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The soldierless termites of Africa (Isoptera: Termitidae)

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... The presence of termites in cocoa farms has been documented since the 1950s (Harris, 1954) but their damage to cocoa trees were not documented. The first signs of termite damage on cocoa was reported by Sands (1972) who described them as minor, illustrated by galleries running off the tree. Entwhistle (1972) the same year published the first catalog of major pests in cocoa where termites were not mentioned. ...
... We identified the soldier castes to genus and sometimes to species levels using dichotomous keys of Emerson (1928), Bouillon & Mathot (1965) and Sands (1965Sands ( , 1972Sands ( , 1998, and through the reference collections of Institut de Recherche Agricole pour le Développement (IRAD) Nkolbisson, Yaoundé (Cameroon). Whenever possible, specimens were identified to species level, and when this proved impossible, unidentified species were added the suffix "sp." to the generic names because many species are not easy to be identified with certainty in some genera (Darlington et al., 2008). ...
... The slides were then observed under an Omax microscope at a magnification of 40x. The enteric valve structures were compared to previous features using Sands (1972Sands ( , 1998 or to existing prepared reference slides preparations of enteric valves structures of termite collections stored at IRAD. ...
Thesis
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Termites are increasingly recognized as a major constraint to cocoa production in West and Central Africa. Their attacks and damages in cocoa agroforestry systems and cocoa trees may significantly affect tree survival and yield. Termite control relied almost exclusively on persistent organochlorine insecticides which are currently under restrictive use due to increasing concern over damage to human health and the environment. Previous studies reported the limited effect of some entomopathogenic fungi as biological control agents in the field. In addition, very little is known about the type of termite responsible for damages to cocoa trees or how environmental conditions and land-use management shape their bio-ecology including that of biological control agents such as mites and parasitic flies. This study was therefore undertaken to assess the damaging activities of termites and to screen biological control agents against key termite pests of cocoa in order to develop new control formulation. The first objective of the study focused on identifying and characterizing the different functional groups of termites in cocoa plantations through a sampling of termites on infestation sites of randomly selected cocoa trees. The study then evaluated the type of ecosystem services and disservices that termites provide to cocoa by quantifying termites’ role on soil fertility and estimating the yield gap induced by their damages. The study further isolated, characterized and identified fungi and parasitic mites and flies as potential biological control agents using molecular tools and techniques. This study identified fifty-four termite species associated with cocoa agroforestry systems with the rustic shaded systems recording the highest termite taxonomic and functional diversity (37 species). The reduction of shade trees coupled with increasing temperatures resulted to a significant increase in the number of cocoa trees attacked by termites. The study further found that cocoa trees were infested both aboveground and belowground but shade trees tend to reduce belowground infestation. The Taylor’s power law was the best-fitted model to describe the aggregative dispersion pattern of termites on cocoa trees and help to estimate sampling plans using Green’s sequential sampling. The study provided detailed information on the damage patterns of key genera Microtermes, Microcerotermes, Ancistrotermes, Nasutitermes and Odontotermes in cocoa agroforestry systems. Termite infestations affected tree growth, pod formation and flowering. Termites reduced cocoa yield by about 29% and could reach 55% in case of severe infestation. The study also isolated fifty nine species of potential biological control agents of termites using morphological and molecular tools. They included 26 fungi species, 19 mite species and 14 parasitoid flies. Fungi were mainly Ascomycota species among which Aspergillus spp. were the most frequently isolated fungi. Mites belonged all to Super-Order Acariforme and were xxii mainly composed of acarid species. Parsitoid flies isolated from termites were mainly the family Phoridae and included Megaselia spp. and Melaloncha sp.. A more diversified community of biological control agents was recorded in heavily shaded cocoa agroforestry systems. The study did not isolate any mites on termites collected in full sun cocoa agroforestry systems. This study is the first to document potential damage patterns, sampling plans and biological control agents of termites on cocoa. Results provided could be used by researchers and farmers to assess the severity of termite damages in plantations and to develop IPM programs to manage termite pest species in cocoa agroforestry systems. This study also highlighted the contribution of shade trees in limiting termite outbreaks in cocoa agroforestry systems. The study provided new insight on good candidate biological control agents (Cordyceps tenuipes, Penicillium paxilli, P. citrinum and Fusarium oxysporum, Acotyledon sp. and Melaloncha sp.) that can further be explored in an Attract-and-Kill strategy to manage termites in agricultural lands or in environments where termites are potential threats.
... Soldiers were later discovered in one Speculitermes species (Roonwal & Chhotani, 1966), but remain unknown, and are possibly absent, in several Oriental species. In his revision of African soldierless termites, Sands (1972) expanded the Apicotermitinae subfamily and distinguished two 'branches': (i) the Apicotermes-branch, including Apicotermes and related African soldiered genera; and (ii) the Anoplotermes-branch, including the soldierless taxa, the soldiered genus Firmitermes Sjöstedt, and the Oriental genera Speculitermes, Indotermes and relatives. More recently, based on gut anatomy, Noirot (2001) recognized three 'groups': (i) the Apicotermes-group, including the soldiered African taxa and the soldierless genus Skatitermes Coaton; (ii) the Speculitermes-group, including all Asian Apicotermitinae; and (iii) the Anoplotermes-group, including the African and Neotropical soldierless species plus the soldiered genus Firmitermes. ...
... This makes the Apicotermitinae one of the most diverse subfamilies of Termitidae. It is also the most understudied, probably because many species, being soldierless, can only be distinguished morphologically by tedious and difficult dissections of the worker digestive tract (Grassé & Noirot, 1955;Sands, 1972Sands, , 1998Noirot, 2001;Bourguignon et al., 2016b). The shape of the mesenteron-proctodeum junction forming a mixed segment and the morphologically variable enteric valve armature are the conventional characters used for the Apicotermitinae taxonomy (Grassé & Noirot, 1955;Sands, 1972Sands, , 1998Noirot, 2001). ...
... It is also the most understudied, probably because many species, being soldierless, can only be distinguished morphologically by tedious and difficult dissections of the worker digestive tract (Grassé & Noirot, 1955;Sands, 1972Sands, , 1998Noirot, 2001;Bourguignon et al., 2016b). The shape of the mesenteron-proctodeum junction forming a mixed segment and the morphologically variable enteric valve armature are the conventional characters used for the Apicotermitinae taxonomy (Grassé & Noirot, 1955;Sands, 1972Sands, , 1998Noirot, 2001). The gizzard ornaments are also useful as taxonomic characters, particularly in the African soldierless species (Noirot, 2001;Romero Arias et al., 2020). ...
Article
Soil-feeding termites are abundant in tropical regions and play an important role in soil bioturbation and in the organic matter cycle. The Apicotermitinae are arguably the most diverse lineage of soil-feeding termites, but they are also the most understudied, probably because many species are soldierless, which makes identification difficult. Although the backbone of the termite phylogenetic tree is now well-resolved, the relationships among representatives of Apicotermitinae are still largely unknown. Here, we present phylogenetic trees inferred from 113 mitochondrial genomes of Apicotermitinae representative of the group diversity. Our analyses confirm the monophyly of the Apicotermitinae and the basal position of soldiered taxa, within which two lineages of soldierless species are nested. We describe two new monotypic genera, whose phylogenetic position appeared of special interest: Koutabatermes gen. n., lies on a long branch among soldiered taxa, and Apolemotermes gen. n., is sister to Adaiphrotermes. We resolved, with high support, the position of Asian genera as sister group of a clade comprising the monophyletic neotropical Anoplotermes-group and the small African clade including Adaiphrotermes and Apolemotermes gen. n.. Our trees cast light on the intergeneric and interspecific relationships within Apicotermitinae and reveal the polyphyly of several genera, including Ruptitermes, Astalotermes and Anoplotermes. Biogeographic reconstructions revealed two dispersal events out of Africa, one to the Oriental realm and one to the Neotropical realm. Overall, the timing of Apicotermitinae diversification and dispersal, following the Eocene–Oligocene boundary, matches that found for other groups of Neoisoptera. Nomenclatural acts are registered in ZooBank: http://zoobank.org/urn:lsid:zoobank.org:pub:CA1A21B6-573E-4855-8C88-372453C922F7.
... In soldierless groups, the workers developed other forms of defense, such as the rupture of the thorax exposing the often sticky product of dehiscent organs (Sands 1982;Poiani & Costa-Leonardo 2016) or the intestinal content itself (reflex gastrotomy, Sands 1972; it is important highlight that this kind of mechanisms is not exclusive of workers, see e.g. Dentispicotermes, Serritermes, Orthognatotermes, Neocapritermes), and the depositing of fecal fluids onto antagonistic organisms (Coaton 1971;Grassé 1986). ...
... The terminology used to describe worker mandibles follows Sands (1972) and Deligne (1999), while worker digestive tube descriptions follow Noirot (2001). ...
