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Research Article
Cryptotermes pugnus (Blattodea, Isoptera, Kalotermitidae), a new drywood termite species from the Brazilian Caatinga dry forest and key to South American Cryptotermes Banks, 1909
expand article infoRudolf H. Scheffrahn‡, Alexandre Vasconcellos§
‡ University of Florida, Davie, United States of America
§ Universidade Federal da Paraíba, João Pessoa, Brazil

Corresponding author: Rudolf H. Scheffrahn ( rhsc@ufl.edu )

Academic editor: Fred Legendre
© 2023 Rudolf H. Scheffrahn, Alexandre Vasconcellos.
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: Scheffrahn RH, Vasconcellos A (2023) Cryptotermes pugnus (Blattodea, Isoptera, Kalotermitidae), a new drywood termite species from the Brazilian Caatinga dry forest and key to South American Cryptotermes Banks, 1909. ZooKeys 1182: 11-18. https://doi.org/10.3897/zookeys.1182.108243
ZooBank: urn:lsid:zoobank.org:pub:B49C6966-A3DE-415E-9D6A-B9EACEECBA99
Open Access

Abstract

A new termite species, Cryptotermes pugnus sp. nov., is described from northeastern Brazil. The winged imago of C. pugnus is distinguished from most congeners by the lack of arolia and the multiple branches connecting the median vein to the radial sector. The soldier is unique among South American Cryptotermes by its cuboidal head capsule and very rugose postclypeus. The new species constitutes the fourteenth Cryptotermes species on the continent for which we provide a key to soldiers.

Key words

Arolium, Bahia, imago, Paraíba, soldier, South America, venation

Introduction

The cosmopolitan termite genus Cryptotermes Banks, 1906 is most diverse in the Neotropics with 32 of the 72 species described worldwide (Constantino 2020). Three of the Neotropical species are exotic pests, including C. brevis (Walker, 1853) (only the populations outside its endemic region of coastal Chile and Peru), C. dudleyi Banks, 1918, and C. havilandi (Sjostedt, 1900) with a previous fourth, C. domesticus Haviland, 1898, now deemed absent from the New World (Scheffrahn et al. 2009; Scheffrahn 2021). Until now, mainland South America (and Trinidad and Tobago) was habitat to 11 endemic Cryptotermes species: C. aequacornis Scheffrahn & Křeček, 1999; C. brevis; C. camelus Scheffrahn, 2021; C. chacoensis Roisin, 2003; C. colombianus Casalla et al., 2016; C. contognathus Constantino, 2000; C. cubicoceps (Emerson, 1925); C. cylindroceps Scheffrahn & Křeček, 1999; C. mangoldi Scheffrahn & Křeček, 1999; C. rhicnocephalus Bacchus, 1987; and C. verruculosus (Emerson, 1925). Cryptotermes mangoldi and C. cylindroceps were originally described from the West Indies until Casalla et al. (2016) reported their mainland distribution. Here, we describe a new endemic mainland species, C. pugnus sp. nov., from northeastern Brazil and provide a key to the described Cryptotermes from South America.

Material and methods

Photomicrographs were taken as multilayer montages using a Leica M205C stereomicroscope controlled by Leica Application Suite v. 3 software. Preserved specimens were taken from 85% ethanol and suspended in a pool of Purell Hand Sanitizer to position the specimens on a transparent Petri dish background. Comparisons with other South American Cryptotermes species were made from specimens in the University of Florida Termite Collection (Scheffrahn 2019).

Taxonomy

Cryptotermes pugnus Scheffrahn & Vasconcellos, sp. nov.

https://zoobank.org/B7EB068D-34FD-4B36-A5B9-C37A230F05FD

Comparison

The imago of C. pugnus groups with C. brevis, C. chacoensis Roisin, 2003, C. kirbyi Moszkowski, 1955, and C. darwini (Light, 1935) in having the arolium absent between the tarsal claws (Fig. 1C). The forewing venation of the C. pugnus imago is atypical for most of the genus in having several branches splitting from the media and intersecting the radial sector (Fig. 1D). This character is only known from C. brevis, C. darwini (see Light 1935), and C. kirbyi (see Bacchus 1987). The latter two species may be found in future studies to by synonyms of C. brevis. Roisin (2003) did not describe the venation of C. chacoensis.

Figure 1. 

Imago of Cryptotermes pugnus sp. nov. (SA470) A dorsal view of head and pronotum B lateral view of head and pronotum C distal tarsomere of foreleg D left forewing.

Among mainland South American Cryptotermes soldiers, C. pugnus is unique in having, in dorsal view, a cuboidal head capsule and a very rugose, rounded and projecting postclypeus (Fig. 2). The postclypeus of C. brevis and C. chacoensis soldiers are closest to C. pugnus, but the head capsules of the former two are constricted (Fig. 3). Along with C. pugnus, only C. aequacornis, C. cylindroceps, and C. rhicnocephalus have both frontal and genal horns projecting the same length anteriorly (Fig. 3H, L, M).

Figure 2. 

Soldier of Cryptotermes pugnus sp. nov. (SA470) A dorsal view of head and pronotum B lateral view of head and pronotum C oblique view of head and pronotum D ventral view of head and pronotum E lateral view of cephalic horns. FF = frontal flange, FH = frontal horn, and GH = genal horn.

