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Roisinitermes ebogoensis gen. & sp. n., an outstanding drywood termite with snapping soldiers from Cameroon (Isoptera, Kalotermitidae)


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Termites have developed a wide array of defensive mechanisms. One of them is the mandibulate soldier caste that crushes or pierces their enemies. However, in several lineages of Termitinae, soldiers have long and slender mandibles that cannot bite but, instead, snap and deliver powerful strikes to their opponents. Here, we use morphological and molecular evidence to describe Roisinitermesebogoensis Scheffrahn, gen. & sp. n. from near Mbalmayo, Cameroon. Soldiers of R.ebogoensis are unique among all other kalotermitid soldiers in that they possess snapping mandibles. The imago of R.ebogoensis is also easily distinguished from all other Kalotermitidae by the lack of ocelli. Our study reveals a new case of parallel evolution of snapping mandibles in termites, a complex apparatus responsible of one of the fastest biological acceleration rates measured to date.
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Roisinitermes ebogoensis gen. & sp. n., an outstanding drywood termite... 91
Roisinitermes ebogoensis gen. & sp. n., an outstanding
drywood termite with snapping soldiers from
Cameroon (Isoptera, Kalotermitidae)
Rudolf H. Scherahn1, omas Bourguignon2,3, Pierre Dieudonné Akama4,
David Sillam-Dussès5,6, Jan Šobotník3
1 Fort Lauderdale Research and Education Center, Institute for Food and Agricultural Sciences, 3205 College
Avenue, Davie, Florida 33314, USA 2 Okinawa Institute of Science & Technology Graduate University,
1919-1 Tancha, Onna-son, Okinawa 904-0495, Japan 3 Faculty of Forestry and Wood Sciences, Czech Uni-
versity of Life Sciences, Prague, Czech Republic 4 Département des sciences biologiques, Ecole normale supérieu-
re, Université de Yaoundé I, BP 47 Yaoundé, Cameroon 5 University Paris 13 - Sorbonne Paris Cité, LEEC,
EA4443, Villetaneuse, France 6 IRD – Sorbonne Universités, iEES-Paris, Bondy, France
Corresponding author: Rudolf H. Scherahn (rhsc@u.edu)
Academic editor: P. Stoev | Received 5 July 2018 | Accepted 27 August 2018 | Published 2 August 2018
Citation: Scherahn RH, Bourguignon T, Akama PD, Sillam-Dussès D, Šobotník J (2018) Roisinitermes ebogoensis
gen. & sp. n., an outstanding drywood termite with snapping soldiers from Cameroon (Isoptera, Kalotermitidae).
ZooKeys 787: 91–105.
Termites have developed a wide array of defensive mechanisms. One of them is the mandibulate soldier
caste that crushes or pierces their enemies. However, in several lineages of Termitinae, soldiers have
long and slender mandibles that cannot bite but, instead, snap and deliver powerful strikes to their
opponents. Here, we use morphological and molecular evidence to describe Roisinitermes ebogoensis
Scherahn, gen.& sp. n. from near Mbalmayo, Cameroon. Soldiers of R. ebogoensis are unique among
all other kalotermitid soldiers in that they possess snapping mandibles. e imago of R. ebogoensis is also
easily distinguished from all other Kalotermitidae by the lack of ocelli. Our study reveals a new case of
parallel evolution of snapping mandibles in termites, a complex apparatus responsible of one of the fast-
est biological acceleration rates measured to date.
Ethiopian Region, mandibles, ocellus, taxonomy
ZooKeys 787: 91–105 (2018)
doi: 10.3897/zookeys.787.28195
Copyright Rudolf H. Scheffrahn et al. This is an open access article distributed under the terms of the Creative Commons Attribution License
(CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
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Rudolf H. Scherahn et al. / ZooKeys 787: 91–105 (2018)
Termites are extremely abundant (Martius 1994, Eggleton et al. 1996) and colonies
may contain millions of individuals attracting a wide variety of predators (Deligne et
al. 1981). Additionally, termites experience strong intra- and inter-specic competi-
tion (Levings and Adams 1984, orne and Haverty 1991). To combat against the
plethora of agonistic opponents, termites have developed a rich array of defensive strat-
egies. e most important defenses are expressed in the soldier caste that is ancestral to
all extent termites (Roisin 2000).
