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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
http://zoobank.org/C6973DAD-84F4-4C54-87D0-4EDFBEDFF161
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. https://doi.org/10.3897/zookeys.787.28195
Abstract
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.
Keywords
Ethiopian Region, mandibles, ocellus, taxonomy
ZooKeys 787: 91–105 (2018)
doi: 10.3897/zookeys.787.28195
http://zookeys.pensoft.net
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.
RESEARCH ARTICLE
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Rudolf H. Scherahn et al. / ZooKeys 787: 91–105 (2018)
92
Introduction
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)
94
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.
Results
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.
Systematics
Roisinitermes Scherahn, gen. n.
http://zoobank.org/9AE40F98-CA9E-45AC-849E-A034F19E8DAE
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)
96
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.
http://zoobank.org/129573FB-E5DE-4673-9E1B-EF062D413FEB
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)
98
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)
100
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.
Discussion
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 gen.et sp. n.
Rudolf H. Scherahn et al. / ZooKeys 787: 91–105 (2018)
102
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.
Acknowledgements
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).
Roisinitermes ebogoensis gen. & sp. n., an outstanding drywood termite... 103
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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
(http://opendatacommons.org/licenses/odbl/1.0/). 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.
Link: https://doi.org/10.3897/zookeys.787.28195.suppl1
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