Research Article |
Corresponding author: Tiago F. Carrijo ( tiagocarrijo@gmail.com ) Corresponding author: Daniel Castro ( jcastro@sinchi.org.co ) Corresponding author: Rudolf H. Scheffrahn ( rhsc@ufl.edu ) Academic editor: Fred Legendre
© 2023 Tiago F. Carrijo, Daniel Castro, Menglin Wang, Joice P. Constantini, Thomas Bourguignon, Eliana M. Cancello, Yves Roisin, Rudolf H. Scheffrahn.
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.
Citation:
Carrijo TF, Castro D, Wang M, Constantini JP, Bourguignon T, Cancello EM, Roisin Y, Scheffrahn RH (2023) Diminishing the taxonomic gap in the neotropical soldierless termites: descriptions of four new genera and a new Anoplotermes species (Isoptera, Termitidae, Apicotermitinae). ZooKeys 1167: 317-352. https://doi.org/10.3897/zookeys.1167.100001
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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.
Enteric valve armature, Linnean shortfall, mitogenome sequencing, soil-feeder, species distribution, taxonomy
Termites are dominant fauna of tropical ecosystems, having a high abundance, density, and biomass, especially in tropical soils (
Soil-feeding, the ability to digest intractable organic elements from the soil, is exclusive to the family Termitidae (
Identification of termite workers requires examination of gut morphology, especially that of the enteric valve armature (EVA), one of the most important diagnostic characters for them (
Currently, 57 species belonging to 17 genera of soldierless termites are recognized in the Neotropics (
In this paper, we describe four monotypic soldierless genera and a new Anoplotermes species based on the morphology of the worker and imago castes. We also provide a molecular phylogenetic analysis based on the complete mitogenome, including all but two neotropical Apicotermitinae genera. Finally, an illustrated dichotomous key for the worker caste is provided based on the diagnostic characters of the genera published to date.
The specimens examined in this study were collected and preserved in 75%, 85%, or 92% ethanol and were deposited at the Museu de Zoologia da Universidade de São Paulo, São Paulo, Brazil (
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
Molecular phylogenetic analyses were performed on a dataset including the four new genera in addition to 15 of the 17 previously described genera of Apicotermitinae from the New World (Fig.
The alignment of the 13 mitochondrial protein-coding genes and the two ribosomal RNAs was performed separately with the Mafft Multiple Alignment plugin (
Distribution maps (Fig.
Anoplotermes Müller, 1873
Holotype. Worker from lot
Brazil. Rondônia, Porto Velho, Mutum-Paraná, -9.6375, -65.0567.
Paratypes. Brazil. Rondônia, Porto Velho, Abunã, (-9.6257, -65.4423), 15.MAY.2010, 122 m, TF Carrijo and MM Rocha coll. (
The EVA has a “star-like” plaque attached to only one cushion; all cushions are striated. The cushions are adorned with a few small spines in the proximal portion (P1 junction). The mixed segment has an inflated mesenteric tongue.
Imago
(Fig.
Measurements of imagoes (range and mean in mm). L: length; W: width; c = number of colonies; n = number of individuals.