Soldiers are specialized sterile colony defenders possessing exaggerated morphology
of the head and mandibles (Prestwich 1984). One of their most intriguing defenses is
exemplied by long and slender snapping mandibles (Deligne et al. 1981). e snap-
ping mandibles are paired with muscles to store potential energy which, when released,
delivers a powerful strike producing one of the fastest accelerations known among
animals (Seid et al. 2008). All termite species with snapping soldiers described so far
belong to the Termitinae (Bourguignon et al. 2017), suggesting that snapping soldiers
evolved several times independently within this subfamily. Alternatively, soldiers with
snapping mandibles might have evolved once, and independently reverted to a biting
strategy in several lineages.
e monophyletic family Kalotermitidae (Inward et al. 2007) constitutes almost
half of all “lower termite” genera and species (Krishna et al. 2013) and has fossil re-
cords to the mid-Cretaceous (Engel et al. 2009). Kalotermitids live entirely in wood as
“one-piece” nesters (Abe 1987) which facilitates transoceanic dispersal (Scherahn and
Postle 2013). Kalotermitids occur in all ecozones and numerous genera have vast dis-
tributions (e.g. Calcaritermes, Cryptotermes, Glyptotermes, Kalotermes, Marginitermes,
Neotermes, and Procryptotermes). A few species of Cryptotermes (Scherahn et al. 2009)
and Incisitermes (James et al. 2013, Yasuda et al. 2003) have also been dispersed by hu-
man activity. A few species are major pests of dry wood (Su and Scherahn 2000) or
minor pests of tree crops (Constantino 2002).
e monumental revision of the Kalotermitidae by Krishna (1961) provided the
morphological diagnoses for all extant genera with the exception of the recently de-
scribed Longicaputermes (Ghesini et al. 2014). Aside from Longicaputermes, all new
kalotermitid species described after Krishnas 1961 revision, ca. 115, have been as-
signed to one of the 21 genera he recognized. e soldier caste of several genera has
unmistakable characters: e.g., the scooped out frons of Eucryptotermes, the massive
third antennal article of Marginitermes, the large ovoid head of Pterotermes, or the
spur on the fore tibia of Calcaritermes. We herein describe a new genus and species
of Kalotermitidae, Roisinitermes ebogoensis, which possesses equally unmistakable
soldiers. e soldier of R. ebogoensis is the rst outside the Termitinae to have snap-
ping mandibles.
Roisinitermes ebogoensis gen. & sp. n., an outstanding drywood termite... 93
Material and methods
Illustrations and measurements
Images of individuals were taken as multi-layer montages using a Leica M205C stere-
omicroscope with a Leica DFC 425 module run with Leica Application Suite software
version 3. Preserved specimens, stored in 85% ethanol, were positioned in a transparent
petri dish lled with Purell hand sanitizer (70% EtOH). Measurements (Tables1–2)
were obtained using an Olympus SZH stereomicroscope tted with an ocular microm-
eter. A eld photograph of live specimens placed in a small paper-lined Petri dish was
taken with a Canon EOS 5DS R combined with a Canon EF 100mm f/2.8L Macro IS
USM lens. Morphological terminology follows that of Krishna (1961).
Phylogenetic analyses
DNA was extracted from ve individuals of R. ebogoensis, after removal of the di-
gestive tract. e full mitochondrial genome was amplied with TaKaRa LA Taq in
two long PCR reactions using primers specically designed for termites (Bourguignon
etal. 2016). Long PCR fragments were pooled in equimolar concentration, and 75-
bp paired-end reads were obtained using Illumina MiSeq. We subsampled a total of
10,000 reads and assembled the full mitochondrial genome with SPAdes, under default
parameters (Bankevich et al. 2012). e total coverage of the assembly was 82-fold.
We used the mitochondrial genomes of ten species of Kalotermitidae, including
one sample of Roisinitermes ebogoensis sequenced in this study. We used four non-Ka-
lotermitidae termite species as outgroups to root the tree: Zootermopsis angusticollis,
Hodotermopsis sjostedti, Coptotermes sjostedti, and Termitogeton planus. All mitochon-
drial genomes, except that of R. ebogoensis, have been published recently (Suppl. mate-
rial1: Table S1). Each gene of the mitochondrial genome was aligned separately using
MAFFT v7.300b with the option “--maxiterate 1000 --globalpair” for higher accuracy.
For protein-coding genes, we rst aligned genes as protein, then converted protein se-
quence alignments into the corresponding codon alignments using PAL2NAL (Suyama
et al. 2006). e 22 tRNAs and the two ribosomal RNAs were aligned as DNA. e
resulting alignments were concatenated with FASTconCAT v1 (Kück and Meusemann
2010). Alignments were separated in ve partitions: one for each codon position of the
protein-coding genes, one for the combined ribosomal RNA genes, and one for the
combined tRNA genes.
