Resource adaptation drives the size-complexity rule in termites.

The size-complexity rule posits that the evolution of larger cooperative groups should favour more division of labour. Examples include more cell types in larger multicellular organisms, and more polymorphic castes in larger eusocial colonies. However, a correlation between division of labour and group size may reflect a shared response of both traits to resource availability and/or profitability. Here, this possibility was addressed by investigating the evolution of sterile caste number (worker and soldier morphotypes) in termites, a major clade of eusocial insects in which the drivers of caste polymorphism are poorly understood. A novel dataset on 90 termite species was compiled from the published literature. The analysis showed that sterile caste number did increase markedly with colony size. However, after controlling for resource adaptations and phylogeny, there was no evidence for this relationship. Rather, sterile caste number increased with increasing nest-food separation and decreased with soil-feeding, through changes in worker (but not soldier) morphotype number. Further, colony size increased with nest-food separation, thus driving the false correlation between sterile caste number and colony size. These findings support adaptation to higher energy acquisition as key to the rise of complex insect societies, with larger size being a by-product.

Amazonian forest termites: a species checklist from the State of Acre, Brazil / Térmitas da Floresta Amazônica: uma lista de espécies do Estado do Acre, Brasil

Abstract The Brazilian state of Acre is located in the southwestern Amazon and it is characterized by a humid tropical forest vegetation that covers plains and mountains. Up to this point, the composition of termite species in the state is not known. The aim of this study was to provide a checklist of termite species or recognizable taxonomic units for the state of Acre. Sampling was conducted through field expeditions at the Serra do Divisor National Park, Chandless State Park, Humaitá Forest Reserve, and Chico Mendes Environmental Park using a standardized rapid termite inventory protocol in the first two areas and active searching collections in the others, without a specific protocol. This study also included occurrence records published in the scientific literature. A total of 128 species and morphospecies of termites were found in Acre, distributed across 59 genera and four families. The most frequently occurring species in Acre was Heterotermes tenuis (Hagen, 1858). The study also identified six new species records for Brazil. The predominant feeding groups were soil-feeders and wood-feeders, as expected from data obtained from surveys in humid tropical forests. Despite the significant number of new records for Acre (112), it is concluded that a larger sampling effort is still required, as many areas of the state have not yet been studied for termites.

Resumen O estado brasileiro do Acre está localizado no sudoeste da Amazônia e é caracterizado por uma vegetação de floresta tropical úmida que cobre planícies e montanhas. Até então, a composição de espécies de térmitas no estado não é conhecida. O objetivo desse estudo foi construir um checklist de espécies ou unidades taxonômicas reconhecíveis de térmitas para o estado do Acre. A amostragem foi conduzida através de expedições de campo no Parque Nacional da Serra do Divisor, no Parque Estadual Chandless, na Reserva Florestal Humaitá, e no Parque Ambiental Chico Mendes utilizando o protocolo rápido de diversidade de térmitas nas duas primeiras áreas e coletas avulsas nas demais, sem um protocolo específico. Este estudo também incluiu registros de ocorrência publicados na literature científica. Um total de 128 espécies e morfoespécies de térmitas foram encontradas no Acre, distribuídas em 59 gêneros e quatro famílias. A espécie de ocorrência mais frequente no Acre foi Heterotermes tenuis (Hagen, 1858). O estudo também identificou seis novos registros de espécies para o Brasil. Os grupos alimentares predominantes foram os humívoros e xilófagos, como esperado a partir de dados obtidos de pesquisas em florestas tropicais úmidas. Apesar do número significativo de novos registros para o Acre (112), conclui-se que ainda é necessário um esforço amostral maior, uma vez que muitas áreas do estado ainda não foram estudadas para térmitas.

The Impact of Constrictotermes cyphergaster (Termitidae: Nasutitermitinae) Termites on Semiarid Ecosystems in Brazil: A Review of Current Research.

Termites have global distributions and play important roles in most ecosystems, often with high nest densities and interesting associations with other organisms. Constrictotermes cyphergaster, is a termite endemic to South America, widely distributed and very conspicuous, and has therefore been considered a good model for filling in gaps in general termite ecology and their relationships with other organisms (e.g., termitophily). A systematic review (content and bibliometric analyses) was used to gather all published scientific knowledge related to C. cyphergaster as well as to observe trends, verify gaps, and direct new perspectives for future studies of this species. We identified 54 studies, of which more than 50% were published in the last five years (28 articles). The majority of the articles investigated the relationships between C. cyphergaster and macroorganisms (44.4%), followed by specific aspects of its biology (25.9%). The collaboration network revealed that links between researchers are still limited and modular, but trending topics have changed over time. Additionally, there are differences in the aims of the studies being carried out in the Caatinga and Cerrado domains, with some information focusing only on one of those environments. Our results show that some gaps in the biology and ecology of C. cyphergaster remain to be explored, although collaborative efforts between researchers open opportunities for suggesting future studies that would make relevant contributions to the general knowledge of termites.

