Proneotermesmacondianus, a new drywood termite from Colombia and expanded distribution of Proneotermes in the Neotropics (Isoptera, Kalotermitidae) Author Casalla, Robin Author Scheffrahn, Rudolf H. Author Korb, Judith text ZooKeys 2016 623 43 60 http://dx.doi.org/10.3897/zookeys.623.9677 journal article http://dx.doi.org/10.3897/zookeys.623.9677 1313-2970-623-43 5E560CB94AE34E8AACBDB8E8DEC0D799 5E560CB94AE34E8AACBDB8E8DEC0D799 Taxon classification Animalia Isoptera Kalotermitidae Proneotermes macondianus sp. n. Diagnosis. The Proneotermes macondianus soldier is smaller and the head capsule lighter than those of Proneotermes latifrons and Proneotermes perezi . In Proneotermes macondianus , the lateral margins of the genal horns angle outward from the sides of the head capsule while, in the other two species, the lateral margins of the genal horns remain in line with the head capsule. The mandibular humps of Proneotermes macondianus are more pronounced and rounded than in Proneotermes latifrons and Proneotermes perezi . Both Proneotermes latifrons and Proneotermes perezi have more robust rugosity on the frons than Proneotermes macondianus . The imago of Proneotermes macondianus is smaller and has much shorter, wider, darker, and more punctate wings than that of Proneotermes perezi . Description. Imago (Figs 2, 3A, Table 2). Head dorsal view: yellowish weakly trapezoidal, eyes moderately protruding and small, diameter 0.30 mm (Fig. 2 A-B ). Ocellus oval and almost touching eye (Fig. 2B). Antenna with 15 articles. Pronotum broader than head (Fig. 2A). Forewing with all major veins running parallel; subcosta running from suture to costal margin about 1/5 length of wing, radius to 1/3 wing length, radial sector with 4-6 branches, media less sclerotized than anterior veins, and cubitus unsclerotized Wings brownish, especially near scale suture, membrane nodular; unusually wide and relatively short. Fore wing with a very long suture line margin; scale much darker that body pigmentation (Fig. 3A). Measurements are reported in Table 2. Figure 2. Proneotermes macondianus sp. n. imago: A Head in dorsal B lateral, and C whole body in lateral. Scale bars: 0.5 mm. Figure 3. Proneotermes macondianus sp. n. Alate: A fore- and hindwing, scale bar: 5 mm. Soldier: B mandibles, scale bar: 1 mm. Table 2. Morphometrical measurements for Proneotermes macondianus sp. n. imagoes.
No. Measurements in mm (n = 10). Mean SD Range
Soldier (Figs 3B, 4, Table 3). Head in dorsal view with postclypeus almost black, grading to ferruginous orange near frontal flange, and yellow at occiput (Fig. 4A). Dorsal view with head elongate and sides parallel, frons wide and shallow and faint frontal flange (Fig. 4B). Eye spots distinct, unpigmented. Mandibles completely black (Fig. 3B, 4 A-C ). Pronotum yellowish with anterior borders brown. Frons angles below vertex approx. 43°. Rugosity vestigial on the frons or vertex regions of the head. Frontal horns robust and project towards the front (Fig. 4A). Genal horns prominent in dorsal view, angled antero-laterally about 45°. Mandible tips bend about 60-65° from longitudinal axis of mandibular blade, prominent dentition, with rounded and pronounced mandibular humps: left hump larger than right (Figs 3B, 4A). Postmentum somewhat constricted in middle, as cup-shaped (Fig. 4C). Third antennal article enlarged and sclerotized, formula 2<3>4=5=6 and 11 articles. Pronotum as broad as head ; anterior emarginate. Measurements are reported in Table 3. The soldiers from the Tayrona National Park (Santa Marta, Magdalena) showed slightly darker coloration than those from the samples sites at El Ceibal (Santa Catalina, Bolivar ) and Coloso ( Coloso , Sucre). Figure 4. Soldier heads of Proneotermes macondianus sp. n. ( A-C ), Proneotermes latifrons ( D-F ), Proneotermes perezi ( G-I ). Head in dorsal (A, D, G), lateral (B, E, H) and ventral position (C, F, I). Arrow in 4C denote genal horn projected prominently in Proneotermes macondianus sp. n. Scale bars: 1 mm. Table 3. Morphometrical measurements for Proneotermes macondianus sp. n. soldiers.