A. susanae ♀ (c = 2, n = 3) | A. susanae ♂ (c = 1, n = 1) | H. kanzakii ♂ (c = 1, n = 1) | K. daironi ♀ (c = 1, n = 2) | K. daironi ♂ (c = 1, n = 1) | O. giblinorum ♀ (c = 1, n = 3) | O. giblinorum ♂ (c = 1, n = 2) | |
---|---|---|---|---|---|---|---|
Head length to lateral base of mandibles | 0.41–0.43 (0.44) | 0.37 | 1.10 | 0.46–0.48 (0.47) | 0.55 | 0.6–0.63 (0.61) | 0.63–0.68 (0.65) |
Maximum head width | 0.42–0.44 (0.43) | 0.42 | 1.42 | 0.50–0.54 (0.52) | 0.55 | 0.78–0.8 (0.79) | 0.73–0.75 (0.74) |
Inter-eye distance | 0.40–0.41 (0.40) | 0.04 | 1.27 | 0.34–0.37 (0.36) | 0.54 | 0.73–0.75 (0.74) | 0.70–0.73 (0.71) |
Max. diameter ocellus | 0.07–0.08 (0.07) | 0.07 | 0.15 | 0.07–0.08 (0.07) | 0.12 | 0.13–0.13 (0.13) | 0.10–0.11 (0.10) |
Min. diameter ocellus | 0.05–0.06 (0.06) | 0.05 | 0.10 | 0.05–0.05 (0.05) | 0.09 | 0.09–0.10 (0.10) | 0.08–0.09 (0.08) |
Max. diameter eye | 0.19–0.20 (0.19) | 0.17 | 0.37 | 0.19–0.22 (0.21) | 0.25 | 0.25–0.28 (0.26) | 0.26–0.28 (0.27) |
Min. diameter eye | 0.15–0.17 (0.16) | 0.15 | 0.35 | 0.16–0.16 (0.16) | 0.20 | 0.23–0.25 (0.24) | 0.23–0.24 (0.23) |
Pronotum length | 0.27–0.30 (0.28) | 0.27 | 0.77 | 0.34–0.35 (0.34) | 0.37 | 0.48–0.50 (0.49) | 0.45–0.48 (0.46) |
Pronotum width | 0.41–0.45 (0.43) | 0.42 | 1.35 | 0.42–0.43 (0.42) | 0.55 | 0.73–0.78 (0.76) | 0.71–0.73 (0.72) |
Hind tibia length | 0.45–0.47 (0.47) | 0.37 | 1.725 | 0.35–0.40 (0.37) | 0.70 | 0.98–1.05 (1.01) | 0.98–1.05 (1.01) |
Fore tibia length | 0.37–0.37 (0.37) | 0.37 | 1.325 | 0.36–0.38 (0.37) | 0.55 | 0.75–0.75 (0.75) | 0.70–0.70 (0.70) |
Fore tibia width | 0.06–0.07 (0.07) | 0.07 | 0.19 | 0.07–0.07 (0.07) | 0.12 | 0.13–0.13 (0.13) | 0.13–0.13 (0.13) |
Fore tibia W/L ratio | 0.17–0.20 (0.19) | 0.20 | 0.14 | 0.19–0.19 (0.19) | 0.23 | 0.173–0.173 (0.17) | 0.185–0.185 (0.18) |
Worker
(Fig.
Measurements of workers (range and mean in mm). LH bm: length of head to lateral base of mandibles; WH max: maximum width of head; LHT: length of hind tibia; LPT: length of protibia; WPT: width of protibia; RPT: ratio WL of protibia. c = number of colonies; n = number of individuals.
Species | LH bm | WH max | LHT | LPT | WPT | RPT |
---|---|---|---|---|---|---|
Anoplotermes susanae (c = 3, n = 10) | 0.26–0.41 (0.35) | 0.43–0.60 (0.50) | 0.30–0.46 (0.36) | 0.29–0.40 (0.33) | 0.09–0.14 (0.10) | 0.22–0.38 (0.32) |
Hirsutitermes kanzakii (c = 8, n = 21) | 0.53–1.40 (0.83) | 0.90–1.28 (1.13) | 0.61–1.28 (0.91) | 0.68–1.05 (0.85) | 0.16–0.29 (0.22) | 0.20–0.31 (0.25) |
Krecekitermes daironi (c = 10, n = 35) | 0.26–0.55 (0.40) | 0.57–0.63 (0.61) | 0.32–0.47 (0.40) | 0.35–0.42 (0.39) | 0.10–0.12 (0.11) | 0.24–0.33 (0.29) |
Mangolditermes curveileum (c = 9, n = 56) | 0.73–0.99 (0.82) | 0.75–1.40 (1.13) | 0.45–1.05 (0.76) | 0.4–0.91 (0.68) | 0.12–0.27 (0.21) | 0.25–0.41 (0.32) |
Ourissotermes giblinorum (c = 3, n = 10) | 0.58–0.70 (0.64) | 0.70–0.80 (0.76) | 0.42–0.80 (0.69) | 0.48–0.65 (0.60) | 0.17–0.23 (0.19) | 0.28–0.37 (0.32) |
Gut (Fig.
