Research Article |
Corresponding author: Robin Casalla ( casallar@uninorte.edu.co ) Academic editor: Eliana Cancello
© 2021 Robin Casalla, Rudolf H. Scheffrahn, Judith Korb.
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:
Casalla R, Scheffrahn RH, Korb J (2021) Rugitermes ursulae (Isoptera, Kalotermitidae), a new drywood termite from the Caribbean coast of Colombia. ZooKeys 1057: 23-36. https://doi.org/10.3897/zookeys.1057.65877
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Rugitermes ursulae sp. nov. is described from a sample collected inside a dead branch in a tropical dry forest of Colombia’s Caribbean coast using molecular information and external morphological characters of the imago and soldier castes. Rugitermes ursulae sp. nov. soldiers and imagoes are the smallest among all described Rugitermes species. The imago’s head capsule coloration is dark castaneous, while the pronotum is contrastingly pale yellow. Our description includes soldier characters, such as subflangular elevation and shape of the antennal sockets, that can help in identification of samples lacking imagoes.
DNA barcoding, imago, northern Colombian coast, soldier, taxonomy, tropical dry forest
Records of termites from Colombia have increased in recent years (
Rugitermes Holmgren, 1911, is a genus from the family Kalotermitidae, mainly found in the Neotropics (
Comparison of head measurements (mm) and coloration of soldiers and imagoes of different Rugitermes species.
No. | Species | Distribution* | Soldier | Imago | Reference | ||||
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Head width | Mean | Head width | Mean | Head color | Pronotum color | ||||
1 | R. ursulae sp. nov. | SA (Colombia) | 1.12–1.22 | 1.17 | 1.10–1.20 | 1.15 | Dark castaneous | Pale yellow | This publication |
2 | R. flavicinctus | SA (Guyana) | 1.19–1.35 | 1.27 | 1.19 | 1.19 | Black | Yellow |
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3 | R. rufus | SA (Bolivia) | 1.16–1.56 | 1.42 | 1.19–1.26 | 1.23 | Reddish |
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4 | R. magninotus | SA (Guyana, Peru) | 1.38–1.48 | 1.43 | 1.25 | 1.25 | Black | Yellow |
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5 | R. volcanensis | SA (Bolivia) | 1.24–1.88 | 1.54 | 1.21 | 1.21 | Black |
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6 | R. athertoni | OC (Polynesia) | 1.47–1.73 | 1.60 | 1.26 | 1.26 | Brown/Black |
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7 | R. aridus | SA (Peru) | 1.44–1.72 | 1.60 | NA | NA | Black | Yellow |
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8 | R. niger | SA (Brazil) | 1.50–1.80 | 1.65 | 1.26–1.34 | 1.30 | Black |
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9 | R. kirbyi | CA (Costa Rica, Panama) | 1.65 | 1.65 | 1.60 | 1.60 | Black | Yellow |
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10 | R. tinto | SA (Colombia) | 1.52–1.90 | 1.72 | 1.31 | 1.31 | Black | Yellow |
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11 | R. unicolor | CA (Honduras) | 1.80 | 1.80 | 1.60 | 1.60 | Yellow-brown | Yellow |
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12 | R. panamae | CA (Panamá) | 1.8–2.0 | 1.90 | 1.4 | 1.40 | Black | Yellow |
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13 | R. bicolor | SA (Guyana) | 1.80–2.07 | 1.94 | 1.51 | 1.51 | Black | Yellow |
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14 | R. occidentalis | SA (Argentina) | 2.00 | 2.00 | 1.35 | 1.35 | Black | Brown |
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15 | R. laticollis | SA (Bolivia, Ecuador) | 1.83–2.43 | 2.06 | 1.5–2.00 | 1.68 | Black |
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16 | R. costaricensis | CA (Costa Rica) | 2.10 | 2.10 | 1.80 | 1.80 | Yellow-brown | Yellow |
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17 | R. nodulosus | SA (Brazil) | NA | NA | NA | NA | Black | Yellow |
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18 | R. rugosus | SA (Brazil) | 1.21 (1.7?) | NA | NA | NA | Black |
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Rugitermes species have few species-specific diagnostic characters. The dorsal antennal ridge and the anterolateral corner of the frontal ridge of the soldier head, the size of the eyes of imagoes and soldiers as well as the imago’s head shape can provide useful information to describe a new species (Krishna et al. 1961;
Here, we describe the soldiers and imagoes of Rugitermes ursulae sp. nov. from a sample collected inside a dead branch in the tropical dry forest of Colombia’s Caribbean coast. In addition, we performed molecular analyses based on the marker COII (cytochrome oxidase II) and including representatives of other genera of Kalotermitidae to support species description.