Schievitermesglobicornis, a new genus and species of Termitinae (Blattodea, Termitidae) from French Guiana.

Asymmetrical snapping mandibles have evolved several times in termites. In the Neotropics, asymmetrical snapping mandibles are found in soldiers of four genera: Neocapritermes, Planicapritermes, Cornicapritermes and Dihoplotermes. Here, I describe Schievitermesglobicornis, new genus and species, from French Guiana. This genus is characterized by an absence of a frontal prominence and slightly asymmetrical mandibles in the soldier caste. The morphology and anatomy of the worker reveal a wood-based diet, and suggest that Schievitermes, Planicapritermes and Neocapritermes constitute a monophyletic group, which is consistent with mtDNA data.

Cryptotermes camelus (Isoptera: Kalotermitidae), a new drywood termite species from the Bolivian Chaco.

Cryptotermes Banks, 1906 is the third most diverse kalotermitid genus worldwide after Glyptotermes Froggatt, 1897 and Neotermes Holmgren, 1911, with its greatest diversity found in the Neotropics (Krishna et al. 2013a). Furthermore, the greatest number of species of Cryptotermes are known from the Caribbean Basin (Scheffrahn Krecek 1999, Casala et al. 2016, Scheffrahn 2019). Although Araujo (1977) and Bacchus (1987) list Cryptotermes domesticus (Haviland, 1898) from Trinidad (treated as mainland) and Panama, respectively, Scheffrahn Krecek (1999) and Scheffrahn et al. (2009) doubt the existence of this Asian species in the New World. Without C. domesticus, the total extant Neotropical diversity of Cryptotermes is 29 endemic and three exotic species (Constantino 2020).

Morphology of the antennal sensilla of two species of Hoplopyga Thomson, 1880 (Coleoptera, Scarabaeidae, Cetoniinae)

ABSTRACT Antennal sensilla are important functional elements of sensory systems in insects. This study aimed to determine the morphology and structure of the sensilla of two species of the genus Hoplopyga. Adults of Hoplopyga liturata (Olivier, 1789) were collected in traps with sugarcane juice as an attractant. Thereafter, larvae of Hoplopyga albiventris (Gory and Percheron, 1833) were collected in mounds of termites (Cornitermes cumulans (Kollar, 1832) Isoptera). Then, they were reared in the laboratory for adult observations. Antennae of H. liturata and H. albiventris have sensilla chaetica, trichodea, placodea (type I and II), coeloconica (type I and II), and ampullacea (or pore). Females of H. liturata have a total of about 10657 sensilla and males have about 12512, whereas females of H. albiventris have about 16490 sensilla and the males 24565 sensilla. Sensilla placodea are predominant in the antenna of males and females of both species.

Termitaphididae, a family of true bugs (Heteroptera) new to Brazil, with the redescription of Termitaradus trinidadensis.

The family Termitaphididae and the species Termitaradus trinidadensis (Morrison) are recorded for the first time from Brazil. This species had been found only in Trinidad and Tobago and is redescribed based on specimens collected in Heterotermes tenuis (Hagen) termite nests from forested and open areas in the Brazilian Amazon. Scanning electron micrographs of integumental ornamentation, illustrations of male and female terminalia, and pictures of live specimens are provided for the first time.

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.

Consumption Rate of Lichens by Constrictotermes cyphergaster (Isoptera): Effects of C, N, and P Contents and Ratios.