We reconstructed phylogenetic trees using Maximum Likelihood and Bayesian
approaches. We ran the analyses twice, once with the third codon position included,
and once without third codon position. e Bayesian phylogenies were implemented
Rudolf H. Scherahn et al. / ZooKeys 787: 91–105 (2018)
in MrBayes 3.2 (Ronquist et al. 2012) with unlinked partitions, each of four chains
(three hot and one cold). e chain length was of two million generations with sam-
pling every 2000 generations. 800,000 generations were discarded as burnin, to en-
sure that the chain reached convergence, as determined by Tracer 1.5 (Rambaut and
Drummond 2007). We ran two replicates of each analysis to ensure consistency of the
results. For each partition of the data, we assigned an independent Generalized Time
Reversible model with gamma-distributed rate variation across sites and a propor-
tion of invariable sites (GTR + G +I). e reconstruction of Maximum Likelihood
phylogenies was carried out with RAxML (Stamatakis et al. 2008). We used the GTR-
GAMMA model of rate heterogeneity across sites. Node support was estimated using
1000 bootstrap replicates.
Phylogenetic analysis
Our phylogenetic analyses supported the monophyly of Kalotermitidae (Figure 1).
e four analyses yielded identical tree topologies, with one exception: in the Bayesian
analysis without third codon position Rugitermes was the sister group of Neotermes +
Cryptotermes + Incisitermes + Roisinitermes, while in the other three analyses Rugitermes
+ Neotermes sp. A formed the sister group of Neotermes insularis + Cryptotermes +
Incisitermes + Roisinitermes. Roisinitermes was consistently placed next to N. insularis.
Roisinitermes Scherahn, gen. n.
Type-species. Roisinitermes ebogoensis Scherahn sp. n.
Winged Imago. Ocelli not visible either by pigmentation or cuticular protrusion
(Figure 2A–C). Fore wing with unsclerotized media and cubitus arising from a com-
mon vein distal from scale suture; radial sector with 5–6 anterior branches; subcosta
very close and dicult to discern from costal margin (Figure 2D). Hind wing with
radial sector and cubitus arising from a common vein distal to suture. Tibial spurs
3:3:3; tarsi without arolia. e left imago/nymph mandible with anterior margin of
their marginal tooth ca. 1.5 times longer than length of the posterior margin of the rst
plus second marginal tooth; right mandible with posterior margin of second marginal
tooth 1.4 times as long as molar plate (Figure 3).
Diagnosis. e lack of visible ocelli is unique among all other Kalotermitidae. In
Krishnas 1961 generic key, Roisinitermes would lead to couplet 2 (Epicalotermes).
Roisinitermes ebogoensis gen. & sp. n., an outstanding drywood termite... 95
Figure 1. Phylogenetic tree of Kalotermitidae based on full mitochondrial genomes. e tree depicted
was reconstructed with RAxML using the data matrix without third codon position. Node labels are the
Maximum Likelihood bootstrap supports and the Bayesian posterior probabilities in the following order,
from left to right: posterior probability of the analysis with third codon position included, posterior prob-
ability of the analysis without third codon position, bootstrap support of the analysis with third codon
position included, bootstrap support of the analysis without third codon position, *indicates 100% boot-
strap support and 1.0 posterior probability for all four analyses.
Soldier. Monomorphic (Figs 4, 5). Eye spots prominent; large, dark brown. Frons
bilobed in dorsal view, crested with rugose longitudinal wrinkles, rugosity below frons
oriented longitudinally. Small horn-like projection at terminus of ventral genae. Man-
dibles sticklike; downward arching in lateral view. Dentition very weak; basal humps
project sharply.
Diagnosis. Stick-like mandibles unique among all other kalotermitid soldiers. In
Krishnas 1961 key, Roisinitermes leads to couplet 17 (Allotermes). In dorsal view, the
mandibular blades of Allotermes, especially A. denticulatus Krishna 1962, somewhat
resemble those of Roisinitermes as those of the former are long, rather narrow and with
rudimentary dentition. In lateral view, however, the Roisinititermes mandibles dier
from all other kalotermitids with projecting mandibles in that the Roisinitermes mandi-
bles arch downward. Although the Roisinitermes imago venation and dentition is very
similar to those of Epicalotermes, the soldier of Roisinitermes shares no major characters
with the Epicalotermes soldier.
Etymology. e genus is named in honor of Dr. Yves Roisin for his many contri-
butions to the study of termites.
Rudolf H. Scherahn et al. / ZooKeys 787: 91–105 (2018)
Figure 2. Imago of Roisinitermes ebogoensis gen. & sp. n. A Dorsal view of head and thorax B Oblique
view of head C Lateral view of head and thorax D Right forewing (arrow on subcosta) and right hind wing.
Roisinitermes ebogoensis Scherahn, sp. n.
Material examined. Holotype. Soldier from colony UF no. AFR3327. CAMEROON:
Ebogo II, (+3.37723N, +11.46135E), 647 m elev., 18FEB18, col. Raphael Onana,
AFR3327 ca. 500 alates, 50 soldiers, and many pseudergates, nymphs, larvae, and
eggs. Paratypes. CAMEROON, Ebogo II (+3.38273N, +11.46190E), 664 m elev.,
10DEC2016, col. Jan Šobotník and collaborators, AFR2982 4 soldiers (1 damaged),
one female dealate, and 46 brachypterous nymphs.