... In Sands (1972, fig. 2), a variation of the tooth that he calls the 'marginal subsidiary' is represented, which may or may not be hidden by the molar prominence (depending on the position it occupies) and indicates that the state of this tooth is a good generic characteristic. ...
Article
A new genus, Dissimulitermes Constantini & Cancello gen. nov ., is described from the Neotropical region. The main morphological feature of the new genus is remarkable defensive organs. A new species of Dissimulitermes is described: D. invisibilis Constantini & Cancello, sp. nov . Description, comparisons, measurements, a distribution map, histology of their dehiscent organ, and an illustration of all the fundamental morphological aspects are provided.
... The systematics of Apicotermitinae has been mostly based on the anatomy of the digestive tract (Grassé and Noirot 1954;Sands 1972Sands , 1998, and in particular, on the configuration of the mesenteric-proctodeal junction and the armature of the enteric valve (Noirot 2001). The diversity of structures in the digestive tract suggests that species differ in their feeding ecology. ...
... Three species belonging to different genera had soldiers, the remaining 14 species were soldierless. According to Donovan et al.'s (2001a) criteria for feeding group assignment in Apicotermitinae, eight species were characterised by a highly sclerotised enteric valve corresponding to feeding group IV (true soil feeders), while nine species exhibited low sclerotisation corresponding to feeding group III (soil-wood feeders) (Sands 1972(Sands , 1998; see also Online Resource 1). Individuals were kept in 80% ethanol for morphological species identification, gut content extraction and Cr-Gi measurements. ...
... In contrast, the sympatric Hospitalitermes papuanus, which carries balls of collected microepiphytes between its mandibles back to the nest, possesses a normal-sized crop (Roisin and Pasteels 1996). The nesting habits of Acholotermes sp03 and Ateuchotermes retifaciens are unknown, but Acidnotermes praus is the sole soldierless species in our sample to build an epigeous nest (Sands 1972). A voluminous crop might therefore be useful to limit the time spent foraging outside the safety of the nest. ...
Article
Termites are dominant organisms of tropical ecosystems. Their success is partly due to the diversity of their feeding substrates, from dead plant tissues to mineralised soils. The Apicotermitinae is one of the richest subfamilies of soil-feeding termites, which are traditionally classified in feeding groups according to anatomical criteria, deemed to the reveal whether species feed on organic-rich layers (group III) or on mineralised soil (group IV). Previous studies based on δ15N isotopic values showed that this subfamily's niche covers a broad range along the gradient of humification. We hypothesised that this broad feeding range could be reflected in the crop-gizzard (Cr-Gi) content and volume. We investigated 17 African species distributed between the two feeding groups. Our results showed a variation of Cr-Gi volume and a consistent composition of content among Apicotermitinae species. Some small-bodied species had a very large Cr-Gi volume relative to their size, indicating a difference in foraging behaviour. These species might use this enhanced storage capacity to forage for longer periods of time. Cr-Gi content was dominated by clay (kaolinite) suggesting that a dietary specialisation could be based on the quality of organic compounds from organo-mineral aggregates. Variations in crystalline solids (quartz) between species indicate either differences in the abundance of mineral grains between feeding patches or active discrimination among particles by foragers. The similar composition of Cr-Gi contents in afrotropical Apicotermitinae suggests that the anatomical criteria used to assign species to feeding groups III or IV are not appropriate.
... The Apicotermitinae subfamily was first proposed by Grassé and Noirot (1954) and defined by Sands (1972) in the most extensive taxonomic work on this group of termites carried out in Africa. Both morphological and molecular data corroborate the monophyly of Apicotermitinae (Inward et al. 2007;Bourguignon et al. 2017). ...
... The taxonomic work of Sands (1972) described 51 new species, redescribed 9 species, and established 16 new genera. Sands (1972) considered the genus Anoplotermes Müller to be exclusively Neotropical. ...
... The taxonomic work of Sands (1972) described 51 new species, redescribed 9 species, and established 16 new genera. Sands (1972) considered the genus Anoplotermes Müller to be exclusively Neotropical. At the time, this was the sole apicotermitine genus of the region. ...
Article
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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.
... Noirot and Noirot-Thimothée (1969) composed the chapter on the termite digestive system in the first volume of a two-volume work, the fundamental Biology of Termites (Krishna & Weesner, 1969, 1970, to this day an obligatory reference for termitologists and a "Bible" for the Isoptera until the publication of the treatise "Termitologia" by Grassé (1982Grassé ( , 1984Grassé ( , 1986. Sands (1972), in his study on soldierless termites of Africa, proposed a new classification of the Termitidae, broadening the definitions of Apicotermitinae (to include Anoplotermes) and Termitinae (to include Amitermitinae). For this purpose, he studied in detail the general external morphology, morphology of the mandibles, and the anatomy of the gut of workers. ...
... For example, the numbers of antennomeres and tibial spurs, in addition to the mandibles, were reported in detail by Weesner (1969), Grassé (1982), and Krishna et al. (2013). Characters of the internal morphology such as the gut coiling in situ, the mixed segment, the shape of the mesenteric tongue, and the insertion and arrangement of the Malpighian tubules (usually numbering four), and the enteric valve are derived mainly from the studies by Grassé and Noirot (1954), Noirot and Kovoor (1958), Kovoor (1969), Sands (1972), and Noirot (2001. ...
... The P2, with variable organization and ornamentation, is invaginated in the EVS. Certain recurring features aid in the delimitation of morphospecies and establishment of identity for Apicotermitinae; many of these are derived from the studies of Sands (1972Sands ( , 1998 and Noirot (2001). The foreleg shows several characters that deserve attention. ...
Article
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This contribution deals with the worker caste of the Neotropical termite fauna. It is a compilation of present knowledge about the morphology of pseudergates and workers, including the literature discussing the origin and evolution of this caste, the terminology used in the different taxonomic groups, and the techniques used to study these individuals, especially examination of the gut, mandibles, legs, and nota. In order to assist in identifying workers, it includes a key for the families that occur in the Neotropical Region and a characterization of workers of all families, especially the subfamilies of Termitidae, with descriptions and illustrations of diagnostic morphological features of genera. We point out advances and gaps in knowledge, as well as directions for future research.
... bugeserae Bouillon & Vincke, 1971) and the information about the various enteric valves seemed to pass unnoticed for over a quarter of a century. Sands (1972) was the first to use the enteric valves thoroughly in his revision of the African soldierless termites and to use them as generic and specific markers. Johnson (1979) recognised the variability of the proctodeal diverticulum in the genus Cubitermes. ...
... To that must be added a "premolar tooth" Pasteels 1969, Deligne 1999) situated between the third marginal tooth and molar tooth, only on the left mandible. This tooth is also called the "subsidiary marginal tooth" (Sands 1972;Uys 1994) or "4 th marginal tooth" (Sands 1992) (Fig. 3). ...
... The upstream and downstream ends (anterior and posterior ends in Uys 2002) refer to the flow of the alimentary bolus. Sands (1972Sands ( , 1998 numbered the PCs in a bilateral way with the cushion n°1 being innermost to the coiling of the gut and n°4 opposite n°1, while the same number (n°2) was given to the two cushions located on both sides of n°1 and the same number (n°3) to the two cushions on both sides of n°4 (Fig. 4B). This numbering is convenient for enteric valves with hexaradial, triradial or bilateral symmetries. ...
Article
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Seventy-four species, forms and varieties of Cubitermes Wasmann, 1906 have been studied, including taxa placed in synonymy with other species. Within this group of taxa, the enteric valve, mainly of the workers and soldiers, provided the best and the only clear-cut criteria for distinguishing some major subsets that are proposed here as species groups.The genus Cubitermes is re-described; the morphologies of the enteric valves are described in detail including the number of spatulae at the downstream end of the primary cushions; the possible presence of crests or bulges near the downstream end of the primary cushions; the overall shape of the primary cushions; the number of lateral supporting bristles; the kind of symmetry of the valve; and the structure of the secondary cushions. These characteristics are used to define nine species groups. For now, these groups have no taxonomic ranking but are helpful as regards species recognition. Identification keys for species groups are provided for soldiers and workers together with a partial key for imagines. Geographical ranges of the groups are also provided. On the basis of enteric valve morphology, some synonymies can no longer be validated: (a) C. planifrons Sjöstedt, 1924 is not a synonym of C. fungifaber (Sjöstedt, 1896); (b) C. kemneri Emerson, 1928 is not a synonym of C. zenkeri (Desneux, 1904); and (c) C. fungifaber var. elongata Sjöstedt, 1924 does not belong to the species C. fungifaber. Cubitermes planifrons and C. kemneri become valid again and C. fungifaber var. elongata is an invalid name.
