farming system selected and the agricultural practices implemented in the field, with some rubber crop systems, like commercial rubber plantations and clonal fields, recovering termite diversity better than others, such as mixed plantations.

Three new species of Rugitermes (Isoptera, Kalotermitidae) from Peru and Bolivia.

The soldier of Rugitermes aridus sp. nov. is described from a xeric, termite-depauperate region of central Peru. Rugitermes rufus sp. nov. and R. volcanensis sp. nov. are described from soldiers and dealated imagos collected in a mesic forest of Amboró National Park in western Bolivia. The imago of R. rufus is unique among all described Rugitermes species in that the head capsule is reddish orange and the pronotum is brown. The imago head and pronotum are both brown in R. volcanensis. A phylogenetic and GMYC barcode analyses were performed with the COI gene. These analyses confirmed the three new species and revealed a high undescribed diversity of Rugitermes in the New World.

Corrigendum: Echinotermes biriba, a new genus and species of soldierless termite from the Colombian and Peruvian Amazon (Termitidae, Apicotermitinae). ZooKeys 748: 21-30. https://doi.org/10.3897/zookeys.748.24253.

[This corrects the article DOI: 10.3897/zookeys.748.24253.].

Rustitermes boteroi, a new genus and species of soldierless termites (Blattodea, Isoptera, Apicotermitinae) from South America.

We present the description of a new genus and species of soldierless termites from South America. Rustitermes boteroi Constantini, Castro & Scheffrahn, gen. et sp. nov. can be identified by the morphology of the enteric valve, with six slightly asymmetric cushions, each one forming a central pouch made of scales smaller than those between the cushions. The new genus features two characteristic rows of thick bristles on the interior margin of the fore tibia, and is supported by COI molecular sequence data. This species is distributed from Tobago to northern Argentina.

Microbial Communities of the Gut and Nest of the Humus- and Litter-Feeding Termite Procornitermes araujoi (Syntermitinae).

The evolution of the symbiotic association with microbes allowed termites to decompose ingested lignocellulose from plant-derived substrates, including herbivore dung and soil humus. Representatives of the Syntermitinae (Termitidae) range in their feeding habits from wood and litter-feeding to humus-feeding species. However, only limited information is available about their feeding ecology and associated microbial communities. Here we conducted a study of the microbial communities associated to the termite Procornitermes araujoi using Illumina sequencing of the 16S and ITS rRNA genes. This species has been previously included in different feeding guilds. However, most aspects of its feeding ecology are unknown, especially those associated to its symbiotic microbiota. Our results showed that the microbial communities of termite guts and nest substrates of P. araujoi differed significantly for bacteria and fungi. Firmicutes dominated the bacterial gut community of both workers and soldiers, whereas Actinobacteria was found in higher prevalence in the nest walls. Sordariomycetes was the most abundant fungal class in both gut and nest samples and distinguish P. araujoi from the grass/litter feeding Cornitermes cumulans. Our results also showed that diversity of gut bacteria were higher in P. araujoi and Silvestritermes euamignathus than in the grass/litter feeders (C. cumulans and Syntermes dirus), that could indicate an adaptation of the microbial community of polyphagous termites to the higher complexity of their diets.

Echinotermes biriba, a new genus and species of soldierless termite from the Colombian and Peruvian Amazon (Termitidae, Apicotermitinae).

A new Apicotermitinae genus and species Echinotermes biriba is described from workers collected on the Andean-Amazon Piedmont in Colombia and Peru. The enteric valve armature of Echinotermes biriba Castro & Scheffrahn, gen. et sp. n. is a remarkably diagnostic character. A Bayesian phylogenetic analysis using the COI gene and including all other Neotropical Apicotermitinae genera, supports the new genus as a distinct terminal.

Tonsuritermes, a new soldierless termite genus and two new species from South America (Blattaria: Isoptera: Termitidae: Apicotermitinae).

A new genus, Tonsuritermes Cancello Constantini gen. nov., is described from South America. The main morphological features of the new genus are a remarkable frontal gland and protibia with two rows of spine-like bristles. Two new species of Tonsuritermes are described: T. tucki Cancello Constantini sp. nov. and T. mathewsi Cancello Constantini sp. nov. Comparisons, measurements, a map, histology of the frontal gland, and illustration of all fundamental morphological aspects are provided.