Diagnosis. See generic diagnosis above.
Description. Winged Imago (Figure 2, Table 1) Head and pronotum light brown-
ish orange; eye ovoid, anterior margin truncate abdominal tergites lighter, concolorous
with legsand labrum; postclypeus nearly hyaline. Compound eyes black, of medium
size and protrusion; ellipsoid but truncated near antennal socket, composed of approx-
imately 85 facets. Ocelli not visible either by pigmentation or cuticular protrusion.
Antennae with more than nine articles; formula 1>2=3=4<5. Pronotum width twice
that of median length; several long and shorter setae project from lateral margins. Fore
wing scale with basal origins of all major veins; wing membrane covered with papillae.
Tibial spurs 3:3:3; tarsi without arolia.
Soldier (Figs 4–6; Table 2) Monomorphic. In dorsal view, head capsule yellowish
orange in posterior grading to orange in middle and reddish brown from frons to ante-
Roisinitermes ebogoensis gen. & sp. n., an outstanding drywood termite... 97
clypeus. ree proximal antennal articles sepia brown; distal articles light brown. Post
clypeus and labrum yellowish with brown highlights. Eye spots prominent; large, dark
brown, elliptical; formed from a mass of discrete ommatidia. Pronotum concolorous
with posterior head capsule. Head capsule in dorsal view, subrectangular; lateral mar-
gins nearly parallel, length 1.5 times width. Posterior corners of head evenly rounded;
Figure 3. Brachypterous nymph of Roisinitermes ebogoensis gen. & sp. n. Top: Dorsal view of mandibles.
Bottom: lateral view of head and thorax.
Rudolf H. Scherahn et al. / ZooKeys 787: 91–105 (2018)
posterior margin rectate. In lateral and oblique view, head capsule almost cylindrical
with only slight dorso-ventral compression; frons bilobed in dorsal view, crested with
rugose longitudinal stripes, rugosity lateral below frons to mandibles. In lateral view,
frons sloping from vertex ~45°; mandibles bow upward to form a 15° arch. Setae short
and sparse on pronotum and head capsule. Periantennal carina rugose, in dorsal view
partially eclipsing the rst antennal article. Small horn-like projection at terminus of
ventral genae. Mandibles stick-like; long, blade narrower in middle than distal third,
dentition very weak; left mandible with faint equilateral tooth approx. three fths
from base, serrations along blade from tooth to tip. Right mandible with single tooth
approx. one third distance from base; blade narrowest before tooth; after tooth blade
widens slightly and then gradually narrows at tip. In dorsal view, basal humps project
sharply as rugose hemispheres. In lateral view, humps are columnar and equal in height
to that of the mandibles. Anteclypeus shallowly incised in middle; labrum linguiform
with gradual point; 4–5 long terminal setae. Antennae with 12–13 articles, third an-
tennal article subclavate, barely shorter than fourth and fth combined. Pronotum
collar-shaped; much wider than long. Anterior margin weakly concave; lateral margins
weakly convex, posterior margin forming 25° angle with incised middle. Femora mod-
erately inated, tibial spurs 3:3:3. Habitus as in Figure 6.
Brachypterous nymph (Fig. 3, Table 3) Body hyaline. Head, thorax, and abdo-
men similar in shape and pilosity of imago. Compound eyes with approx. 85 dark
Table 1. Measurements (mm) of Roisinitermes ebogoensis alates from a single colony.
Males (n=6) Females (n=6)
Measurement max min mean max min mean
Head max. width 1.05 0.95 1.00 1.05 1.00 1.03
Pronotum max. width 1.00 0.89 0.96 1.05 0.93 1.01
No. antennal articles 15 14 14.67 17.00 14.00 15.17
Max diam. eye 0.40 0.32 0.36 0.39 0.35 0.37
Body length with wings 9.63 8.63 9.10 9.88 9.50 9.65
Fore wing length (suture to tip) 7.50 6.80 7.20 7.80 7.20 7.43
Table 2. Measurements of Roisinitermes ebogoensis soldier (n=17 from two colonies).
Measurement Max Min Mean
Head length to lateral mandible base 1.92 1.60 1.79
Head width, maximum 1.28 1.18 1.22
Head height with gula, max. 1.08 0.92 1.00
Pronotum length 0.70 0.56 0.65
Pronotum width 1.18 1.05 1.13
No. antennal articles 14 10 12.70
Left mandible width @ basal humps 0.35 0.21 0.26
Left mandible width @ middle 0.18 0.16 0.17
Max. diam. eye 0.26 0.18 0.21
Length left mandible from condyle (ventral) 1.78 1.46 1.66
Roisinitermes ebogoensis gen. & sp. n., an outstanding drywood termite... 99
Figure 4. Soldier (holotype) of Roisinitermes ebogoensis gen. & sp. n. Dorsal (A), lateral (B), and ven-
tral(C) views of head and pronotum.