In July 2014, a survey was done in the tropical dry forest of Colombia’s Caribbean coast (
Rugitermes ursulae sp. nov. was compared with Rugitermes samples from the University of Florida Termite Collection (UFTC), USA, and with descriptions and measurements from the literature (
We considered COII sequences for 26 species of Kalotermitidae (if available three species per genus) and the woodroach, Cryptocercus punctulatus, as an outgroup (Suppl. material
We inferred a phylogenetic tree based on the maximum likelihood (ML) approach. We selected the best fitting model using ModelFinder (
We calculated p-distances between COII sequences with MEGA X v.10.1.8 (
Specimens were suspended in hand sanitizer and images were taken with a Leica M205 C stereomicroscope coupled to a Leica MC190 HD digital camera. Helicon Focus software was used to stack pictures. Measurements were done following
Holotype
soldier. Colombia: Colosó, Sucre (9.5435, -75.34884), 400 meters a.s.l., 11.JUL.2014, R. Casalla, ADD-2014-10A. Paratypes. One additional soldier, 12 pseudergates, a pair of reproductive, same colony sample as holotype ADD-2014-10B. Voucher specimen are held at the Universidad del Norte, Colombia. Holotype soldier (ADD-2014-10A) and one reproductive paratype of Rugitermes ursulae sp. nov. (ADD-2014-10B-1) will be deposited at the Natural History Museum of the Alexander von Humboldt Institute of Bogotá (MIAvH, Colombia) and a paratype soldier (ADD-2014-10B-2) and reproductive (ADD-2014-10B-3) at the collection of the American Museum of Natural History, New York. (
The soldier of R. ursulae sp. nov. is the smallest of all congeneric soldiers (Fig.
Character | Holotype | Measurements |
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Head length with mandibles | 2.81 | 2.81, 2.90 |
Head length to lateral base of mandibles | 1.87 | 1.87, 1.99 |
Head width max. | 1.12 | 1.12, 1.22 |
Head height with postmentum | 1.06 | 1.06, 1.20 |
Postmentum width min. | 0.22 | 0.22, 0.24 |
Postmentum length | 1.34 | 1.34, 1.38 |
Pronotum width | 1.27 | 1.27, 1.39 |
Pronotum length | 0.63 | 0.63, 0.70 |
Third antennal article length | 0.16 | 0.16, 0.17 |
Left mandible length (from dorsal condyle) | 1.17 | 1.17, 1.22 |
Frontal angle (° degrees) | 34 | 29, 34 |
The imago of R. ursulae sp. nov. is also the smallest of all Rugitermes species (Fig.
Measurements (mm) of Rugitermes ursulae sp. nov. dealated imagoes (N = 2).
Character | Measurements |
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Eye diameter max. | 0.30, 032 |
Ocellus diameter max. | 0.09, 0.10 |
Head width (max. with eyes) | 1.10, 1.20 |
Head length to tip | 1.44, 1.49 |
Head height | 0.78, 0.87 |
Pronotum width max. | 1.25, 1.27 |
Pronotum length min. | 0.64, 0.70 |
“Los primates” Colosó, Sucre, Colombia (Fig.
Soldier
(Fig.
Imago
(Fig.