Phylogeography of Nasutitermes corniger (Isoptera: Termitidae) in the Neotropical Region.

BACKGROUND: The Neotropical Region is known for its biodiversity and ranks third in number of known termite species. However, biogeographic and phylogeographic information of termites of this region is limited compared to other world geographic regions. Nasutitermes corniger is widely distributed in the region and is of considerable economic importance. The goal of this study was to describe the phylogeography of N. corniger in the Neotropical Region, to better understand its evolutionary processes. RESULTS: The sampled populations of N. corniger showed high genetic variation. Results indicated strong geographic structure among N. corniger populations, with most haplotypes not broadly shared among separated locations. Phylogeographic analyses showed a dispersal route for N. corniger from Central America into South America via the Isthmus of Panama, with subsequent dispersal through the highlands east of the Andes and into eastern regions of the continent. The majority of haplotypes were limited in distribution to proximal regions, corresponding to particular biomes (Atlantic Forest, Amazonia, Chaco, Cerrado and Caatinga). CONCLUSIONS: Nasutitermes corniger is suggested to be a good model for biogeographic and phylogeographic studies in the Neotropical Region. This study clarified the phylogeographic history of N. corniger and can contribute to the understanding of biogeographic dispersion processes in the Neotropical Region.

Disjunctitermes insularis, a new soldierless termite genus and species (Isoptera, Termitidae, Apicotermitinae) from Guadeloupe and Peru.

Disjunctitermes insularisgen. n. & sp. n. is described from workers collected on Guadeloupe and in Peru and is the first soldierless termite found on a deep-water island. As with many soldierless and soil-feeding termite species, the enteric valve morphology is an essential diagnostic character of D. insularis. The D. insularis sequence cluster, derived from a barcode analysis with twelve other described genera of New World Apicotermitinae, is well resolved. Results of a stochastic dynamic spread model suggest that the occurrence of D. insularis on Guadeloupe may be the result of a pre-Colombian overwater dispersal event from mainland South America.

Sandsitermes gen. nov., a new nasute termite genus from South America (Isoptera, Termitidae, Nasutitermitinae).

We establish herein a new genus of Neotropical termites 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.

Uncitermes almeriae, a new termite species from Amazonia (Isoptera, Termitidae, Syntermitinae).

The Neotropical termite genus Uncitermes Rocha & Cancello, 2012 was known from a single species, Uncitermes teevani (Emerson, 1925). In this paper a new species, Uncitermes almeriae sp. n., is described and illustrated from worker and soldier castes, along with observations on the Uncitermes nest. A distribution map with the occurrences of both species is presented. The new species is distinguished from its congener by the presence of short bristles covering the head capsule and frontal tube.

The nasus gland: a new gland in soldiers of Angularitermes (Termitidae, Nasutitermitinae).

Termites have developed many exocrine glands, generally dedicated to defence or communication. Although a few of these glands occur in all termite species, or represent synapomorphies of larger clades, others are morphological innovations of a single species, or a few related species. Here, we describe the nasus gland, a new gland occurring at the base of the nasus of Angularitermes soldiers. The nasus gland is composed of class 1, 2, and 3 secretory cells, a rare combination that is only shared by the sternal and tergal glands of some termites and cockroaches. The ultrastructural observations suggest that the secretion is produced by class 2 and 3 secretory cells, and released mostly by class 3 cells. The base of the nasus has a rough appearance due to numerous pits bearing openings of canals conducting the secretion from class 3 secretory cells to the exterior. We tentatively assign a defensive function to the nasus gland, although further research is needed to confirm this function. Although the gland is described only from species of Angularitermes, other genera of Nasutitermitinae also present a rough nasus base, suggesting the presence of a similar, possibly homologous, gland.

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).