Wood is the main dietary item for most termites; however, supplementation with certain nutrients may occur via the ingestion of other available food resources in the ecosystem. The objective of this study was to evaluate the consumption of lichens with different C, N, and P contents by Constrictotermes cyphergaster under laboratory conditions, and estimate the intake of this substrate by this species in a semi-arid area of Northeast Brazil. The foraging activities of fifteen field colonies were monitored over 15 days from 6:00 p.m. to 6:00 a.m., and the lichens that were consumed were identified. Blocks of lichen thallus (1.5 × 1.5 cm) of four lichen species were offered to the termites in the laboratory. The mean total consumption rate of lichen by C. cyphergaster was 0.032 mg lichen/g termite (fresh weight)/day. Dirinaria confluens was the lichen most consumed by termites (0.010 mg lichen/g of termite (fresh weight)/day), followed by Lecanora spp. and Haematomma persoonii at a mean consumption of 0.008 and 0.006 mg lichen/g termite (fresh weight)/day, respectively. Based on the size of the C. cyphergaster populations, the estimated lichen consumption rate was 105.12 g lichen/ha/year. Lichen consumption was significantly affected by the N content and the C:N and C:P ratios, with the N content being the factor that best explained the consumption by the termites. The results suggest that C. cyphergaster can use lichens as a supplemental source of nutrients, especially nutrients that are found in low concentrations in wood.

Population structure of Guyanagaster necrorhizus supports termite dispersal for this enigmatic fungus.

Understanding how species accomplish dispersal of their propagules can shed light on how they are adapted for their ecosystem. Guyanagaster necrorhizus is a sequestrate fungus, meaning its dispersal propagules, or spores, are entirely enclosed within a fruiting body, termed a sporocarp. This fungus is most closely related to Armillaria and its allies. While Armillaria species form mushrooms and have forcibly discharged spores, G. necrorhizus spores have lost this ability, and by necessity, must be passively dispersed. However, G. necrorhizus does not possess characteristics of other sequestrate fungi with known dispersal mechanisms. Repeated observations of termites feeding on G. necrorhizus sporocarps, and spores subsequently adhering to their exoskeletons, led to the hypothesis that termites disperse G. necrorhizus spores. To test this hypothesis, we used microsatellite markers and population genetics analyses to understand patterns of clonality and population structure of G. necrorhizus. While Armillaria individuals can spread vegetatively over large areas, high genotypic diversity in G. necrorhizus populations suggests spores are the primary mode of dispersal. Spatial genetic structure analyses show that G. necrorhizus sporocarps within 238 m of each other are more closely related than would be expected by chance and conservative estimates from population assignment tests suggest gene flow no longer occurs between sporocarps separated by 2 km. These distances are consistent with previous studies analysing foraging distances of the termites found associated with G. necrorhizus sporocarps. Termites have rarely been recorded to specifically target fungal sporocarps, making this a potentially novel fungal-insect interaction.

Reexamination of the identity and status of the termite taxa described by Czerwinski (1901) from southern Brazil (Isoptera: Termitidae).

Czerwinski (1901) (sometimes spelled Tscherwinsky) described four new termite taxa in a publication about the termite collection of the former Zoological Cabinet of the Imperial Warsaw University (currently Museum and Institute of Zoology of the Polish Academy of Sciences, MZPW): Eutermes burmeisteri and Eutermes rippertii iheringi from southern Brazil, Eutermes canariensis from the Canary Islands, and Eutermes nasonovi from Madagascar.

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.

New species of Corotoca Schiødte, 1853 from South America and description of first instar larva.

Corotoca Schiødte, 1853 is a Neotropical genus of termitophile beetles, with five species. The establishment of this genus marks the first record of insects living associated to termites. A new species, Corotoca pseudomelantho Zilberman, sp. nov. from Brazil and Argentina, based on adult and first larval instar, is described and illustrated. The first instar larva of C. melantho Schiødte, 1853, a related species, is redescribed and illustrated. This is the first record of Corotoca species from Argentina.

Tropical wood resistance to the West Indian drywood termite Cryptotermes brevis: If termites can't chew .

BACKGROUND: The importance and impact of invasive species are usually considered based on their economic implications, particularly the direct damage that they cause. The West Indian drywood termite Cryptotermes brevis (Walker) is an example and is a concern in structural lumber, furniture, and other wood products. Despite its importance, its tropical wood preferences and the wood physical characteristics contributing to resistance have not been investigated to date. Here, we developed wood testing units to allow the X-ray recording of termite colonization and then subsequently tested tropical wood resistance to the termite through free-choice and no-choice bioassays using these wood testing units. The relevance of wood density and hardness as determinants of such resistance was also tested, as was termite mandible wear. RESULTS: The wood testing units used allowed the assessment of the termite infestation and wood area loss, enabling subsequent choice bioassays to be performed. While pine (Pinus sp.), jequitiba (Cariniana sp.) and angelim (Hymenolobium petraenum) exhibited the heaviest losses and highest infestations; cumaru (Dipteryx odorata), guariuba (Clarisia racemosa), and purpleheart (Peltogyne sp.) showed the lowest losses and infestations; courbaril (Hymenaea courbaril), eucalyptus (Eucalyptus sp.), and tatajuba (Bagassa guianensis) exhibited intermediary results. CONCLUSION: Wood hardness and in particular wood density were key determinants of wood resistance to the termites, which exhibited lower infestations associated with greater mandible wear when infesting harder high-density wood. © 2017 Society of Chemical Industry.