Anoplotermes susanae can be distinguished from all other Anoplotermes species by the “star-like” plaque of the EVA. Although Anoplotermes parvus also has a sclerotized plate on the EVA, it forms an asymmetrical spiny mass (Fig.
The phylogeny recovered this species as the sister group of the four other Anoplotermes species (including the type species) included in this study, except A. meridianus, which was recovered as the sister group of Humutermes. The EVA and COI sequence alone suggested this species should be placed in a new genus; however, the external and gut morphology, and the complete mitogenome, corroborate it as a new Anoplotermes species.
This species is distributed in Ecuador, Brazil, and French Guiana (Fig.
This species is named in honor of Susan G. Scheffrahn, wife of RHS.
Hirsutitermes kanzakii sp. nov.
Enteric valve armature (EVA) embellished with hundreds of thin spines on the posterior rim. The EVA seating is tubular, trilobed, and long. The mesenteric tongue is long but not inflated.
Imago
(Fig.
Worker
(Fig.
The EVA of Hirsutitermes is closest to that of Longustitermes; however, the spines of Hirsutitermes are longer and thinner than those of Longustitermes and the anterior portion of the cushions is more rounded in the latter than pyriform in the former. In addition, the external morphology of these two genera is very different, Longustitermes is smaller in size and has a trilobed EVS, while Hirsutitermes has a tubular EVS.
The position of the genus is not clear. In the BI analysis, Hirsutitermes was recovered as the sister group of the clade composed of Krecekitermes and Anoplotermes with a posterior probability of 0.96 (Fig.
From the Latin hirsutus, meaning hairy, rough, refers to the hirsute appearance of the EVA armature.
Holotype. Worker from lot
Colombia. Caquetá, Florencia, Palmichar, 1.7145, -75.6148.
Paratypes. Bolivia. Beni, Puerto Leigue, (-14.2126, -64.9402), 29.MAY.2013, 149 m, TF Carrijo, JA Chase, R Constantino, JR Mangold, A Mullins, J Křeček, T Nishimura, and RH Scheffrahn coll. (
As described for the genus.
Imago
(Fig.
Worker
(Fig.
Gut (Fig.
See remarks for the genus.
Hirsutitermes kanzakii is distributed from Bolivia to Trinidad and Tobago. The material examined for this study did not include samples from Ecuador, but
This species is named in honor of Dr. Natsumi Kanzaki, a Japanese nematologist who has a keen interest in the nematodes of termites.
Krecekitermes daironi sp. nov.
Dehiscent organs fill the hemocoel anterior to the crop. The EVS forms three large lobes. The Mesenteron/P1 junction is well marked, but the mixed segment is short. The EVA armature of Krecekitermes is composed of cushions with six sclerotized and crown-shaped termini adorned with thorns.
Imago
(Fig.
Worker
(Fig.
Dissimulitermes and Ruptitermes present dehiscent organs like Krecekitermes; however, Ruptitermes does not present armature in the EVA, except for R. bandeirai, but this species is much larger, and the spines of the EVA armature are concentrated in only three cushions. Dissimulitermes presents an EVA armature with sclerotized plates and Krecekitermes an EVA armature formed only by spines, without plates. Enteric valve seating (EVS) is tubular in Dissimulitermes and is trilobed in Krecekitermes.
This genus was recovered as the sister group of Anoplotermes, in a clade composed of Hirsutitermes + Krecekitermes + Anoplotermes (Fig.
We named this genus in honor of Dr. Jan Křeček, a retired Czech terminologist who, along with RHS, contributed to many termite descriptions while at the University of Florida, Ft. Lauderdale R.E.C.
Holotype. Worker from lot
Brazil. Rondônia, Porto Velho, Mutum-Paraná, -9.4513, -64.8439.