Food preference of giant anteater and collared anteater (Pilosa, Myrmecophagidae) regarding the termite defense strategies / Preferência alimentar de Tamanduá Bandeira e de Tamanduá Mirim (Pilosa, Mirmecophagidae) em relação à estratégia de defesa dos cupins

Giant (Myrmecophaga tridactyla) and collared anteaters (Tamandua tetradactyla) are common mammals in the Cerrado biome. They are specialized in eating termites (Isoptera, Blattaria) and ants (Formicidae, Hymenoptera). This study tested the preference of the giant anteater for termites with different defense strategies: 1) soldier with chemical defense and a soft nest (Nasutitermes), and 2) soldier with mixed defenses - chemical and mechanical - and a hard nest (Cornitermes). Pieces of nests of both genera of termites were provided to captive giant anteaters, their behaviors were observed, and the time spent feeding in each termite nests was recorded. The anteaters exploited both termite species, although no significance difference was found, they spent more time feeding on Cornitermes than on Nasutitermes. The stomach contents of one road-killed giant anteater and one collared anteater were analyzed. The collared anteater fed on a wider diversity of termite species with different defense strategies, but showed a preference for Cornitermes. We argue that the preference of anteaters for a termite species that has a harder nest, and soldiers with mixed defense, may be due to the presence of terpenoids in the chemical apparatus of Nasutitermes, absent in Cornitermes. Also, the much higher proportion of soldiers in Nasutitermes may influence the anteaters' choice.

Tamanduá-bandeira (Myrmecophaga tridactyla) e Tamanduá-mirim (Tamandua tetradactyla) são mamíferos comuns no bioma Cerrado. São espécies especializadas em se alimentar de cupins (Isoptera, Blattaria) e de formigas (Formicidae, Hymenoptera). Este trabalho testou a preferência de tamanduás-bandeira por cupins com diferentes estratégias de defesa: 1) soldados com defesa química e ninho macio (Nasutitermes) e 2) soldados com defesa mista ­ química e mecânica ­ e ninho duro (Cornitermes). Pedaços de ninhos de ambos os gêneros de cupins foram fornecidos para tamanduás-bandeira em cativeiro, seu comportamento foi observado e o tempo gasto para se alimentar em cada ninho foi registrado. Os tamanduás exploraram os dois tipos de ninhos, apesar de não haver diferença significativa, eles permaneceram mais tempo se alimentando de Cornitermes do que de Nasutitermes. Os estômagos de um tamanduá bandeira e de um tamanduá-mirim atropelados em rodovias foram utilizados para analisar a dieta das espécies. O conteúdo estomacal do tamanduá-mirim revelou uma maior diversidade de espécies de cupins com diferentes estratégias de defesa, mas mostrou uma preferência por Cornitermes. Discutimos que a preferência de tamanduás por espécies de cupins com soldados de defesa mista e que constroem ninhos duros, pode ser devido à presença de terpenos no sistema de defesa de Nasutitermes, ausente em Cornitermes. Além disso, uma proporção superior de soldados em Nasutitermes pode influenciar a escolha dos tamanduás.

Nest plasticity of Cornitermes silvestrii (Isoptera, Termitidae, Syntermitinae) in response to flood pulse in the Pantanal, Mato Grosso, Brazil

Nest plasticity of Cornitermes silvestrii (Isoptera, Termitidae, Syntermitinae) in response to flood pulse in the Pantanal, Mato Grosso, Brazil. The Pantanal is one of the largest wetlands in the world. Since many areas in Pantanal are flooded during part of the year, it is expected that plants and animals would have mechanisms for their survival during the flooded period. This study investigated the existence of differences in nest shape and inquilines of Cornitermes silvestrii in areas influenced by the flood pulse. We measured the volume, height, width, and height/width ratio of 32 nests in flooded areas and 27 in dry areas, and performed an one-way-Anova with the quasi-Poisson distribution to determine if there were differences in the nest measurements between the points. To analyze the relationship of nest inquilines to flood pulse and nest shape, we performed a regression with a Poisson distribution with the inquiline richness and flood pulse, and the above measurements. The nests of C. silvestrii in flooded areas were significantly higher than nests in dry areas, and had a larger height/width ratio. Colonies in periodically flooded areas would probably make a larger effort to extend their nests vertically, to maintain at least some portion of the structure out of the water and prevent the entire colony from being submerged. Neither the size of the nest nor the flood pulses influenced the assemblage of 11 species found in nests of C. silvestrii.