The Role of Resource Density on Energy Allocation in the Neotropical Termite Nasutitermes aff. coxipoensis (Termitidae: Nasutitermitinae).

Organisms acquire energy from environment and must allocate it among different life traits (growth, maintenance and reproduction). Social insects must manage the energy allocation to various levels such as colony growth and caste functions. Here, we addressed the question of whether resource density affects the energy allocation to the number of individuals and caste functions as well as nest's growth rate in the Neotropical termite Nasutitermes aff. coxipoensis (Homgren) (Termitidae: Nasutitermitinae). In a manipulative field experiment, colonies of N. aff. coxipoensis, with known volume, were maintained in plots with three different resource's density (0.32, 0.64 and 1.92 baits/m2) over 3 months. After this period, the number of individuals as well as the caste identity and nest volume were measured. Surprisingly, our results showed that colonies reared in the extremes of resource's density (0.32 and 1.92 baits/m2) produced a higher number of individuals compared with colonies reared with intermediate resource density (0. 64 baits/m2). The mean number of workers increased linearly with resource density; however, the average number of immature was higher in colonies reared with 0.32 baits/m2 compared with colonies reared with 0.64 and 1.92 baits/m2. No significant differences of resource density were observed in the mean number of soldiers, worker/soldier ratio as well as in the nest's growth rate. In conclusion, the resource's density seems to play an important role in determining the investment of energy in the number of individuals and caste in N. aff. coxipoensis colonies.

Hexapod decomposers of Serra de Santa Catarina, Paraíba, Brazil: an area with high potential for conservation of Caatinga biodiversity / Hexápodes decompositores da Serra de Santa Catarina, Paraíba, Brasil: uma área com elevado potencial para a conservação da biodiversidade da caatinga

Abstract: In detritus-based trophic systems, springtails, dung beetles, saprophagous calyptrate flies and termites consume and fragment organic matter and control populations of decomposer microorganisms, exerting a strong influence on energy and nutrient fluxes. A faunal inventory of these four groups of hexapods was performed in Serra de Santa Catarina (SSC), an area of arboreal-shrub caatinga located in the state of Paraíba, with the purpose of characterizing the assemblages of these taxa, and highlighting their unique links to the local ecosystem. Samplings were performed in May, 2014, and April, 2015, both during the rainy season. Standard sampling protocols for biodiversity inventory of the various taxa were used, including both active and passive sampling methods. In general, 114 species of hexapods were captured, with 26 species of springtails, 20 dung beetles, 30 saprophagous calyptrate flies and 38 termites, with sampling sufficiency varying from 69.5 to 96.8% of total estimated richness. Species richness of the groups are among the highest recorded for a single area of Caatinga, with some taxa being recorded for the first time for the domain. Several morphospecies had indeterminate taxonomic status, especially springtails and termites, and are quite likely new species to science. The structure of the assemblages of springtails, dung beetles, saprophagous calyptrate flies and termites recorded in SSC, suggest that this conserved area is unique within the highly impacted landscape of Caatinga, and has great potential for the conservation of biodiversity of this domain in the Northeast Region of Brazil.