Paratypes. Bolivia. Cochabamba, Chapare, Villa Tunari, (-16.9704, -65.2100), 26.MAY.2013, 247 m, TF Carrijo, JA Chase, R Constantino, JR Mangold, A Mullins, J Křeček, S. Kuswanto, T Nishimura, and RH Scheffrahn coll. (
As described for the genus.
Imago. (Fig.
Worker
(Fig.
Dehiscent organs present, visible in most individuals. Gut (Fig.
See remarks for the genus.
This species is distributed from Bolivia to French Guiana. Krecekitermes was found in Amazonian and Pacific Forest (Fig.
This species is named in honor of the late Dairon Cárdenas (1957–2022), a Colombian botanist, co-founder of the Colombian Amazon Herbarium COAH of the SINCHI institute, who greatly contributed to the knowledge of the Amazon plants.
Mangolditermes curveileum sp. nov.
In the left view, P1 makes a curve dorsally. The enteric valve seating (EVS) is weakly trilobed. The EVA is unarmed and composed of six elongated and inflated cushions, with well-marked fringed pentagonal scales in the distal half of the cushions. On the proximal ends, each cushion has about 10–20 tiny triangulate spines.
Imago. Unknown.
Worker
(Fig.
Mangolditermes resembles Aparatermes, including the P1 shape, but can be distinguished by: 1) the mesenteric tongue (MT, mixed segment) morphology, that is elongated in the new genus and short in Aparatermes; 2) EVA with the pointed scales in the proximal portion of the cushions in Mangolditermes and towards the central portion of the cushion in Aparatermes; and 3) by the relatively inflated protibia of Mangolditermes. Mangolditermes has unarmed EVA, as Hydrecotermes and Rustitermes, but it can be separated from the latter two by its very long S-shaped P1 in left view. In ventral view, the mesenteric tongues of unarmed Anoplotermes have a whitish spherical mesenteric nodule, that is absent in Mangolditermes. Also, unarmed Anoplotermes have spatulate EVA cushions, while in Mangolditermes they are trapezoid and truncated at the base. Disjunctitermes can easily be confused with Mangolditermes; however, Mangolditermes is larger (WH max >0.75 mm) than Disjunctitermes (WH max < 0.7 mm); Mangolditermes EVA cushions are wider, while Disjunctitermes cushions are narrower, giving an elongated appearance. The cushions of the Disjunctitermes have very marked scales throughout the entire cushion, while in Mangolditermes the scales are only marked from the central to the anterior region of the cushion. Tonsuritermes is closely related to Mangolditermes, but Tonsuritermes workers have a very prominent fontanelle.
This genus was recovered as the sister group of Tonsuritermes, with which it forms the sister group of a large clade composed of Dissimulitermes, Disjunctitermes, Ourissotermes, Tetimatermes, Aparatermes, Compositermes, and Ruptitermes arboreus. The last four form a well-corroborated group, but all the others should be interpreted as a polytomy.
We named this genus in honor of Dr. John Mangold, a retired American terminologist who collected and contributed to many Neotropical termite studies.
Holotype. Worker from lot
Brazil. Rondônia, Porto Velho, Abunã, -9.6089, -65.3769.
Paratypes. Bolivia. Santa Cruz, Ñuflo de Chaves, (-16.4935, -62.6529), 28.MAY.2013, 299 m, TF Carrijo, JA Chase, R Constantino, JR Mangold, A Mullins, J Křeček, T Nishimura, and RH Scheffrahn coll. (
As described for the genus.
Imago. Unknown.
Worker
(Fig.
Gut (Fig.
See remarks for the genus.
This species is widely distributed in South America and the Caribbean region. It is present from Paraguay to Trinidad and Tobago. It is also recorded from the Amazonian, Cerrado, and Chaco forests (Paraguay) (Fig.
The epithet curveileum is a compound noun formed from the words curved and ileum in reference to the curved P1 (the ileum), visible in the left view of the digestive tract.