... Morphological examination of the enteric valve armature (EVA) and digestive tube was done on the worker caste. The terminology used for the worker digestive tube and mandibles follows Noirot (2001), Sands (1972), and Deligne (1999). We used the term "molar process" proposed by Constantini et al. (2020). ...
... Since most samples from museums and collections are yet to be studied and there is a lack of inventories in many areas, the geographic gap for these termites is even greater than for other groups. This is also true in the other parts of the world, where the taxonomic gap is not so deep because of the older available literature (Grassé and Noirot 1954;Sands 1972), but the geographical gap still exists due the lack of researchers for a long period (Romero Arias et al. 2021). ...
Article
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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, Anoplotermes susanae Scheffrahn, Carrijo & Castro, sp. nov. and four new species in four new genera are described: Hirsutitermes kanzakii Scheffrahn, Carrijo & Castro, gen. nov. et sp. nov., Krecekitermes daironi Scheffrahn, Carrijo & Castro, gen. nov. et sp. nov., Mangolditermes curveileum Scheffrahn, Carrijo & Castro, gen. nov. et sp. nov., and Ourissotermes giblinorum 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.
... Harvested individuals are sorted and kept in bottles containing 70 ethyl alcohol. They are identified using a determination key of [10,11,12,13] . The identification was based on the caste of the soldiers according to the identification keys mentioned above. ...
... The identification was based on the caste of the soldiers according to the identification keys mentioned above. For those without soldiers, they were identified using the key of [13] . After identification, each species is classified in the trophic groups (Champignonnist, humivorus, xylophageous). ...
Article
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This work was done in the East of Côte d'Ivoire in Abengourou, in order to compare the termite diversity and their impact on two cultivated cocoa varieties « Mercedes » and « Tout venant ». One hundred (100) of 2-year-old cocoa plants were delineated on three plots by variety of cocoa. The results obtained show a comparatively identical specific richness in the two varieties. The unselected variety "Tout venant" records 18 species against 17 species for the selected variety "Mercedes". The comparison of the specific richness, the Shannon diversity index (H'), the Simpson index (SI) and the Equitability index (E) do not vary significantly between the two varieties of cocao. The density of termites varies significantly depending on the variety in the soil. The densities of the varieties "Mercedes" and "Tout venant" were respectively 2755.50±397.43 ind/ m 2 and 1612.37±208.74 ind/ m 2. Densities at the depth level were also compared for both varieties. Statistical analyzes show that the "Mercedes or CNRA" variety has the highest density at all depths. This density decreases with the depth of the two varieties. Eleven (11) species of termites have been identified, 8 of which are common to both varieties. The calculated Jaccard coefficient was 0.74. This value does not reflect any difference in the diversity of termites pests in both varieties. The attack rates of the "Mercedes" and "Tout venant" varieties were respectively 25.00 ± 1.15% and 26.33 ± 1.20%. These rates do not vary significantly between the two varieties. The species Ancistrotermes guineensis and Microtermes sp1 are the most aggressive. A. guineensis, with an attack rate of 34.68 ± 1.70 %, is more aggressive on the variety "Mercedes" and Microtermes sp1 on the variety "Tout venant" with an attack rate of 28.74±1.97%. This work shows that the new cocoa variety "Mercedes" does not influence the biodiversity and the attacks of the cocoa pest termites but shows a difference of the termite population in the soil depending on the variety and specificity of the termites pests depending on the species.
... The EV was photographed with a Leica CTR 5500 compound microscope with phase-contrast optics using the same montage software. Terminology of the worker gut follows that of Sands (1972) and Noirot (2001). Mandible terminology as in Sands (1972) except for the left subsidiary fourth marginal tooth which was clarified and redefined as the "premolar process" . ...
... Terminology of the worker gut follows that of Sands (1972) and Noirot (2001). Mandible terminology as in Sands (1972) except for the left subsidiary fourth marginal tooth which was clarified and redefined as the "premolar process" . ...
Article
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Chasitermes pax 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.
... Termite soldiers from the pill boxes in each batch are observed. The genus and species of each lot are determined using the identification keys of Hamad (1950) [15] , Bouillon and Mathot (1965) [16] , Roy-Noel (1966) [17] , Sjöstedt (1926) [18] and Sands (1965Sands ( , 1972Sands ( , 1992 [19,20,21] . ...
... Termite soldiers from the pill boxes in each batch are observed. The genus and species of each lot are determined using the identification keys of Hamad (1950) [15] , Bouillon and Mathot (1965) [16] , Roy-Noel (1966) [17] , Sjöstedt (1926) [18] and Sands (1965Sands ( , 1972Sands ( , 1992 [19,20,21] . ...
Article
Termites are one of the most important components of soil invertebrates and often cause several damages to fruit trees and cash crops, infecting their productivity. The objective of this study is to inventory the termite species that make up a primary forest and a corossol tree plantation in M'Brimbro (southern Côte d'Ivoire). The Transect standard method was used to sample over an area of 5000 m2 of corossol tree planting, i.e. half a hectare composed of 528 corossol trees in the plantation and 5000 m2 in the primary forest. Sampling made it possible to collect termites from the litter and tree trunks up to 1.5 m high. The obtained results have shown that the primary forest contained a more diversified termite population, with 15 species, than the corossol tree plantation (8 species). 6 species common to both environments have been identified. These are: Ancistrotermes guineensis, Ancistrotermes cavithorax, Macrotermes subhyalinus, Macrotermes bellicosus, Microtermes thoracalis and Pseudacanthotermes militaris. Among those species, the group of fungus-growers is the most represented with 8 species. All the identified species were grouped into 9 genera all found in primary forests and only 5 in corossol tree plantations. This study will allow us to know the biological diversity of the corossol plantation in this area of Côte d'Ivoire.
... From 1950 to 2013, thirty (30) termite species (Table 1) were recorded in The Gambia based on the works of Sands [1,[5][6][7]; Williams & Perez-Morales [8], Johnson et al. [9] and Krishna et al. [4]. ...
... The works of Sands focusing more on Nasutitermitinae [18] and on the genus Amitermes [19] are also used. Identification of the soldierless species of Apicotermitinae has been made after sands [5,7] on the basis of the morphology of the digestive tube: mesenteron-proctodeum junction and dissected enteric valves are observed under the stereomicroscope. Cubitermes species identification is based on the combination of the morphological characters of soldiers [3,[11][12][13]17] and the shape of the cushions of workers' enteric valves [20]. ...
Article
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From 28 October to 5 November 2013, a termite study was undertaken in 3 protected sites in The Gambia (West Africa). The aim of the study is to investigate the diversity of termites in three protected areas in the western region of the country. Termite sampling is carried out in 100 m × 2 m transects that are replicated three (3) times in each site. A total of thirty-one (31) termite species, that belong to fungus growing (11), harvester (1), humuvorous (12) and xylophagous (7), were recorded. The following nineteen (19) species are new to The Gambia: Coptotermes intermedius,
... Fontes (1986) described Aparatermes as a new genus, with Aparatermes abbreviatus (Silvestri 1901), formerly Anoplotermes, as the type species. Although Fontes (1986) recognized the importance of including digestive tube morphology in soldierless termite descriptions, as shown by Sands (1972) and Mathews (1977), he did not describe the enteric valve of A. abbreviatus. In addition, Fontes (1998) later transferred Anoplotermes cingulatus (Burmeister, 1839) to Aparatermes. ...
... The tube was immersed in PVA mounting medium (Bioquip, Rancho Dominguez, California, USA), to completely detach the enteric valve from surrounding muscle tissue, then cut longitudinally to splay open the enteric valve for mounting in the medium. Description of the worker gut follows the terminology of Sands (1972) and Noirot (2001). Enteric valves of specimens from A. abbreviatus, A. cingulatus, and A. silvestrii from previous samples deposited at the University of Florida, Termite Collection were dissected and included for comparison. ...
Article
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The worker of Aparatermes thornatus is described as the fourth species of a widespread Neotropical termite genus, Aparatermes, from the worker caste. This species occurs in the northern Amazonia. The enteric valve armature of A. thornatus and the unique coloration of its worker’s head capsule distinguished it from its three congeners. Molecular data recovered the proposed new species as a sister to A. silvestrii. Type material is deposited at Colección Entomológica Forestal, Universidad Distrital “Francisco Jose de Caldas” (CEFUDFJC- 45 National Registry of Collections) en Bogotá, Colombia and the University of Florida, Termite Collection (UFTC) in Fort Lauderdale Research and Education Center.
... Because these structures are delicate, PVA can continue acting on and damaging the tissue. Some classic studies of termite taxonomy use other methods for mounting the blades, such as Swan's Berlese (Sands, 1972), the Hoyle Medium (Constantino, 1999), and potassium hydroxide (KOH), which can be used to remove musculature. Visualizing the insertion of Malpighian tubules requires a specialized technique. ...