‘Los Primates’ is located in the mountains of the municipality of Colosó, Sucre. It is a regional forest reserve created in 1983, containing primary and secondary tropical dry forest. The mean annual temperature is 26.7 °C (min: 25.8 °C; max: 27.8 °C) with an annual precipitation of around 1337 mm (INDERENA, 1983;
“Ursulae” derived from a diminutive of the Latin ursa, which means “little bear”, in line with the small size of the species. Ursula is also the name of José Arcadio Buendía’s wife in the novel “One hundred years of solitude” written by Gabriel García Márquez and represents an apology/symbolism for the spiritual engine, entrepreneurship, and hard and silent work of many women around the world.
The topology and splits inferred from the multiple sequence alignment of the COII fragment for all Kalotermitidae genera available in NCBI, and including our new species, revealed a COII ML gene tree that clearly separated R. ursulae sp. nov. from the two other Rugitermes species with maximal BS support. Furthermore, it suggests that the genus Rugitermes is monophyletic (maximal BS support) and that it is the sister taxon of Postelectrotermes, however support values for the latter are low (19% BS; Fig.
Maximum likelihood (ML) tree inferred from COII mtDNA gene sequences of 30 species of Kalotermitidae, including the woodroach, Cryptocercus punctulatus, as outgroup. Numbers before the splits show statistical bootstrap support (BS). Terminals are labeled with the respective NCBI access numbers and genus or species name. Rugitermes ursulae sp. nov. is shown in bold.
The p-distance analyses revealed that the barcode sequences most similar to R. ursulae (accession number: MW600961) belonged to Rugitermes sp. A TB-2014 (accession number: KP026284.1) and Rugitermes ADD 2015-29 (accession number: MW600962); they shared 87% and 86% sequence similarity (p-distance), respectively (Suppl. material
1 | Soldier with dark coloration of anterior head capsule. Anterolateral corners of frontal ridges project to form acute angles. Maximum soldier head width (mean) 1.72 mm; imago with a black head capsule; pronotum brownish orange, compound eye small, nearly circular. Ocellus very small; maximum head width (mean) 1.31 mm | Rugitermes tinto |
– | Soldier with light yellowish orange of anterior head capsule. Anterolateral corners poorly developed. Maximum soldier head width (mean) 1.17 mm; imago with a dark-castaneous head capsule; pronotum pale yellowish, compound eye large, nearly circular. Ocellus small, oval shape; head width (mean) 1.15 mm | Rugitermes ursulae sp. nov. |
The Caribbean Region of Colombia is rich in Kalotermitidae and the tropical dry forest supports a high species diversity for this family (
The phylogenetic relationships within the Kalotermitidae are not clearly resolved among the most recently discovered genera (
Molecular markers often allow a clear separation between Rugitermes congeners (
Our study shows the importance of further surveys at isolated sites in the tropics as they continue to reveal many new species. This is also essential for phylogenetic studies to infer the evolutionary history of the Kalotermitidae, and any taxonomic lineage in a broad way.
We thank the Universidad del Norte Barranquilla, Colombia, for a research grant to RC as part of its Strategic Research Area in Biodiversity, Ecosystem Services and Human Well-being, the University of Freiburg, and COLCIENCIAS-Colfuturo for financial support. We are also grateful to the National Agency of Environmental Licenses for research permit no. 739/ANLA/MADS (8 July 2014). We thanks to John Warner for English editing, Karen Meusemann for phylogenetic analyses and discussions, and two reviewers and the editor for helpful comments and careful review of the manuscript.
Tables S1, S2
Data type: Sequence accession identifier and Measures of genetic distance
Explanation note: Table S1. GenBank accession numbers for COII gene sequences of Kalotermitidae used in this study. Table S2. Nucleotide p-distances for COII sequences between Rugitermes ursulae sp. nov. (bold) and other species belonging to Kalotermitidae. Cryptocercus punctulatus used as outgroup.
Figure S1
Data type: Multimedia: Photo
Explanation note: Biome for Rugitermes ursulae sp. nov. (Tropical dry forest, Colosó, Colombia, July 2014).
Supplementary references
Data type: References