Rudolf H. Scherahn et al. / ZooKeys 787: 91–105 (2018)
Figure 5. Dorsal view of frons and mandibles of Roisinitermes ebogoensis gen. n. sp. n. Inset: oblique
ventral view of columnar hump (arrow).
facets; both eyes and facets smaller than imago. Antennae with 15 articles; formula
1>2>3=4=5. Left mandible with anterior margin of marginal tooth 1.5 times longer
than length of the posterior margin of the rst plus second marginal tooth. Right man-
dible with posterior margin of second marginal tooth 1.4 times as long as molar plate.
Biology and distribution. e type colony of R. ebogoensis was collected in the
forest on an island in the Nyong River near the Ebogo II village. e colony lived in a
Table 3. Measurements (mm) of Roisinitermes ebogoensis brachypterous nymph (n=10).
Measurement Max Min Mean
Head max. width 1.10 1.00 1.07
Pronotum max. width 1.16 1.08 1.11
No. antennal articles 15 15 15
Maximum diam. eye 0.20 0.20 0.20
Roisinitermes ebogoensis gen. & sp. n., an outstanding drywood termite... 101
relatively thin (3 cm) and long (over 3 m) broad-leaf tree branch suspended from the
canopy approximately 2 m above the ground. e colony contained roughly 2,000
members. A second colony of R. ebogoensis was collected in a nearly pristine rain forest
near the village of Ebogo II. e colony was taken from a dead liana branch (ca. 15
mm diam.) hanging from the canopy at a height of approx. 1 m above the ground.
Liana stems have been generally overlooked as a colonization site for Kalotermitidae
(Scherahn et al. 2018). In light of Emersons 1925 description of Cryptotermes cubio-
ceps from a single soldier collected from a dead liana, this host should be probed rou-
tinely as a colonization site for kalotermitids.
Etymology. e species is named for the village of Ebogo II, the type locality for
this termite.
Kalotermitids inhabit a single woody item and are largely unable to move to a new
food source once the original is exhausted. e lone exception is Paraneotermes sim-
plicicornis that builds underground galleries connecting several wood pieces (Light
1937). e ability to feed on sound wood represents a defensive adaptation in itself
as the hard food source acts as an ecient physical barrier against intruders. Kaloter-
mitids thus show low soldier-to-worker ratios (see Haverty 1977) and soldiers reach
a high level of polymorphism, reected especially in the development of the headcap-
Figure 6. Live habitus of soldier and brachypterous nymphs of Roisinitermes ebogoensis sp. n.
Rudolf H. Scherahn et al. / ZooKeys 787: 91–105 (2018)
sule and mandibles. Some genera such as Bicornitermes, Cryptotermes, Eucryptotermes,
Calcaritermes, or Glyptotermes, possess very short mandibles and a plug-like headcap-
sule to prevent intruder entry into a nest gallery (phragmosis). In C. cryptognathus
from Jamaica, the mandibles are reduced to small stubs that do not project beyond the
frontogenal boundaries of the head capsule, and therefore cannot be used to bite op-
ponents (Scherahn et al. 1998). Some other genera (e.g., Biditermes, Epicalotermes,
Incisitermes, Kalotermes, Neotermes) possess long mandibles with robust dentition
(crushing mandibles sensu Prestwich 1984) used to injure an opponent mechanically.
is is often combined with release of defensive secretions originating in the labial
glands (Šobotník et al. 2010, Sillam-Dussès et al. 2012). Epicalotermes pakistanicus
has particularly long and serrated mandibles (Akhtar 1974). e defensive strategy
of Roisinitermes soldiers does not match any of these; instead, Roisinitermes employs a
unique strategy of snapping, achieved by long and slender mandibles pressed against
each other in a defensive encounter. When this potential energy is released, the left
mandible springs over the right and the resultant snap is forced onto the opponent if
it is in the path of the strike. is singular mandibular modication was previously
known in several lineages of Termitinae (Deligne et al. 1981, Prestwich 1984, Seid et
al. 2008), and was portrayed as a defensive strategy unique to this group. Roisinitermes
represents the rst undisputable evidence of parallel evolution of snapping soldiers.