Article
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Apicotermitinae are soldierless termites highly abundant in tropical forests. The taxonomy of this subfamily is based on characters of worker cast and winged forms when present. However, the procedures necessary to dissect termite workers to observe their external and internal morphological characteristics are not well detailed in any study. Here, we describe a step-by-step protocol for worker dissection of soldierless termite species. We suggest the use of Polyvinyl Alcohol (PVA) for cleaning and visualization of the gizzard and enteric valve, and glycerin to remove tergites and sternites and describe in detail the dissection of the gizzard and enteric valve, and how visualize the insertion of Malpighian tubules.
... The investigation of such outliers may provide additional insights into the evolution of alarm behaviors, and into the ecological pressures driving them. Another interesting factor possibly influencing alarm communication is the presence of a soldier caste, which was lost at least three times independently in (i) Apicotermitinae, (ii) Orientotermes and Protohamitermes, and (iii) Invasitermes (all Termitidae [97][98][99] ). Workers in these groups are fully responsible for colony defense, and they thus reveal high levels of agonism and sometimes also developed unique defense strategies, such as body rupturing 34,47,100 . ...
Article
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Termites (Blattodea: Isoptera) have evolved specialized defensive strategies for colony protection. Alarm communication enables workers to escape threats while soldiers are recruited to the source of disturbance. Here, we study the vibroacoustic and chemical alarm communication in the wood roach Cryptocercus and in 20 termite species including seven of the nine termite families, all life-types, and all feeding and nesting habits. Our multidisciplinary approach shows that vibratory alarm signals represent an ethological synapomorphy of termites and Cryptocercus. In contrast, chemical alarms have evolved independently in several cockroach groups and at least twice in termites. Vibroacoustic alarm signaling patterns are the most complex in Neoisoptera, in which they are often combined with chemical signals. The alarm characters correlate to phylogenetic position, food type and hardness, foraging area size, and nesting habits. Overall, species of Neoisoptera have developed the most sophisticated communication system amongst termites, potentially contributing to their ecological success. The vibroacoustic and chemical alarm communication in the wood roach Cryptocercus and in 20 termite species are studied to reveal that the Neoisoptera species have developed the most sophisticated communication system, which could contribute towards their ecological success.
... Terminology follows that of Sands (1972) for mandible dentition and that of Noirot (1995Noirot ( , 2001 for gut anatomy. Measurements, as described in Roonwal (1969), were taken to the nearest 0.005 mm with a Wild MMS 235 length-measuring set fitted to a Wild M6 stereomicroscope. ...
Article
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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 Schievitermes globicornis , 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.
... Specimens, more specifically soldiers taken at random from each lot, are observed with a brand LEICA of the series EZ4 binocular loupe. The genus and species of each lot is thus determined using the identification keys [25][26][27][28]. ...
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Termites play an important ecological role. Thus, this study was carried out in the forest adjoining restaurant "the palm trees of the university campus of Cocody" to determine the species of termites and their impacts on the trees of this forest. Three transects of 100m long and 2m wide each subdivided into 20 sections and Tropical soil biology and fertility (TSBF) monoliths were developed. Then a systematic excavation was carried out along transects in search of termites. Then the plot containing transects was prospected in order to identify termite mounds and biogenic structures. After inventory, we identified 7 species of termites: Macrotermes bellicosus, Microtermes sp, Basidentitermes sp, Odontotermes sp, Pericapritermes sp, Amitermes guineensis and Ancistrotermes guineensis. Wood-feeders, fungus-growers and soil-feeders are the trophic groups found on the site. Fungus-growers were the group with the most species. We recorded a 25% attack rate on the trees from transects. A total of 13 nests were counted including 9 epigeal termite mounds and 4 arboreal nests. The low richness recorded shows that our plot is heavily anthropized. The conservation or maintenance of this site would allow species to recolonize this space.
... The total number of termite species identified was eighteen for all the habitats with species belonging to Rhinotermitidae (two species) and Termitidae (sixteen species) families. Amitermes (Silvestri 1901) Amitermes evuncifer (Silvestri 1912) * + Promirotermes (Silvestri 1914) Promirotermes holmgreni infera (Silvestri 1912) + + Apicotermitinae (Sands 1972) Nitiditermes (Emerson 1960) Nitiditermes proximatus (Silvestri 1914 Coptotermes intermedius, Microtermes lepidus, Odontotermes aff. erraticus, Amitermes evuncifer, and Nitiditermes proximatus (Fig. 2) represent "newly recorded species in Burkina Faso" (Table 1). ...
Article
Termites are insects found in tropical regions where they play a major ecological role. The objective of this study was to inventory termite species present in the Gourma province in eastern Burkina Faso in relation to habitat variations in order to contribute to the conservation of biodiversity. For this purpose, we sampled nine sites from three habitats (savannahs, fields, and fallows), i.e. three sites per habitat, using a standard rapid harvesting method for two weeks in August 2018. From this sampling, we identified 1285 individuals from which 18 termite species, 11 genera and 2 families. Our collections documented five species and one genus (Schedorhinotermes) recorded for the first time in Burkina Faso. The analysis of occurrence indices revealed that Macrotermes bellicosus was a constant savannah species, and Nasutitermes sp. and Schedorhinotermes sp. were accessory species whereas the majority of the species were rare. The Shannon and equitability indices showed that savannah supported a greater richness of termites (sixteen species) and a more equitable distribution of species, likely because savannah was less disturbed compared with crop fields and fallows and was therefore richer in food resources. Statistical analysis of functional groups revealed significant variation in termite in groups II and IV abundance among sampled habitats probably in relation to soil, vegetation, climate and anthropic disturbances. Our study has provided an update to the list of termite species and highlighted the ecological importance of this taxonomic group in Burkina Faso.
... The investigation of such outliers may provide additional insights into the evolution of alarm behaviors, and into the ecological pressures driving them. Another interesting factor possibly influencing alarm communication is the presence of a soldier caste, which was lost at least three times independently in (i) Apicotermitinae, (ii) Orientotermes and Protohamitermes, and (iii) Invasitermes (all Termitidae [97][98][99] ). Workers in these groups are fully responsible for colony defense, and they thus reveal high levels of agonism and sometimes also developed unique defense strategies, such as body rupturing 34,47,100 . ...
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Termites (Blattodea: Isoptera) have evolved specialized defensive strategies for colony protection. Alarm communication enables workers to escape threats while soldiers are recruited to the source of disturbance. Here, we studied the vibroacoustic and chemical alarm communication in the wood roach Cryptocercus and in 20 termite species including seven of the nine termite families, all life-types, and all feeding and nesting habits. Using a multidisciplinary approach, we found that vibratory alarm signals represent an ethological synapomorphy of termites and Cryptocercus , while chemical alarms evolved independently in several cockroach groups, and at least twice in termites. Vibroacoustic alarm signaling patterns were the most complex in Neoisoptera, in which they were often combined with chemical signals. The alarm characters correlated to phylogenetic position, food type and hardness, foraging area size, and nesting habits. Overall, species of Neoisoptera have developed the most sophisticated communication system amongst termites, potentially contributing to their ecological success.
... The widespread pantropical distribution of these groups may therefore be explained by their feeding ecology, possibly enabling them to colonize new biogeographic regions more easily than 'true' soil-feeding taxa feeding on, and living in, the soil (figure 2). These considerations may also pertain to the Apicotermitinae, as Adaiphrotermes, an Afrotropical genus sister to the Oriental and Neotropical apicotermitines [42] not included in our analyses, is often found within sound wood [79]. In any case, some soil-feeding lineages appear to be weak colonizers and never dispersed outside Africa, such as the basal Apicotermitinae, Promirotermes and the Cubitermitinae. ...
Article
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Termites feed on vegetal matter at various stages of decomposition. Lineages of wood- and soil-feeding termites are distributed across terrestrial ecosystems located between 45°N and 45°S of latitude, a distribution they acquired through many transoceanic dispersal events. While wood-feeding termites often live in the wood on which they feed and are efficient at dispersing across oceans by rafting, soil-feeders are believed to be poor dispersers. Therefore, their distribution across multiple continents requires an explanation. Here, we reconstructed the historical biogeography and the ancestral diet of termites using mitochondrial genomes and δ13C and δ15N stable isotope measurements obtained from 324 termite samples collected in five biogeographic realms. Our biogeographic models showed that wood-feeders are better at dispersing across oceans than soil-feeders, further corroborated by the presence of wood-feeders on remote islands devoid of soil-feeders. However, our ancestral range reconstructions identified 33 dispersal events among biogeographic realms, 18 of which were performed by soil-feeders. Therefore, despite their lower dispersal ability, soil-feeders performed several transoceanic dispersals that shaped the distribution of modern termites.