Ourissotermes giblinorum sp. nov.
Enteric valve armature composed of six pectinate bunches with 4–6 rows of spines, distal portion (proximal to P3) with longer spines, decreasing posteriorly; smaller spines tooth-like. Proximal portion of cushions with a small spine tooth-like (near P1). Enteric valve (EV) inserted directly into the paunch (P3), without an enteric valve seating (EVS) differentiated.
Imago. (Fig.
Worker. (Fig.
Worker digestive tracts. From left to right: ventral, right, dorsal, and left views A Anoplotermes susanae sp. nov. B Hirsutitermes kanzakii sp. nov. C Krecekitermes daironi sp. nov. Abbreviations: C = crop, DO = Dehiscent organ, MS = mesenteron, MT = mesenteric tongue (mixed segment), P1 = ileum, EVS = enteric valve seating, P3 and P3b = paunch, P4 = colon, P5 = rectum.
Worker digestive tracts A Mangolditermes curveileum sp. nov. B Ourissotermes giblinorum sp. nov. From left to right: ventral, right, dorsal, and left views. Abbreviations: DO = dehiscent organ, C = crop, MS = mesenteron, MT = mesenteric tongue (mixed segment), P1 = ileum, EVS = enteric valve seating, P3 and P3b = paunch, P4 = colon, P5 = rectum.
Enteric valve armature of Ourissotermes is similar to Patawatermes, and Echinotermes EVA, but Ourissotermes can be easily differentiated because EVA is inserted directly in P3, without an EVS extension, while Patawatermes EVS is a tubular extension clearly visible externally; Echinotermes present a trilobed EVS. Ourissotermes present a digestive tract external morphology similar to that of Hydrecotermes; however, the EVA of Hydrecotermes is unarmed.
This genus was recovered as part of a big clade composed of Dissimulitermes, Disjunctitermes, Tonsuritermes, Mangolditermes, Tetimatermes, Aparatermes, Compositermes, and the species Ruptitermes arboreus. The last four form a well-corroborated group, but all the others should be interpreted as a polytomy.
The species name is due to the resemblance of the EV armature with the body spines of a sea urchin, that the translation to Portuguese is “ouriço”.
Holotype. Worker from lot
Brazil. Rondônia, Porto Velho, Jaci-Paraná, -9.4526, -64.3900.
Paratypes. Bolivia. Cochabamba, Chapare, Villa Tunari, (-17.0024, -65.4356), 26.MAY.2013, 332 m, TF Carrijo, JA Chase, R Constantino, JR Mangold, A Mullins, J Křeček, T Nishimura, and RH Scheffrahn coll. (
As described for the genus.
Imago. (Fig.
Worker. (Fig.
Gut (Fig.
Worker enteric valves A–D Mangolditermes curveileum sp. nov. from French Guiana (G18-178), Peru (
Apicotermitinae worker enteric valves A Hydrecotermes arienesho B Hydrecotermes kawaii C Humutermes noiroti D Anoplotermes banksi E Anoplotermes meridianus F Anoplotermes janus G Anoplotermes parvus H Longustitermes manni I Patawatermes nigripunctatus J Patawatermes turricola K Rubeotermes jheringi L Grigiotermes hageni. Scale bars: 100 μm.
See remarks for the genus.
This species was mainly collected from soil and decomposing wood pieces. It was found mainly in Amazon and Atlantic primary forests with a high state of conservation, at sites below 504 m a.s.l. This species was recorded as Anoplotermes-group sp. T in
This species is named in honor of Dr. Robin M. Giblin-Davis and his son Sean Giblin. From 1985 to 2020, “Rob” was a former colleague of RHS at the University of Florida, Ft. Lauderdale R.E.C., until his retirement. He taught RHS to appreciate nematodes in termites. For many years, Sean cleaned debris from thousands of termite vials upon the return of collection expeditions by RHS and colleagues listed in the “materials examined” section herein for
Be aware before using this key: Many soldierless termite genera remain undescribed and are not included in this key. So, if your sample does not fit in any genus here, do not try to force a name to it. Using the EVA to confirm the identification is always recommended.