Resumo: No sistema trófico baseado em detritos, os colêmbolos, besouros escarabeíneos, moscas saprófagas e térmitas atuam no consumo e fragmentação da matéria orgânica e no controle das populações de microrganismos decompositores, exercendo forte influência nos fluxos de energia e nutrientes. Um inventário faunístico desses quatro grupos de hexápodes foi realizado na Serra de Santa Catarina (SSC), uma área de Caatinga arbóreo-arbustiva localizada no Estado da Paraiba, com intuito de caracterizar as taxocenoses desses táxons, salientando as suas peculiaridades ligadas ao ecossistema local. As coletas foram realizadas em maio/2014 e abril/2015, durante o período chuvoso na região. De acordo com o táxon, foram utilizados protocolos amostrais padronizados para inventários de biodiversidade, com métodos passivos e ativos de coleta. No geral, 114 espécies de hexápodes foram capturadas, sendo 26 de colêmbolos, 20 de besouros escarabeíneos, 30 de dípteros e 38 de térmitas, com suficiência amostral variando de 69,5 a 96.8% da total estimada. A riqueza de espécies por grupo está entre as maiores para uma única área de Caatinga, com alguns táxons sendo registrados pela primeira vez para o domínio. Especialmente para os colêmbolos e térmitas, várias morfoespécies tiveram seus status taxonômicos indeterminados, havendo elevada possibilidade de serem nova para a ciência. As estruturas e funcionalidades das taxocenoses de colêmbolos, besouros escarabeíneos, moscas saprófagas e térmitas registradas na SSC, sugerem que esse ecossistema é singular, dentro de um cenário generalizado de impacto antrópico presente na Caatinga, e possui um elevado potencial para a conservação da biodiversidade desse domínio no nordeste brasileiro.

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.

Sub-lethal effects of essential oil of Lippia sidoides on drywood termite Cryptotermes brevis (Blattodea: Termitoidea).

The drywood termite Cryptotermes brevis (Walker, 1853) (Kalotermitidae) is one of the most important wood structural pest in the world. Substances from the secondary metabolism of plants (e.g., essential oils) have been considered an environmentally safer form of control for urban pests, such as termites. In the present study, we analyzed the lethal and sub-lethal effects of essential oil of Lippia sidoides and its major components on C. brevis pseudergates in two routes of exposure (contact and fumigation). The essential oil of L. sidoides and thymol were more toxic to C. brevis pseudergates when applied by contact (LD50 = 9.33 and 8.20µgmg-1, respectively) and by fumigation (LC50 = 9.10 and 23.6µLL-1, respectively). In general, treatments changed the individual and collective behaviors of C. brevis pseudergates, as well as the displacement and walking speed. The essential oil of L. sidoides and its major components showed a high potential to control C. brevis pseudergates, due to the bioactivity in the two routes of exposure and the sub-lethal effects on the behavior and walking, important activities for the cohesion of C. brevis colonies.

First forensic records of termite activity on non-fossilized human bones in Brazil / Primeiros relatos forenses de atividade de térmitas em ossos humanos não fossilizados no Brasil

Abstract The aim of this study was to describe the first records of termite activity on non-fossilized human bones in Brazil. The cases reported in this study resulted from forensic analysis of six human skeletons found in northeastern Brazil between 2012 and 2014. Traces of tunnels and nests commonly produced by termites were found on several human bone surfaces as well as the specimens and characteristic signs of osteophagic activity. In four cases, the species were identified: Amitermes amifer Silvestri, 1901, Nasutitermes corniger (Motschulsky, 1855) (on two skeletons), and Microcerotermes indistinctus Mathews, 1977. In two other cases, the activity of termites on bone surfaces was evidenced by remains of nests and tunnels produced by these insects. At least in the samples of human remains available for this report, the number of termites collected was greater on bones found during autumn, the rainy season in the Northeast of Brazil. The human bones examined showed termites like insects with lots of strength at bone degradation, capable of continuing the process of decomposition of human remains even in completely skeletonized bodies.

Resumo O objetivo deste estudo foi descrever os primeiros relatos de atividade de térmitas em ossos humanos não fossilizados, no Brasil. Os casos relatados neste estudo resultaram da análise pericial de seis esqueletos humanos encontrados no Nordeste do Brasil, entre os anos de 2012 e 2014. Vestígios de túneis e ninhos comumente produzidos por cupins foram encontrados em várias superfícies dos ossos humanos, bem como a presença de espécimes e característicos sinais de atividade osteofágica. Em quatro casos, foram identificadas as espécies: Amitermes amifer Silvestri, 1901, Nasutitermes corniger (Motschulsky, 1855) – em duas ossadas, e Microcerotermes indistinctus Mathews, 1977. Em dois outros casos, a atividade de cupins sobre superfícies ósseas foi evidenciada pelos restos de ninhos e túneis produzidos por esses insetos. Pelo menos nas amostras de restos humanos disponíveis para o presente estudo, o número de térmitas recolhido foi maior em ossos encontrados durante o outono, a estação chuvosa no Nordeste do Brasil. Os ossos humanos analisados ​​apresentaram cupins como insetos com muita força na degradação óssea, com capacidade de prosseguir o processo de decomposição de restos humanos, mesmo em corpos completamente esqueletizados.

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.