... Generally, it is difficult to use any morphological character of the worker caste in the identification of different species. However, studies on the general external morphology, structure of mandibles, and the anatomy of the gut of worker caste have been introduced by Sands (1972). In recent past, almost all new taxa descriptions of family Termitidae and revisionary works included features of the gut and mandibles of the workers (Rocha et al., 2019). ...
... The identification was based on morphological characters on caste soldiers or workers for groups of termites lacking the soldier castes. Several keys such as [8][9][10][11][12][13][14][15][16][17][18][19] have been used for termite identification. ...
... Individuals were first identified to genus and then to species level using a binocular magnifying glass, using the identification keys of [15][16][17][18][19][20][21][22]. After identification, each species was classified into one of the trophic groups (Fungus-growers, Soil feeders, Grass-feeders and Wood-feeders). ...
Article
Aims: This study aimed to assess the impact of human activities on termites in teak plantations in the Korhogo communal area. Methodology: Termites were sampled from October to November 2020 using the transect method recommended by Jones and Eggleton (2000). The study was carried out in three teak plantations undergoing different levels of human activities, with a forest fragment as reference area. Five types of human activity were assessed and the overall proportion of human pressure on each habitat was calculated. The species richness (S), Shannon index (H'), Evenness (E) and the relative abundance were calculated of termites for each habitat type. Analyses of variance (ANOVA) were used to compare the species richness and abundance of termites. Results: The results showed that the village plantation of teak (PVT) had the highest degree of human pressure (50.94%), followed by the teak plantation of the forest of Mount Korhogo (TFMK) (29.24%). The teak plantation of Botanical Garden (TJB) was under low pressure (6.60%). A total of 30 species grouped in 19 genera and 8 sub-families of termites were identified in all plots. Termite diversity was high in the forest fragment (19.67 ± 1.15) and in the teak plantation of Botanical Garden (21.33 ± 2.08), but low in the village teak plantation (11 ± 1). The abundance of termites evolves in the same direction as the species richness. Conclusion: Anthropogenic activities affect the trophic composition of termites, particularly the humivore group. Reconstruction of the fauna and flora of the teak forests would be beneficial for the conservation of termite species. In this region, teak forests would thus play a role as a refuge for termite communities, which are recognised as the main soil fertilising organisms in the tropics.
... Ebogotermes is closest to Aderitotermes Sands, 1972 workers (A. cavator and A. fossor) which Sands (1972) describes as being large (HW= 0.90-0.99 mm) with the fore tibia "scarcely swollen" and a longer mixed segment than Ebogotermes. ...
Article
Ebogotermes raphaeli gen. n. sp. n., is described from workers collected in Cameroon. This soil-feeding termite is the largest soldierless termite from central Africa and aligns with the Anoplotermes subgroup. The enteric valve armature is weakly armed and, as with most apicotermitine species, is uniquely diagnostic.
... The identification of termite species was done using a stereomicroscope and was based on the caste of soldiers or workers for termite species without soldiers (dissection of the enteric valve or comparison with specimens with complete castes). The identification of the termite species sampled was done using various taxonomic determination keys developed by Bouillon and Mathot (1965), Sands (1959Sands ( , 1965Sands ( , 1972Sands ( , 1992, Roy-Noel (1966), Harris (1966), Ruelle (1970). Termite voucher specimens were deposited at the National High School of Applied Biosciences and Biotechnologies (ENSBBA) ...
Article
Termites are considered as important pests of maize plantations in Republic of Benin. However, very little information is available on their assemblage composition, abundance, damages, and on the sensitivity of common maize varieties in Benin. This study aims to fill these gaps and provide baseline for the development of an integrated termite management in maize plantations. Therefore, 60 maize fields across 30 villages located in four agro-ecological zones of southern Benin were prospected using the standardized method of belt transects. In addition, the resistance of five maize varieties to termite attacks was evaluated on-farm during two consecutive seasons (2018 and 2019). Fifteen termite species belonging to 11 genera, 6 subfamilies and 3 families were collected in the prospected maize fields. Four species (Macrotermes subhyalinus Rambur, Coptotermes sjostedti Holmgren, Amitermes evuncifer Silvestri, and Trinevitermes germinatus Wasmann) were found in all agro-ecological zones with A. evuncifer as the most abundant species. High similarity indices of termite assemblages were found between the four agro-ecological zones. Wood-feeders group and species nesting in hypogeal nests were the mostly abundant in all agro-ecological zones. The attack rate of termites in maize fields was 16.9% and did not vary significantly between agro-ecological zones. A. evuncifer and C. sjostedti were significantly the most destructive species on the five evaluated varieties, although Trinevitermes oeconomus Trägardh was most abundant in leaf sheaths; M. subhyalinus and Macrotermes bellicossus Smeathman in the stem; Pseudacanthotermes militaris Hagen and Odontotermes sp. in the roots. Termite damage level to maize depended of the variety. Korga variety was classified as highly resistant to termite attacks; Agbadé kouin winiwini variety as resistant; Igbado tchakpa variety as moderately resistant. While, Elèkpèrè and Forkui varieties were classified as highly susceptible. The three maize varieties identified as resistant to termite attack should be included in the development of an integrated termite management program and in future breeding programs.
... The identification of termite species was done using a stereomicroscope and was based on the caste of soldiers or workers for termite species without soldiers (dissection of the enteric valve or comparison with specimens with complete castes). The identification of the termite species sampled was done using various taxonomic determination keys developed by Bouillon and Mathot (1965), Sands (1959Sands ( , 1965Sands ( , 1972Sands ( , 1992, Roy-Noel (1966), Harris (1966), Ruelle (1970). Termite voucher specimens were deposited at the National High School of Applied Biosciences and Biotechnologies (ENSBBA) ...
Article
Rice (Oryza spp.) is an important food and cash crop in the Republic of Benin. However, despite its production increase in recent years, the yield of cultivated varieties remains low, and the introduction of improved varieties threatens the rice diversity existing in traditional agriculture. Therefore, documenting the on-farm management of rice diversity, farmers’ varietal preferences, and their perceptions of the performance of cultivated varieties and species are important prerequisites for the development of on-site breeding and conservation programs. To fill these gaps in Benin, 418 rice farmers, belonging to 21 ethnic groups, were surveyed in 39 villages using participatory rural appraisal tools. Subject to synonymy, 30 improved varieties and 68 local varieties were registered and their folk nomenclature and taxonomy were documented. The north of Benin had the highest diversity of rice with the greatest number of traditional varieties,making this region the best place for an in situ conservation program. The number of rice varieties maintained per village varied from 1 to 15 (six on average). The foursquare analysis revealed that the improved variety IR 841 was by far the most popular variety.Most ofNERICA varieties were abandoned in the south, while the north still hosts a wide range of local varieties. Twenty-one reasons explained varietal abandonment by farmers, varying according to geographic areas and ethnic groups. The seed system was both formal and informal in the study area. The participatory evaluation revealed the necessity to create and introduce tolerant/resistant rice varieties to drought and flooding stresses in Beninese agriculture that meet farmers’ preferences. Our results showed that north Benin would be the most suitable place for in situ conservation of local rice diversity.
... It is based on the caste of the soldiers using the identification keys of: [20,21,22,23] and [24,25] . The termites without soldiers, the workers were analyzed using the key of [26] . ...
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The general objective of this study is to know the diversity of termites in papaya plantations in southern Côte d'Ivoire and the damages caused by those termites. A papaya plantation was sampled using the standardized method for the rapid estimation of termite biodiversity. Two transects were plotted on diagonals 111 m long, 2 m wide and divided into 10 sections each. A last transect 100 m long and 2 m wide was drawn vertically in the middle of the farm and divided into 5 sections. A total of 25 sections have been delineated and searched. The results obtained show that five species are present in this papaya plantation. The species Pseudacanthotermes spiniger, Ancistrotermes crucifer, with respectively 70.05% and 12.79% of occurrence are the most abundant in the medium. Macrotermes bellicosus records the lowest occurrence at 1.08%. The fungus-growers make up the majority of termites harvested on the plot. And out of 68% of the attacked papaya trees, 38% has recorded structural damages, and 05% of the plants/trees would have been died because of termites, have been noticed. This work is a preliminary study that shows the impact of the papaya crop on termite biodiversity. However, the variation of termite communities according to the age of the plantation deserves to be studied.
... L'identification des termites est basée sur la caste des soldats au moyen des clés d'identifications de : Hamad (1950); Bouillon et Mathot (1965); Harris (1966, 1968) et Sands (1965. Pour ceux dépourvus de soldats, les ouvriers ont été analysés à l'aide de la clé de Sands (1972). 3.3.3 ...
