New record of Dolichorhinotermes lanciarius Engel & Krishna, 2007 (Blattodea, Rhinotermitidae) from Colombia

Dolichorhinotermes Snyder & Emerson, 1949, a genus of Neotropical distribution, includes seven living termite species characterized by the presence of distinct major and minor soldiers. In Colombia, to date, only Dolichorhinotermes longilabius (Emerson, 1924) has been recorded in the Colombian Amazon Basin. A new distribution record of Dolichorhinotermes lanciarius Engel & Krishna, 2007 (Rhinotermitidae, Rhinotermitidae) is reported from an Andean region of the Department of Caldas, Colombia, where it was found at an altitude of 2100 m.

The Colombian termite fauna from the Andean region is poorly known and undersampled. In Colombia, only one species of Dolichorhinotermes, D. longilabius, has been recorded at altitudes up to 500 m in the Amazon region (Beltrán and Pinzón 2018;Castro et al. 2021). I report here D. lanciarius for the first time from Colombia, where this species was found in an Andean region of the Department of Caldas at 2100 m of altitude.

Methods
Termites were manually collected from pieces of rotten logs in contact with the ground. Specimens were stored in 85% ethanol and deposited in the Colección Entomológica Forestal CEFUDFJC (RNC 045), and a COI sequence was published in the Barcode of Life Data (BOLD). Measurements and photographs of specimens were made using a Discovery V8 Carl Seizz stereomicroscope utilizing an automated multi-layer system and combined using ZEN or Helicon Focus software. A map showing the current known distribution of D. lanciarius was produced using the ArcGIS software v. 10.6. (Fig. 2).
Dolichorhinotermes lanciolarius was identified using the keys to genera and species (Krhisna et al. 2013;Castro and Scheffrahn 2019) and the original description of D. lanciarius (Engel and Krishna 2007). The sample collected in 1998 did not require a collection permit, while the sample from 2016 was collected under the permit ANLA 2014 0738. Figure   Dolichorhinotermes lanciarius is the largest species in the genus, particularly the soldier caste (Engel and Krishna 2007). In the collected specimens, the length of the major soldier head, including mandibles (2.62 mm), the head width (1.68 mm), and the pronotum width (0.89), coincide with the size of the soldiers from the original description. On the other hand, the minor soldier metrics, such as head length, including the labrum, labrum length, head width, and pronotum length and width, are slightly larger than those metrics given Engel and Krishna (2007). The anterior border and shape of the pronotum of a minor soldier of the Colombian specimens differ from the original description of the species (Engel and Krishna 2007: 4, fig. 2A). While the original description indicates a rounded anterior border, the Colombian samples have a wide anterior border with three lobes (Fig. 1D).

Dolichorhinotermes lanciarius Engel & Krishna, 2007
The dentition of the major soldier mandibles (not shown) is another significant character in the recognition of D. lanciolarius. According to Engel and Krishna (2007: 5): "left mandible with two stout teeth, first just apical of midpoint, second just basal of midpoint, first tooth distinctly longer than second tooth, first tooth slightly curved along posterior surface, second tooth straight; right mandible with two teeth near midpoint, first tooth shorter than second tooth and both shorter than those of left mandible, second tooth extending straight out from inner mandibular surface, first tooth running closer along mandibular surface".
Another feature unique to D. lanciolarius is the shape of the major and minor soldier labra. Major soldiers (Fig. 1A, B) have "labrum elongate, slightly wider at apex, apex rounded, without emargination, with slight dorsal, longitudinal depression extending posteriorly from apical margin about one-half of labral length" (Engel and Krishna 2007: 3), as shown in Figure 1A. In addition, the labrum from the minor soldier exhibits a pronounced apical labral forking, as shown in Figure 1C, and "… several erect, pale yellow setae at apex, otherwise dorsal surface with scattered, short, pale yellow setae, setae slightly more numerous in apical half" (Engel and Krishna 2007: 4). Minor soldier head also presents "gently convex, sides converging anteriorly, and slightly converging from midpoint posteriorly to a gently rounded posterior border than basal half" (Engel and Krishna 2007: 3), as illustrated in Figure 1D. Noteworthy of this species minor soldier is the lance-like labrum shown in Figure 1E.
Alate nymphs are substantially larger than the other castes ( Fig. 1G-I), having the body, legs, head, and antennae yellow with light brown highlights. The head capsule is oval, almost as wide as the pronotum, and covered with abundant long bristles. The moniliform antenna has 20 antennomeres covered with long bristles, being the basal flagellomere distinctly larger than the second. Eyes are dark brown, bordered by clear margins, and there are two light, non-prominent circular ocelli distant from the compound eyes. The clipeus is oblong with a rounded margin and covered by long bristles. The pronotum has rounded borders, wider anteriorly, covered by long sparse bristles and several shorter erect bristles in the anterior border behind the head. In contrast, the posterior border is slightly emarginate at the center. The head width of a single alate nymph is 2.1 mm, the head length to the labrum apice is 2,1 mm, and the pronotum width is 2.5 mm.
Ecological observations. An adult of the genus Peneta sp. (Tenebrionidae, Prenapatinae), living in association with D. lanciarius in a heavily infested log, is shown in Figure 1J.  2017; Vargas et al. 2005), and there are few records from above 2000 m altitude. Here, Dolichorhinotermes lanciarius is reported for the first time in the country, from a mountainous region in the Department of Caldas at 2100 m altitude. This region has a humid climate with average annual precipitation of 2000 mm and a mean annual average temperature of 17 °C.

Distribution
The samples were collected in rotten logs in a remnant of secondary forest surrounded by an area dedicated to livestock and commercial forest plantations. However, D. lanciolarius was not found to be damaging living trees or crops. This is consistent with previous accounts in which D. lanciolarius has not been observed in forest plantations and crops, and is not considered a pest species in Bolivia and Brazil (Constantino 2002;Krishna et al. 2013). In the nearby collecting region, no other species of termites are known to occur to date.
Only one type of soldier of D. lanciarius was collected in each collecting season in Colombia, suggesting that the season influenced the occurrence of the soldier caste. Likewise, fewer major soldiers were found than minor soldiers, coinciding with Engel and Krishna's (2007)

findings in Ecuador.
This new record occurred outside the Amazon region, where most known extant Dolichorhinotermes species occur. The known distribution of D. lanciarius is extended more than 1000 km north and at an altitude greater than 1000 m above that of the type locality of the species.