No. Measurements in mm (n = 11). Mean SD Range
Genetic analysis. The COII, 12S, and 16S sequences obtained in this study are deposited in GenBank under accession numbers KX267090-KX267098 (Table 1). The combined COII, 12S, and 16S nucleotide data of three different haplotypes of Proneotermes macondianus from the northern Colombian coast (separated ~200 km from each other) revealed genetic distances of about 2.5% (p-distance, SE 0.004; 38bp / 1488bp) ( Table 4). Our Bayesian phylogenetic tree shows more than 74% Bayesian Posterior Probability (BPP) support for all nodes (Fig. 5). The COII based tree suggests that Proneotermes is the sister taxon to a cluster consisting of [( Marginitermes + Epicalotermes ) + Bifiditermes ] + [( Cryptotermes + Procrypotermes ) + Incisitermes ]. Since Kalotermitid sequences for 12S and 16S were very sparse in public databases such as The Barcode of Life Data Systems (BOLD) or NCBI, we only used COII to estimate genetic distances between species and for phylogenetic tree inference. Inter-generic COIIp-distances for available genera (Table 5) showed that Bifiditermes is closest to Proneotermes (0.153 SE 0.015). Figure 5. Bayesian inference tree, inferred with MRBAYES from COII sequence data (nodes show posterior probability support). Kalotermitidae distribution (X), Species distribution known (O), unknown (?) and established introductions from other regions (I). Neartic = Nea, Neotropic = Neo, Ethiopian = Eth, Paleartic = Pal, Oriental = Ori, Australian = Aus, Papuan = Pap. Distribution based on Krishna et al. 2013 . Table 4. Nucleotide distances for combined analysis of COII, 12S rDNA and 16S rDNA genes between localities of Proneotermes macondianus sp. n. (p-distance). Standard error estimates are shown above the diagonal.
No. Species (n = number of sequenced specimens) Localities CE CO PT
Proneotermes macondianus CE
Proneotermes macondianus CO
Proneotermes macondianus PT
Table 5. Nucleotide distances for COII sequences between sample localities Proneotermes macondianus sp. n, the cockroach Cryptocercus punctulatus and different Kalotermitidae (p-distance). Standard error estimates are shown above the diagonal.
No. Specie 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16
Proneotermes macondianus CEKX267097
Proneotermes macondianus COKX267098
Proneotermes macondianus PTKX267096
Bifiditermes improbus AF189080.1
Postelectrotermes howa EU253883.1
Neotermes castaneus HQ215844.1
Incisitermes immigrans AB109542.1
Epicalotermes mkuzii DQ442125.1
Kalotermes flavicollis DQ442147.1
Glyptotermes brevicornis AF189096.1
Cryptotermes cavifrons FN377810.1
Comatermes perfectus EU253878.1
Procryptotermes leewardensis EU253884.1
Calcaritermes temnocephalus EU253877.1
Marginitermes KJ907844.1
Cryptocercus punctulatus DQ007637.1
Ecological notes. Proneotermes macondianus sp. n was found in tropical dry forests of the Colombian Caribbean near to coastal areas up to 25 km inland (Fig. 1, Appendix 1 - Figure S1). Encounters of Proneotermes were scarce. In line transects that covered a total area of 1500m x 2m, only 0.82% of all termite samples (n = 1102) were Proneotermes macondianus (n = 9). All samples were from thin pieces of drywood branches: less than 2 cm diameter on the ground, with a maximum of 20 individuals per branch. Pellets were hexagonal in shape, beige in colour and had a length of 0.92 +/- 0.04 mm (Appendix 1: Figure S1, S2, Table S1). It was impossible to identify the plant species from the small dry branches where Proneotermes macondianus sp. n. was found. Material examined. Holotype colony: Colombia: Municipality of Santa Marta, Magdalena. Tayrona National Natural Park, Gairaca Bay: 11.3152°N , 74.1032°W (Fig. 1), 6m, 27.VI.15 by R. Casalla. COLPT4K1-206. Holotype: Soldier, paratypes: 5 soldiers, 2 reproductives, and few pseudergates, two used for DNA analysis. Municipality of Santa Catalina, Bolivar . Protected area "El Ceibal": 10.6336°N , 75.2517°W , 25m, 30.VIII.14 R. Casalla. COLCE3F5-155, COLCE3G5-158, COLCE3H2-160: Paratypes: 4 soldiers, 5 functional reproductives, and few pseudergates, two used for DNA analysis. Municipality of Coloso , Sucre. Serrania de Coraza y Montes de Maria . Protected area "Los Primates ": 9.5332°N , 75.3479°W , 223m, 27.VII.14 R. Casalla. COLCO4F4-226: Paratypes: 1 soldier, 1 winged imago, 2 dealated imagoes, one used for DNA analysis. Measurements for holoype, paratype soldiers and imagoes are reported in Table 2 and 3. The holotype and clearly colored paratype soldiers from COLCE3F5-155 will be deposited in the Arthropod Collection of the Natural History Museum of the Alexander von Humboldt Institute of Bogota , Colombia (MIAvH). A paratype soldier from holotype colony, will be deposited in the American Museum of Natural History, New York, United States. Morphotype imagoes, paratype soldiers and pseudergates will be part of the collection of the Department of Chemistry and Biology at the University del Norte, Barranquilla, Colombia. Etymology. Macondianus : In honour of Nobel laureate Gabriel Garcia Marquez and the fictional town "Macondo" in his novel "One hundred years of solitude". "Macondiano/a" is also a Spanish world used in Colombia to describe an incredible, rare or surprising event that could only be compared with the fictional universe and magical realism of this novel.