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Objectifs : Elle a pour objectif d’évaluer l’effet insecticide, sur les termites, de l’extrait aqueux de Tithonia diversifolia dosé à 131,25 g/L comparé à un termiticide chimique, le pyriforce (chlorpyrifos éthyl) dosé à 480 g/L dans les parcelles de riz nerica1. Méthodologie et résultats : Cette étude a été menée sur une parcelle expérimentale de l’Institut National Polytechnique Houphouët-Boigny (INP-HB) de Yamoussoukro (Côte d’Ivoire). Trois carrés de fouille (25 cm de côté sur 30 cm de profondeur) ont été réalisés dans chaque parcelle élémentaire (5mx4m). Au total, 18 carrés de fouille par traitement ont été effectués. Au total, 11 espèces de termites ont été récoltées dans l’ensemble des parcelles. Les champignonnistes avec 6 espèces, constituent le groupe dominant. Quatre (4) espèces d’humivores et une (1) espèce de xylophage ont été récoltées. Seuls les champignonnistes sont présents dans les milieux traités aux extraits de plantes et au chlorpyrifos ethyl. Au stade maturité, c’est à dire 4 mois après semis, la densité des termites dans les parcelles traitées s’est montrée plus faible avec l’extrait aqueux de T i t h o n i a d i v e r s if o li a (913,78 ind/m2). Conclusion : Le chlorpyrifos éthyl pourrait être utilisé à titre curatif pour son action rapide sur les termites tandis que le T i t h o n i a d i v e r s if o li a pourrait être utilisé pour les cultures à cycle long.
... As in the soldierless Apicotermitinae (Sands, 1972), the Cubitermes enteric valve armature (EVA), mainly of the worker caste, can be used for differentiating species groups (Josens & Deligne, 2019). In the subfamily Cubitermitinae, the basic enteric valve is a funnel bearing six elongated primary, prominent cushions of the valve (PC) alternating with non-prominent, secondary cushions of the valve (SCs) (Noirot, 2001). ...
Article
The phylogeny of the genus Cubitermes Wasmann was reconstructed using two mitochondrial genes (COI and COII) and a fragment of the nuclear 28S rDNA, including samples of Apilitermes Holmgren, Crenetermes Silvestri, Megagnathotermes Silvestri and Thoracotermes Wasmann. Based on our analyses, we recovered these genera within a paraphyletic Cubitermes clade. Cubitermes species are distributed between five main clades, highly reflective of their enteric valve armatures (EVAs). Consequently, Cubitermes is here divided into five monophyletic genera based on phylogenetic analyses and EVAs: Cubitermes (sensu stricto), Isognathotermes Sjöstedt, Nitiditermes Emerson, Polyspathotermes Josens & Deligne, gen.n., and Ternicubitermes Josens & Deligne, gen.n. Moreover, the two species of Megagnathotermes included in this study, exhibiting different EVAs, are phylogenetically distant: M. katangensis Sjöstedt belongs in the genus Polyspathotermes, while M. notandus Silvestri remains as a monotypic genus. During the evolution of the Cubitermes complex, sclerotized EVA spatulae appeared twice independently: three or six spatulae in Polyspathotermes (sulcifrons and oblectatus patterns) and two jaw‐like spatulae in Nitiditermes (sankurensis pattern), which is absent in some species (oculatus pattern). Zoobank registration: http://zoobank.org/urn:lsid:zoobank.org:pub:576AAFF8‐63C6‐4962‐A8F5‐75C9317AEA7B.
... Whilst in the laboratory, the samples were examined and photographed using a MOTIC ST-30C binocular magnifier equipped with an AmScope MD130 camera connected to a computer. All specimens collected were identified to the level of species or morphospecies using the keys of Sjöstedt (1926), Emerson (1928), Ahmad (1950), Bouillon and Mathot (1965), Grassé (1986), Sands (1959Sands ( , 1972Sands ( , 1998, Roy-Noel (1966), Harris (1966). After identification, the species were classified into the following respective trophic groups. ...
Article
Termites play an important role in the process of decomposition of organic matter in tropical ecosystems and participate in the evolution of soil structure. This study was carried out in the north of Côte d’Ivoire to evaluate the influence of cashew tree orchards’ age on termite assemblages. The standardised method for rapid estimation (RAP) of biodiversity was used to collect termites in four (4) classes of cashew tree orchards (Class 1: ˂ 5 years, Class 2: 5–10 years, Class 3: 11–20 years, Class 4: 21 years and above) in comparison to savannah. The results showed that species richness, were low in young orchards of class 1 (9 species). This increased progressively with age to its maximum in older orchards of class 4 (30 species). These old orchards were even richer in termite species than the savannah (27 species). Shannon-Weiner diversity index varied in the same way as species richness. The abundance of termites, were low in young orchards (45 occurrences), higher in older orchards of class 4 (423 occurrences), with a predominance of fungus-growing termite species. This study showed the recovery of termite diversity in cashew tree orchards after the stoppage in agricultural activities. Suggesting that, this form of agroforestry would allow for the conservation of diversity of the soil fauna while ensuring food security in the area.
... However, when soldiers are not present, either in the foraging party sampled or because the termite genus in question is soldierless (e.g. the highly diverse Apicotermitinae of African and South American rainforests and wetter savannas), identification is more challenging. The traditional approach has been to perform gut dissections (Sands 1972), but these are time-consuming and require high levels of expertise. We instead recommend that DNA barcoding techniques be used for these soil-feeding termites, as well as for other samples without soldiers, using the cytochrome oxidase II mitochondrial gene (Hausberger et al. 2011;Bourguignon et al. 2014Bourguignon et al. , 2016. ...
Article
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Termites are globally dominant and functionally important soil organisms. While their role in ecosystems is being increasingly recognised and understood, methods that adequately sample termite communities across habitats can be challenging and have not advanced at the same pace as studies of termite ecology. Moreover, the appropriateness of sampling methods varies with habitat and biogeographic region due to differences in termite communities. Focusing largely on tropical systems where the majority of termites occur, we review currently available termite sampling methods and provide recommendations for sampling across different biomes and biogeographic regions. Active searching transects are most effective in rainforest habitats, whereas reduced transects, baiting and mound surveys are more appropriate in grassy systems and seasonally dry forests. Baiting is recommended for deserts. Recent advances in termite sampling, such as the use of remote sensing and DNA metabarcoding, and outstanding challenges, such as sampling episodic grass-feeding termites, are also discussed. Improved use of standardised termite sampling methods, as we recommend, should lead to increased knowledge of the patterns and drivers of termite diversity, which will, in turn, facilitate the quantification of the influence termites have on ecosystems and lead to new insights into the functioning of tropical systems.
... The identification of soldierless termites usually requires the examination of the enteric valve armature, which needs to be dissected and mounted on microscope slides (Sands 1972). The gut segment with the valve should be cut and removed, slit longitudinally, and cleaned ( Fig. 10.10). ...
Chapter
Termites are eusocial, polymorphic insects which live in colonies with distinct morphological castes: reproductives, workers, and soldiers. They are important decomposers in natural ecosystems, and several species are urban and agricultural pests. Termite nests may be simple excavated tunnels inside wood, or constructions of variable size and shape, arboreal, epigeic, or subterranean. Collecting termites requires the use of several tools to open their nests, take a representative sample of the various castes present, and preserve them in 70–80% ethanol. Many sampling methods have been used for termites, including belt transects, large plots, soil monoliths, baiting, and alate traps. Each method has advantages and limitations, and some may be used complementarily. Termite identification requires good samples with a good number of specimens of all castes, especially soldiers.
... Termite identification: Termites harvested were identified in the laboratory of biology and tropical ecology of the University Jean Lorougnon Guédé of Daloa. Specimens were determined up to the level of species using various documents such as Hamad (1950), Bouillon and Mathot (1965), Sands (1965Sands ( , 1972Sands ( , 1998. After identification, each species was classified into one of the trophic groups. ...
Article
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To maintain its status of leading producer of cocoa in the world, Côte d'Ivoire must be contain the different types of threats, including the emergence of termites in cocoa farms. This study was conducted in the south-west of the country to evaluate the effect of cropping systems (full sun system, intermediate system and shaded system) on termite attacks. The study was conducted in four localities with different cocoa agroforestry systems. The sampling was made on cocoa trees in quadrats of 30mx30m and the attack rate was calculated. Eleven (11) termite species were identified. The wood-feeders groups were dominant. These termites were responsible for 41.82% of the damage on cocoa trees. Termite Attacks vary between different cropping systems. Full sun systems were most attacked comparing to intermediate shade systems that were least attacked by termites. This work shows that the shade system has an effect on termite attacks in cacao trees. The choice of right system can help to significantly reduce termite attacks in cocoa farms.
... The taxonomic significance of the termite worker gut characters, including the enteric valve armature (EVA), was first addressed by a few authors including Grassé & Noirot (1954), Noirot & Kovoor (1958), Noirot (1966), Sands (1972), Mathews (1977), and Johnson (1979). Gradually, the worker gut has become a prominent character in the descriptions of soil-associated taxa as exemplified by the monumental work of Sands (1998). ...