1 | Fontanelle large, sunken, diameter one quarter to three quarters head width. [see |
Tonsuritermes |
– | Fontanelle smaller, diameter less than one quarter head width. (Figs |
2 |
2 | Fore tibia with long “scooped out” excavation [see |
Tetimatermes |
– | Fore tibia without long “scooped out” excavation | 3 |
3 | Enteric valve inserted directly in P3 (EVS not conspicuous in intact gut), P3 without tubular extension to P1 (Fig. |
4 |
– | Enteric valve seating visible in intact gut (Figs |
5 |
4 | Enteric valve armed (Fig. |
Ourissotermes gen. nov. |
– | Enteric valve unarmed (Fig. |
Hydrecotermes |
5 | Junction of EVS/P3 with ring of 15–20 pectinate paddles [see |
Compositermes |
– | Junction of EVS/P3 without ring of 15–20 pectinate paddles | 6 |
6 | Mesenteric tongue (MS/P1 junction) with a whitish spherical mesenteric nodule in ventral view (Fig. |
7 |
– | Mesenteric tongue without a nodule in ventral view (Figs |
8 |
7 | Enteric valve armature consists of six palmate pads (Fig. |
Humutermes |
– | Enteric valve armature with one to three sclerotized pads or enteric valve unarmed (Figs |
Anoplotermes |
8 | Dehiscent organs present (Fig. |
9 |
– | Dehiscent organs absent (Fig. |
11 |
9 | Enteric valve unarmed; workers head pale yellow to black [See |
Ruptitermes
|
– | Enteric valve armed; workers head whitish to pale yellow | 10 |
10 | Enteric valve seating tubular, EVA armature pads with more than 20 spines [see |
Dissimulitermes |
– | Enteric valve seating trilobed (Fig. |
Krecekitermes gen. nov. |
11 | Enteric valve unarmed | 12 |
– | Enteric valve armed | 15 |
12 | Mesenteric tongue spheroidal or elongated (Fig. |
13 |
– | Mesenteric tongue short (drop shaped), not spheroidal (Fig. |
14 |
13 | Cushion scales are only marked from the central to the anterior portion of the cushion (Fig. |
Mangolditermes gen. nov. |
– | Cushion scales markedly throughout the entire cushion [see |
Disjunctitermes |
14 |
EVA Cuticle between cushions with scales; EVS not trilobed [see |
Rustitermes |
– |
EVA Cuticle between cushions without scales; EVS trilobed [see |
Aparatermes |
15 |
EVA armature resembling a continuous (spines on cuticle between cushions) wreath of hundreds of tiny and thin spines (Figs |
16 |
– | EVA armature in form of comb, pads, or sclerotized plates on EVA cushions, not continuous on cuticle between cushions | 17 |
16 | Enteric valve seating tubular (Fig. |
Hirsutitermes gen. nov. |
– | Enteric valve seating trilobed (Fig. |
Longustitermes |
17 |
EVA armature sclerotized plates, pads or comb present only in five cushions (Fig. |
Rubeotermes |
– | EVAA sclerotized plates, pads or comb present in all the six cushions | 18 |
18 | Enteric valve seating tubular (Fig. |
19 |
– | Enteric valve seating trilobed (Fig. |
20 |
19 | Distal end of EVA armature pads capped with even tuft of spines; EVS elongated [Fig. |
Patawatermes |
– | Distal end of EVA armature pads ringed by unevenly arranged spines; EVS short [Fig. |
Grigiotermes |
20 |
EVA armature pads spiny, spherical [see |
Echinotermes |
– |
EVA armature pads raspy, truncated [see |
Amplucrutermes |
The systematic and taxonomic delineation of Neotropical soldierless termites has been implemented by various methods (
A few groups showed high branch support values and deserve a note. The genera Grigiotermes and Patawatermes (originally described as Grigiotermes) form a clade and share many morphological characters. If future analysis corroborates the results found here, reverting the Patawatermes species to the genus Grigiotermes may be a parsimonious taxonomic decision. A group composed of Compositermes, Aparatermes, Tetimatermes, and the species Ruptitermes arboreus seems to be a very consistent group, despite the lack of morphological synapomorphies, this clade was also recovered in recent works using the COI gene (e.g.,