Our phylogenetic analyses consistently placed Roisinitermes on a long branch,
next to N. insularis. Neotermes insularis is a large termite species from Northern Aus-
tralia with soldiers endowed with biting mandibles of crushing type. e smaller
Roisinitermes shares no obvious similarity with N. insularis, supporting its generic
status. Currently, the number of mitochondrial genomes available for Kalotermitidae
is limited to a handful of genera, and there is a possibility that future phylogenetic
analyses will support anities between Roisinitermes and yet-to-be sampled taxa. In
any case, the highly unusual morphology of Roisinitermes suggests that it shares no
close relatives among modern Kalotermitidae. Future studies should focus on whether
the mechanisms used by soldiers of Roisinitermes to snap are like those of the distantly
related Termitinae.
e authors thank Eliška Cintulová, Crystal Clitheroe and Barbora Křížková for assis-
tance on R. ebogoensis full mitochondrial genome sequencing. J.Š. is grateful to Ebogo
II inhabitants for their help during the eldwork, and for helping saving the Ebogo
forest for future generations. J.Š. and D.S.-D. thank Aleš Buček, František Jůna, and
Margot Archambeu for their hard work during the eld campaigns. e eld work
was supported by the Czech Science Foundation (project no. 16-05318S), by the
Internal Grant Agency of Faculty of Forestry and Wood Sciences, CULS (IGA No.
A_27_18) and by the Grand Agency of the Czech University of Life Sciences (CIGA
No. 20184307).
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Supplementary material 1
Table S1
Authors: Rudolf H. Scherahn, omas Bourguignon, Pierre Dieudonné Akama, Da-
vid Sillam-Dussès, Jan Šobotník
Data type: molecular data
Explanation note: Sources and GenBank accession numbers of mitochondrial se-
quences used for Fig. 1.
Copyright notice: is dataset is made available under the Open Database License
( e Open Database License
(ODbL) is a license agreement intended to allow users to freely share, modify, and
use this Dataset while maintaining this same freedom for others, provided that the
original source and author(s) are credited.
... The systematics of Kalotermitidae has greatly benefited from the revision of Krishna (1961) who proposed a robust interpretation and delineation of the kalotermitid genera. More recently, this pioneering work was complemented with the description of two additional genera (Ghesini & Marini 2014;Scheffrahn et al. 2018). ...
Anewfossil kalotermitid species of the genus Cryptotermes is described and illustrated as Cryptotermes pouilloni n. sp. from the Miocene Dominican amber (Dominican Republic). This new species can be differentiated from the other Cryptotermes species, inter alia, by its small size (one of the smallest Cryptotermes species), anterior margin of anteclypeus concave, 15 antennomeres, and forewing RP with five secondary branches reaching wing anterior margin and M joining RP after wing midlength. The new species originates from La Toca mines while the exact origins of the other Dominican fossil Cryptotermes species are unknown. We summarize the Isoptera diversity from Dominican amber and rapidly discuss the impact of inaccurate dating of the Dominican amber on the time divergence estimate of the isopteran constitutive families.
... These lineages are deemed "basal" and exhibit many traits considered ancestral; however, they have also evolved their own unique traits [31,34,35]. As a consequence, it can be challenging to distinguish between plesiomorphic (ancestral) and apomorphic (derived) traits, and there are still limitations to determine whether a trait evolved multiple times independently, or whether it evolved once and was then lost on repeated occasions [30,37,38]. These uncertainties can create confusion and be the subject of debate among proponents of contradicting hypotheses. ...
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Termites are a clade of eusocial wood-feeding roaches with > 3000 described species. Eusociality emerged ~ 150 million years ago in the ancestor of modern termites, which, since then, have acquired and sometimes lost a series of adaptive traits defining of their evolution. Termites primarily feed on wood, and digest cellulose in association with their obligatory nutritional mutualistic gut microbes. Recent advances in our understanding of termite phylogenetic relationships have served to provide a tentative timeline for the emergence of innovative traits and their consequences on the ecological success of termites. While all “lower” termites rely on cellulolytic protists to digest wood, “higher” termites (Termitidae), which comprise ~ 70% of termite species, do not rely on protists for digestion. The loss of protists in Termitidae was a critical evolutionary step that fostered the emergence of novel traits, resulting in a diversification of morphology, diets, and niches to an extent unattained by “lower” termites. However, the mechanisms that led to the initial loss of protists and the succession of events that took place in the termite gut remain speculative. In this review, we provide an overview of the key innovative traits acquired by termites during their evolution, which ultimately set the stage for the emergence of “higher” termites. We then discuss two hypotheses concerning the loss of protists in Termitidae, either through an externalization of the digestion or a dietary transition. Finally, we argue that many aspects of termite evolution remain speculative, as most termite biological diversity and evolutionary trajectories have yet to be explored.
... Snapping mandibles have evolved in four independent clades of Termitidae and in one clade of Kalotermitidae 10,34,36,37 . These mandibles have been hypothesised to be a defensive weapon, especially in tunnels 6,12 , where both termites and their enemies are confined to narrow spaces (e.g., approximately 2-mm wide, as reported for Pe. ...