Article
Gastrotermes spinatus gen. n. sp. n is described from workers of a single foraging group collected in Cameroon. This soil-feeding termite aligns with the Labidotermes subgroup (Apicotermes group) because of its non-protruding and symmetrical enteric valve armature, its short P1, and its globular P3a. An asymmetrical field of robust sclerotized spines at the opening of the P3a is unique among the other Labidotermes subgroup genera.
... However, they are not totally defenseless as workers can defecate on enemies, self-sacrifice through the rupture of the body wall or utilize labial or frontal defensive glands [59]. Few confirmed records of ants feeding on Apicotermitinae exist in the literature and they all originate from Africa [60]. However, we believe that the two potential prey interactions that we inferred with the CCAs, between Solenopsis subterranea and Anoplotermes ADA8662 and between Wasmannia auropunctata and Microcerotermes arboreus, are plausible. ...
... The collected termites were identified at the Biology and Animal laboratory (Université Péléforo Gon Coulibaly of Korhogo). Specimens were identified to the level of species or morphospecies using standard determination keys such as: [1,5,31,32,33,12,34]. We were aware that morphological identification alone may lead to the confusion of sibling species [30,14] , e.g. in Odontotermes or Microtermes, but a molecular characterization of all samples to sort out complexes of morphologically indistinguishable, closely related species was beyond the scope of this study. ...
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Termites are one of the most abundant invertebrate groups in tropical ecosystems. Despite their important role, termites can cause considerable damage to wooden structures in most tropical countries. This study aim is to evaluate the termite diversity on the campus of the University Peleforo Gon Coulibaly of Korhogo and to show their attacks on trees and buildings. According to the standardized rapid estimation method (RAP) of biodiversity, a total of 12 species of termite were collected. Fungus-growers group with 68% of termite total abundance were the mostly abundant group. The study of termite attacks showed that plants were strongly attacked by termites. Out of 294 trees visited, 274 trees showed traces of termite attacks, a rate of 93%. Termite damage caused leaf drop, dieback and / or death of the plant. Concerning buildings and woodwork, termites caused significant damage on these structures. The results of our study are a preliminary study to draw the special attention of university officials to take action to combat these termites.
... The body rupture performed by these species, followed by the release of a sticky secretion, seems to be a pertinent defensive strategy. Comparative analyses should be encouraged to fully understand the development of these defensive mechanisms and identify their occurrence in other soldierless taxa [1,26,27]. ...
Article
Termite soldiers constitute the defensive frontline of the colonies, despite workers also perform such tasks, especially within the Neotropical Apicotermitinae, in which all species are soldierless. Workers of the genus Ruptitermes display an extreme form of defense, characterized by body rupture and release of a sticky secretion. Previous observations suggested that such behavior may be advantageous against enemies, but the chemical composition of this secretion has been neglected. Here we firstly provide the proteomic profile of the defensive secretion of Ruptitermes reconditus and Ruptitermes pitan workers. Additionally, the mechanisms of action of this behavior was evaluated through different bioassays. A total of 446 proteins were identified in R. reconditus and 391 proteins in R. pitan, which were classified into: toxins, defensins and proteolytic enzymes; sticky components/ alarm communication; proteins related to detoxification processes; proteins involved in folding/conformation and post-translational modifications; housekeeping proteins; and uncharacterized/hypothetical proteins. According to the bioassays, the self-sacrifice is triggered by a physical stimulus, and the defensive secretion may cause immobility and death of the opponents. Assuming that termites are abundant in the tropics and therefore exposed to predators, suicidal behaviors seem to be advantageous, since the loss of an individual benefit the whole colony. Significance Although recent studies have reported the biochemical composition of different weapons in soldiered species of termites, such efforts had not been applied to sordierless taxa up until now. Thus, this is the first report of the defensive mechanisms in soldierless termite species based on proteomic analysis. The diversity of compounds, which included toxin-like and mucin-like proteins, reflect the mechanisms of action of the defensive secretion released by termite workers, which may cause immobility and death of the opponents. Our findings may contribute to the knowledge regarding the development of defensive strategies in termites, especially in groups which lost the soldier caste during the evolution.
... Termite identification was carried out at the Evolutionary Ecology and Biology laboratory at the Université Libre de Bruxelles (ULB), based on caste soldiers or workers for groups of termites lacking the soldier castes. Specimens of the same genus were determined to species using various standard determination keys developed by Sands (1959Sands ( , 1965Sands ( , 1972, Bouillon and Mathot (1965), and Ruelle (1970). ...
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Termites are known to cause significant economic losses in yam fields. Therefore, assessing diversity and abundance of termites in yam fields is critical for their control. In this study, we compared the specific richness, abundance, and diversity of functional groups of termite pests in upland and lowland yam fields, across 15 villages using the standardized belt transect method. The specific termite diversity encountered in yam fields was compared with that of the Agoua classified forest. In general, eight genera classified into four subfamilies (Nasutitermitinae, Macrotermitinae, Amitermitinae and Coptotermitinae) grouped into two families (Termitidae and Rhinotermitidae) were identified. The highest number of the species (13) were identified in upland fields while the lowest number (10) were identified in the lowlands. Termite fauna collected from yam fields and from the Agoua forest were distinct, as showed by the similarity index. Amitermes evuncifer (Silvestri) was the most widely distributed and the most dominant termite species in upland yam fields while in lowland yam fields it was Macrotermes subhyalinus (Rambur). Three functional groups were observed with humus feeders only found in upland yam fields. Principal component analysis classified the 24 sampled transects into four groups with different termites feeding groups. The dominance of non-fungus-growers was observed in the groups mostly composed of transects in uplands yam fields. Three nesting habitats were represented (epigeal mound, wood, and subterranean). The results of this present research could help to develop efficient strategies against the identified key yam termite pests. Les termites sont connues pour causer des pertes économiques importantes dans les champs d'igname. Par conséquent, l'évaluation de la diversité et de l'abondance des termites dans les champs d'igname est essentielle pour leur gestion. Dans cette étude, nous avons comparé la richesse spécifique, l'abondance et la diversité de groupes fonctionnels des termites ravageurs dans les champs d'igname de plateaux (15) et bas-fonds (9), à travers 15 villages en utilisant la méthode normalisée de transect en ceinture. La diversité spécifique de termites rencontrée dans les champs d'igname a été comparée à celle de la forêt classée du Bénin d'Agoua. Huit genres classés en quatre sous-familles (Nasutitermitinae, Macrotermitinae, Amitermitinae et Coptotermitinae) regroupés en deux familles (Termitidae et Rhinotermitidae) ont été identifiés. Le plus grand nombre d'espèces (13) a été identifié dans les champs de plateaux, tandis que le plus petit nombre (10) a été identifié dans les bas-fonds. La faune de termites collectée dans les champs d'ignames et dans la forêt d'Agoua a été distincte sur base des indices de similarité. Amitermes evuncifer (Silvestri) a été l'espèce de termite la plus largement répandue et la plus dominante dans les champs d'ignames de plateaux, tandis que dans les bas-fonds, il s'est agi de Macrotermes subhyalinus (Rambur). Trois groupes fonctionnels ont été observés avec des consommateurs d'humus que l'on ne trouve que dans les champs d'igname de plateaux. Suite à l'analyse en composantes principales, les 24 transects échantillonnés ont été classés en quatre groupes ayant des termites avec différents comportements alimentaires. La prédominance des non-cultivateurs de champignons a été observée dans les groupes Entomologie Faunistique-Faunistic Entomology 2019 72 112 principalement composés de transects des champs d'igname de plateau. Trois habitats de nidification étaient représentés (monticule épigé, bois et sous-sol). Les résultats de la présente recherche pourraient aider à développer des stratégies efficaces contre les principaux termites ravageurs de l'igname identifiés.
... Samples of each maize' termite pest identified by farmers were collected and preserved in labelled boxes containing 70% alcohol for later identification at the Laboratory of Applied Entomology (LEnA) of the Faculty of Sciences and Technology of Dassa (Loko et al. 2015b). The specimens were identified to species level based on the caste soldiers by using various standard determination keys established by Sands (1959Sands ( , 1965Sands ( , 1972, Bouillon and Mathot (1965) and Ruelle (1970). ...