Remarkable diagnostic morphological characters in New World Apicotermitinae occur in a few genera (e.g., Compositermes, Tonsuritermes, Tetimatermes), but there is a lack of remarkable diagnostic characters for differentiating other genera. For example, Anoplotermes and Humutermes share a similar digestive tract, and these genera can only be supported by molecular data. This difficulty becomes even clearer in our phylogeny, where the species Anoplotermes meridianus clustered with Humutermes, making Anoplotermes paraphyletic. In addition, the new species Anoplotermes susanae sp. nov. clustered with the Anoplotermes “stricto sensu”, but with low branch support. Using the barcode gene COI alone (unpublished data), this species was first hypothesized as a new genus. The inclusion of more genetic data, such as nuclear genes, will be important to corroborate our results here. Likewise, we emphasize the large number of species whose type material was not yet examined in recent studies, and synonyms can be generated in the future.
Another critical challenge occurs within species with unarmed EVA, where the characters are even more inconspicuous, making species identification more difficult. In this study, we identified morphospecies records from pictures from other studies (
Despite the large number of descriptions of new taxa of soldierless termites from the Neotropics in the last decade (
The soldierless termites present not only a taxonomic gap (Linnean shortfall), but also a geographical gap (Wallacean shortfall) (
The key that we provide can serve as a support tool for the confirmation of identification of genera that are already described; however, as we discuss here, there is many species not yet described. It is expected that progress in the description of soldierless taxa will highlight the importance of these groups in ecological studies and once ecological studies start naming the soldierless termites, the Wallacean shortfall of these termites will also diminish.
DC thanks to Instituto Amazónico de Investigaciones Científicas SINCHI and to Dr. Clara Peña-Venegas for financing and logistics support in CATAC collection.
The authors have declared that no competing interests exist.
No ethical statement was reported.
This study was partially funded by the São Paulo Research Foundation (FAPESP) through the grants #2020/06041-4 (TFC) and #2014/11982-1 (JPC). YR received support from the Belgian Fonds National de la Recherche Scientifique (F.R.S.-FNRS, grant J.0180.20). EMC received support from the Conselho Nacional de desenvolvimento Científico e Tecnológico (CNPq Proc. Nr. 308408/2019-5).
Conceptualization: RS, TFC, DC. Data curation: YR, TFC, RS, TB, JC, DC, MW. Formal analysis: MW, JC, TFC, DC, RS, EMC. Funding acquisition: YR, TFC, EMC, TB. Investigation: TFC, MW, DC, JC, RS. Methodology: DC, TFC, RS, JC, EMC. Software: DC, TFC, MW. Visualization: MW, TFC, RS, JC, DC, TB. Writing – original draft: TFC, RS, DC. Writing – review and editing: YR, EMC, MW, RS, DC, TFC, TB, JC.
Tiago F. Carrijo https://orcid.org/0000-0001-6308-7252
Daniel Castro https://orcid.org/0000-0002-5060-6129
Menglin Wang https://orcid.org/0000-0003-2206-9503
Joice P. Constantini https://orcid.org/0000-0002-1813-9276
Thomas Bourguignon https://orcid.org/0000-0002-4035-8977
Eliana M. Cancello https://orcid.org/0000-0003-3125-6335
Yves Roisin https://orcid.org/0000-0001-6635-3552
Rudolf H. Scheffrahn https://orcid.org/0000-0002-6191-5963
All of the data that support the findings of this study are available in the main text or Supplementary Information.
Samples used in the phylogenetic analysis
Data type: table (word document)
Maximum Likelihood phylogenetic tree of the New World Apicotermitinae using the complete mitogenome
Data type: figure (word document)