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The asymmetric mandibles of termites are hypothetically more efficient, rapid, and powerful than the symmetric mandibles of snap-jaw ants or termites. We investigated the velocity, force, precision, and defensive performance of the asymmetric mandibular snaps of a termite species, Pericapritermes nitobei. Ultrahigh-speed recordings of termites revealed a new record in biological movement, with a peak linear velocity of 89.7–132.4 m/s within 8.68 μs after snapping, which caused an impact force of 105.8–156.2 mN. High-speed video recordings of ball-strike experiments on termites were analysed using the principle of energy conservation; the left mandibles precisely hit metal balls at the left-to-front side with a maximum linear velocity of 80.3 ± 15.9 m/s (44.0–107.7 m/s) and an impact force of 94.7 ± 18.8 mN (51.9–127.1 mN). In experimental fights between termites and ant predators, Pe. nitobei killed 90–100% of the generalist ants with a single snap and was less likely to harm specialist ponerine ants. Compared with other forms, the asymmetric snapping mandibles of Pe. nitobei required less elastic energy to achieve high velocity. Moreover, the ability of P. nitobei to strike its target at the front side is advantageous for defence in tunnels.
... Termites have been proven to be social cockroaches which are sister to the woodroach Cryptocercus by many comprehensive molecular phylogenetic analyses (Inward et al. 2007;Cameron et al. 2012;Bourguignon et al. 2015). There are 458 species assigning to 23 living genera in termite family Kalotermitidae in the world (Krishna 1961;Krishna et al. 2013;Ghesini et al. 2014;Scheffrahn et al. 2018). The habitat of Kalotermitidae is in wood in small colonies which are with irregular galleries filled with faecal pellets. ...
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In this article, we describe the complete mitochondrial genome of Neotermes koshunensis from the Pingtung County, Taiwan. This mitogenome is 15,589 bp long, containing 13 protein-coding genes, 22 tRNA genes, and 2 rDNA genes. Nucleotide composition of the whole mitogenome is 42.86% for A, 25.42% for T, 19.65% for C, and 12.07% for G. The AT and GC skewness of mitogenomic sequences are 0.255 and −0.239, showing the A-skew and C-skew. Neotermes koshunensis grouped within the clade including the other nine Kalotermitidae species was well supported. The phylogenetic position of Kalotermitidae is sister to Neoisoptera (including Termitidae, Rhinotermitidae, Serritermitidae, and Stylotermitidae) in the current phylogenetic analysis. Mitogenomic data from this study will provide useful information for further studies on the phylogeny of Kalotermitidae.
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Termites are major decomposers in terrestrial ecosystems and the second most diverse lineage of social insects. The Kalotermitidae form the second-largest termite family and are distributed across tropical and subtropical ecosystems, where they typically live in small colonies confined to single wood items inhabited by individuals with no foraging abilities. How the Kalotermitidae have acquired their global distribution patterns remains unresolved. Similarly, it is unclear whether foraging is ancestral to Kalotermitidae or was secondarily acquired in a few species. These questions can be addressed in a phylogenetic framework. We inferred time-calibrated phylogenetic trees of Kalotermitidae using mitochondrial genomes of ∼120 species, about 27% of kalotermitid diversity, including representatives of 21 of the 23 kalotermitid genera. Our mitochondrial genome phylogenetic trees were corroborated by phylogenies inferred from nuclear ultraconserved elements derived from a subset of 28 species. We found that extant kalotermitids shared a common ancestor 84 Mya (75–93 Mya 95% HPD), indicating that a few disjunctions among early-diverging kalotermitid lineages may predate Gondwana breakup. However, most of the ∼40 disjunctions among biogeographic realms were dated at less than 50 Mya, indicating that transoceanic dispersals, and more recently human-mediated dispersals, have been the major drivers of the global distribution of Kalotermitidae. Our phylogeny also revealed that the capacity to forage is often found in early-diverging kalotermitid lineages, implying the ancestors of Kalotermitidae were able to forage among multiple wood pieces. Our phylogenetic estimates provide a platform for critical taxonomic revision and future comparative analyses of Kalotermitidae.
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.
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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.
The New World Kalotermitidae constitute about one-fourth of all termite species in this region. The geographic distributions of all fifteen kalotermitid genera are updated based on records in the University of Florida Termite Collection and in the literature. Range-appropriate exotic records of four pest species are also given. New distribution records are presented on maps, and representative soldier photographs are provided for each genus.
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The complete mitochondrial genome of drywood termite, Incisitermes minor, is reported in this study. The circular mitogenome has a length of 15,970 bp and encodes 37 genes including 13 protein-coding genes (PCGs), 22 transfer RNA (tRNA), two ribosomal RNA (rRNA), and a non-coding control region (D-loop). The percentage of A and T (65.44%) within this mitogenome is much higher than that of G and C (34.56%). The phylogenetic tree revealed that mitogenomes of Kalotermitidae formed one clade. The tree also revealed that I. minor was closest to Cryptotermes secundus, and was a sister group to Neotermes. I. minor is a only species in which mitogenome has been completed so far among the Incisitermes termite. The data provide resource for ecological and evolution analysis within termites especially Kalotermitidae.