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Termite (Isoptera: Termitidae) infestation is an important constraint of maize production in Benin. A survey of 300 households was conducted in 2018 throughout 30 villages in Southern Benin to evaluate farmers’ knowledge, perceptions and management practices of maize termite pests using focus group discussions and individual interviews with a semi-structured questionnaire. The results showed that for most of the farmers, maize is the most susceptible crop to termite attacks and the maturation developmental stage having the highest termite abundance and damage. A total of 43 different names of maize termite pests corresponding to 8 species were recorded in the study area. Size and colour were the main criteria used by farmers to classify and identify maize termite pests. Amitermes evuncifer was perceived as the most damaging to maize during vegetative and maturation stages, while Macrotermes bellicosus was perceived by famers (32.5%) as most damaging during seedling and flowering maize development stages. Erratic rainfall was perceived by maize producers as the most important factor favouring termite infestation in maize fields. The majority of the local maize is considered by farmers as susceptible to termites. The use of synthetic insecticides was the main method to control termites. Four of the 20 pesticides used by farmers have active ingredients that have been listed as highly hazardous (class Ib) by the World Health Organization (WHO). Farmers training on the danger and impact of misuse of insecticides on the development of termite insecticide resistance was recommended. The development of an integrated termite management strategy is discussed and the future research for its implementation identified.
... Ruptitermes workers may burst their dehiscent organs, exceptional structures which contain glycoproteins and are used during this altruistic behavior (Costa-Leonardo 2004;Poiani and Costa-Leonardo 2016). Furthermore, Sands (1972) reported that workers of some Apicotermitinae species perform abdominal dehiscence, breaking the abdominal cuticle and spreading their inner organs to the exterior. Thus, self-sacrifice behavior concomitant to chemical defense occurs in Neotropical soldierless taxa, like in the case of the genus Ruptitermes (Poiani and Costa-Leonardo 2016) and in species which present soldiers such as Neocapritermes taracua (Bourguignon et al. 2015;Sobotinik et al. 2012). ...
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Termite societies are abundant in the tropics, and are therefore exposed to multiple enemies and predators, especially during foraging activity. Soldiers constitute a specialized defensive caste, although workers also participate in this process, and even display suicidal behavior, which is the case with the species Neocapritermes braziliensis. Here we describe the morphology, mechanisms of action, and proteomics of the salivary weapon in workers of this species, which due to the autothysis of the salivary glands causes their body rupture, in turn releasing a defensive secretion, observed during aggressiveness bioassays. Salivary glands are paired, composed of two translucent reservoirs, ducts and a set of multicellular acini. Histological and ultrastructural techniques showed that acini are composed of two types of central cells, and small parietal cells located in the acinar periphery. Type I central cells were abundant and filled with a large amount of secretion, while type II central cells were scarce and presented smaller secretion. Parietal cells were often paired and devoid of secretion. The gel-free proteomic approach (shotgun) followed by mass spectrometry revealed 235 proteins in the defensive secretion, which were classified into functional groups: (i) toxins and defensins, (ii) folding/conformation and post-translational modifications, (iii) salivary gland detoxification, (iv) housekeeping proteins and (v) uncharacterized and hypothetical proteins. We highlight the occurrence of neurotoxins previously identified in arachnid venoms, which are novelties for termite biology, and contribute to the knowledge regarding the defense strategies developed by termite species from the Neotropical region.
... This process of feeding of one colony member by another is known as trophyllaxis. Some termite species belonging to the sub-family Apicotermitinae have no soldiers (Sands, 1972). ...
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The Oriental termite genus, Speculitermes (Termitidae: Apicotermitinae) was previously considered soldierless, as soldiers are rarely encountered. Among the 12 species of Speculitermes, soldier caste is described only for three species. Herewith we report hitherto undescribed soldier caste of S. dharwarensis for the first time with additional records of its presence from the southern India.
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The Centre for Overseas Pest Research (COPR) exists to help overseas territories, particularly the developing countries, solve pest problems in the field of agriculture and public health. It concentrates on pest species which are of international or regional significance either because they migrate between countries, for example, locusts and weaver birds (Quelea), or because they are widespread and endemic, for example, termites and mosquitoes. COPR undertake laboratory research and overseas field investigations into the biology, ecology and pest status of pre-harvest pests of agriculture and certain public health pests. It is also concerned with the development and application of modern pest control techniques, as well as the long-term environmental effects of the use of pesticides in the tropics. It is a new organization in name only, since it has the experience and knowledge gained from over 30 years of pest research and control in overseas territories, as it was formed in June 1971 from already existing scientific units of the British Government's Overseas Development Administration.
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Termites represent a group of eusocial insects that live in colonies containing up to hundreds to millions. They are highly abundant, exceeding in tropics 6,000 individuals per square meter. Due to their abundance, termites represent an important food source for a wide variety of predators. At the same time, termite defensive adaptations allow the colonies to overcome the predator pressures, being extremely evolutionary successful. This achievement is explained by the development of a complex communication system operated by a rich set of exocrine glands. As many as 20 different exocrine organs are known in termites. Some of these organs had received negligible attention being only known by anecdotal observation. One of these was the labral gland. In this study, I examined the structure and ultrastructure of the labrum in soldiers of 28 species, workers of 28 species and imagoes of 33 species across termites’ main representatives, and in the wood roach Cryptocercus. The labral gland was present in all species and castes, and comprises two secretory regions located on the ventral side of the labrum and the dorso–apical part of the hypopharynx, respectively. The epithelium of the gland consisted of class 1 secretory cells, with an addition of class 3 secretory cells in soldiers of few species. A common feature of the secretory cells was the abundance of smooth endoplasmic reticulum (an organelle known to produce lipidic and often volatile secretions), long microvilli with a channel inside, which releases the secretion through a modified cuticle. According to the structure, ultrastructure and behavioural experiments, my results suggest that the labral gland is involved in defensive communication after encounter to an alien. On the other hand, other glands are extensively studied in some castes but have received almost no attention in other castes. It is the case of the frontal gland, an organ without any equivalent among other animals. The frontal gland is well known in soldiers and imagoes but not much was known about it in workers. In order to provide a complete picture of the evolution of this gland in termite workers and consequently in termites, I studied it in 41 additional species across Neoisoptera. The frontal gland of these species was formed by class 1 secretory cells only, and occured as an epithelial without reservoir in all cases. My data suggest that the frontal gland would have caste–specific evolutionary routes, being its ancestral form epithelial with reservoir in soldiers and imagoes, while epithelial thickening in workers. This study was the first to provide a comprehensive picture of the structure of the labral and frontal gland across all termite taxa and castes, providing fundamental information to enhance our understanding about the evolution and social behaviour of Isoptera.
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The anatomy of the workers' digestive tube is essential in taxonomical studies of soil-feeding Apicotermitinae termites, especially in soldierless lineages. Two structures, the mesenteric-proctodeal junction and the enteric valve, have long been important to distinguish genera and species. By contrast, the gizzard (proventriculus) has been almost ignored by taxonomists because of its generally regressed state in soil-feeding termites. In this study, we document in detail for the first time the sclerotized structures and ornamentations in the gizzard in the Apicotermitinae subfamily. We identified two main clusters of species: those without ornamentations and those exhibiting a sclerotized pulvillar armature, which may include spicules or spines of diverse sizes, numbers and dispositions. The latter group comprises the majority of African soldierless species, a widely diverse and dominant group in tropical forests and savannas. We outline the potential role of the anatomy of the gizzard in the taxonomy of Apicotermitinae based on the interspecific anatomical variation of the pulvillar armatures. We suggest that sclerotized ornamentations regulate the flow of food particles and break or lacerate aggregates to facilitate the access of enzymes in the midgut.
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Neocapritermes opacus is a Neotropical termite, which feeds on a variety of food sources, including highly decayed wood and humus. Similar to many Termitidae, this species contains a mixed segment in the intestine, an anatomic distinct region composed of both mesenteric and proctodeal tissues. In Neocapritermes spp., a pair of intestinal protrusions referred to as mesenteric sacs are associated with the mixed segment. In the present study, with the aid of histological and scanning microscopy techniques, we provide a detailed anatomical and histological analyses of the mesenteric sacs in N. opacus workers. The results showed that the paired sacs are contiguous with the mesenteric component of the mixed segment and open into the ectoperitrophic space. The wall of the sacs is an epithelium composed of two types of cells (regenerative and principal) and musculature. The regenerative cells are located at the epithelium base and organized in regenerative crypts, while the principal cells are columnar and enfold the lumen of the sacs. Brush border was observed in the principal cell apex, aligned closely to rod-like bacteria. Secretory activity was observed in the principal cells and vesicles cast from them toward the sacs lumen, probably result of an apocrine activity. The sac epithelium was of mesenteric affiliation but differed from the intestinal midgut. The proctodeal epithelium was markedly different. Although some studies have been interpreted the mesenteric sacs as Malpighian nodules or Malpighian sacs, our results support that these structures are morphologically distinct and that the occurrence of mesenteric sacs is a synapomorphy for Neocapritermes. The precise function of the mesenteric sacs in termite digestion remains unclear; therefore, further investigations should carefully consider their secretory activity and symbiont population, aiming to understand their development and maintenance among these Neotropical species.
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