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The higher termites (Termitidae) are keystone species and ecosystem engineers. They have exceptional biomass and play important roles in decomposition of dead plant matter, in soil manipulation, and as the primary food for many animals, especially in the tropics. Higher termites are most diverse in rainforests, with estimated origins in the late Eocene (∼54 Ma), postdating the breakup of Pangaea and Gondwana when most continents became separated. Since termites are poor fliers, their origin and spread across the globe requires alternative explanation. Here, we show that higher termites originated 42-54 Ma in Africa and subsequently underwent at least 24 dispersal events between the continents in two main periods. Using phylogenetic analyses of mitochondrial genomes from 415 species, including all higher termite taxonomic and feeding groups, we inferred 10 dispersal events to South America and Asia 35-23 Ma, coinciding with the sharp decrease in global temperature, sea level, and rainforest cover in the Oligocene. After global temperatures increased, 23-5 Ma, there was only one more dispersal to South America but 11 to Asia and Australia, and one dispersal back to Africa. Most of these dispersal events were transoceanic and might have occurred via floating logs. The spread of higher termites across oceans was helped by the novel ecological opportunities brought about by environmental and ecosystem change, and led termites to become one of the few insect groups with specialized mammal predators. This has parallels with modern invasive species that have been able to thrive in human-impacted ecosystems.
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Reticulitermes, Heterotermes and Coptotermes form a small termite clade with partly overlapping distributions. Although native species occur across all continents, the factors influencing their distribution are poorly known. Here, we reconstructed the historical biogeography of these termites using mitochondrial genomes of species collected on six continents. Our analyses showed that Reticulitermes split from Heterotermes + Coptotermes at 59.5 Ma (49.9–69.5 Ma 95% CI), yet the oldest split within Reticulitermes (Eurasia and North America) is 16.1 Ma (13.4–19.5 Ma) and the oldest split within Heterotermes + Coptotermes is 36.0 Ma (33.9–40.5 Ma). We detected 14 disjunctions between biogeographical realms, all of which occurred within the last 34 Ma, not only after the break-up of Pangaea, but also with the continents in similar to current positions. Land dispersal over land bridges explained four disjunctions, oceanic dispersal by wood rafting explained eight disjunctions, and human introduction was the source of two recent disjunctions. These wood-eating termites, therefore, appear to have acquired their modern worldwide distribution through multiple dispersal processes, with oceanic dispersal and human introduction favoured by the ecological traits of nesting in wood and producing replacement reproductives. © 2016 The Author(s) Published by the Royal Society. All rights reserved.
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Of the more than 2,300 termite species in the world, 183 species are known to damage buildings and 83 species cause significant damage. Subterranean termites, including mound building and arboreal species, account for 147 (80%) of the economically important species. The genus Coptotermes contains the largest number of pest species (28), whereas the genus Cryptotermes, especially Cr. brevis, is the most widely introduced. The world-wide economic impact figure of termites is uncertain, but the control cost for termite pests in the United States was estimated at $1.5 billion annually in 1994. Because of differences in their life histories, control measures differ between subterranean and drywood species. Insecticide barriers are used for exclusion of soil-borne subterranean termites, whereas slow-acting baits are used for population control of subterranean termite colonies in and near structures. Whole-structure treatments (fumigation and heat), compartmental treatments (heat or cold), and local treatments (wood surface treatments or insecticide injection) are the primary tools for drywood termite control.
The termite family Kalotermitidae constitutes a wood-nesting termite family that accounts for about 15% of all extant termite species. In recent decades, field studies have been carried out to assess termite diversity in various wooded habitats and geographic locations. Three sampling methods have been favored expert, transect, and alate light-trap surveys. Expert collecting is not spatially quantifiable but relies on field personnel to recognize and sample termite niches. The transect method aims to standardize and quantify termite abundance and diversity. Light trapping is a passive method for sampling nocturnal alate flights. We compared our expert survey results and results of published sampling methods for their proportional yields of kalotermitid versus non-kalotermitid encounters. Using an odds ratio statistic, we found that worldwide, there is about a 50.6-fold greater likelihood of encountering a kalotermitid sample versus a non-kalotermitid using the expert survey method and a 15.3-fold greater likelihood using alate trapping than using the transect method. There is about a 3.3 -fold greater likelihood of collecting a kalotermitid specimen versus a non-kalotermitid sample using the expert survey method than using the alate trap method. Transect studies in which only termite species diversity was reported gave similar low Kalotermitidae yields. We propose that multiple biases in sampling methodology include tools, time constraints, habitat type, geographical location, topographical conditions, and human traits account for the divergent outcomes in sampling the abundance and diversity of Kalotermitidae compared to other termite families.