Figure 3. 

South American Cryptotermes soldier head capsules A C. dudleyi (arrow: genal horn) B C. havilandi (arrow: genal horn) C C. colombianus D C. contognathus (arrow: frontal horn) E C. chacoensis (arrow: constriction) F C. brevis (arrow: constriction) G C. cubicoceps (arrow: frontal horn) H C. aequacornis (arrow: postclypeus; bracket: outer span of mandibles) I C. pugnus sp. nov. (arrow: postclypeus; bracket: outer span of mandibles) J C. camelus (arrow: frontal flange) K C. verruculosus (arrow: frontal flange) L C. cylindroceps (arrow: lateral margin of vertex) M C. rhicnocephalus (white arrow: vertex concave, grey arrow: frontal flange without elevated rim) N C. fatulus (black arrow: frontal flange with elevated rim; white arrow: frontal horn) O C. mangoldi (black arrow: frontal flange with elevated rim; white arrow: frontal horn with elevated rim). A, B, F, H, L, M modified from Scheffrahn and Křeček (1999); C modified from Casalla et al. (2016). Scale bars: 1 mm.

Description

Imago (Fig. 1A–D). Head capsule and pronotum pale yellow brown. Compound eye obtusely triangular; ocellus light yellow, about half diameter of eye, roundly ellipsoid, and touching eye margin. Vertex with a few short setae. Pronotum wider than head capsule; anterior margin shallowly concave. Pronotum lateral margins with about one dozen setae each. Antennae with 15 articles, basal article relative lengths 2 = 3 > 4 = 5. Forewing with subcosta joining costal margin at about 1/8 of wing length from suture. Wing membrane pale; veins a shade darker. Costa, subcostal, radius, and radial sector sclerotized; unsclerotized media with several branches intersecting radial sector; media terminating at radial sector about 3/4 wing length, then appearing as a separate branch near tip of wing. Arolium absent. Measurements (mm, mean, n = 3). Head maximum width with eyes 0.96; head maximum width without eyes 0.88; pronotum maximum width 0.94; eye maximum diameter 0.23; ocellus maximum diameter 0.12; total body length 5.3; right forewing length from scale 6.90; body length with wings 8.74.

Soldier (Fig. 2A–E). Head capsule, in dorsal view, strongly rugose; dark castaneous brown from postclypeus grading to orange-brown at occiput. Head capsule widest at posterior third, narrowest at frontal flange. Frontal flange (ridge) V-shaped with deep median cleft. Posterior margin of head capsule truncate, posterolateral corners forming right angles, lateral margins nearly parallel combining to form cuboidal appearance. In lateral view, frontal flange elevated, vertex unevenly concave; frontal horns visible as blunt knobs. Genal horns evenly rounded, slightly posterior to frontal horns (Fig. 2E). Pronotum angled sharply from vertex, narrower than head; anterior margin dark and ruffle; incised in middle with rounded anterior lobes. Eye spots large, narrowly elliptical. In oblique view (Fig. 2C) frons concave. Postclypeus evenly convex, strongly rugose, projecting well beyond frontal flange (Fig. 2A). Antennae with 8 or 9 articles, third fused or divided; or with 10 or 11 articles, third fused or divided. Mandibles wide and short for the genus; rugose, rounded basal hump at half-length when seen from below, outer margin of blade angles about 50°. Measurements (mm, mean, n = 2). Head length to tip of mandibles 1.57; head length to tip genal horns 1.20, frontal flange width 1.11; frontal horns, outside span 0.90; genal horns, outer span 0.95; head width, maximum 1.20; head width, minimum (behind frontal flange) 1.10; head height, excluding postmentum 0.88; pronotum, maximum length 0.95; pronotum, maximum width 1.12; left mandible length, tip to ventral condyle 0.53.

Type materials

Holotype : Brazil • Soldier; Paraíba, São José dos Cordeiros; -7.39056, -36.80833; 526 m a.s.l.; 17 Aug. 2000; A. Vasconcellos leg.; two soldiers (one labelled holotype, Fig. 2), three imagos, and three pseudergates; University of Florida Termite Collection (UFTC) no. SA470, subsample from Federal University of Paraíba Termite Collection (FUPTC) no. 2052. Paratypes: Brazil • Bahia, Curaçá; −9.123, −39.691; 366 m a.s.l.; 4 May 2011; A. Vasconcellos leg.; one soldier and pseudergates; FUPTC no. 4345.

Etymology

Named after the pug dog. The oblique view of the soldier (Fig. 2C) resembles this short-nosed breed.

Key to South American Cryptotermes soldiers

1 In dorsal (or ventral) view, genal horns form anterolateral knobs of head capsule; vertex smooth (introduced species) (Fig. 3A, B) 2
— In dorsal view, genal horns eclipsed by frontal horn or frontal flange (e.g. Fig. 3H) 3
2 Mandibles project more than one third length of head capsule (Fig. 3A) C. dudleyi
— Mandibles project about one fourth length of head capsule (Fig. 3B) C. havilandi
3 Mandibles barely project beyond frons or frontal horns (Fig. 3C, D) 4
— Mandibles clearly project beyond frons or frontal horns (e.g. Fig. 3H) 5
4 Frontal horns not visible (Fig. 3C) C. colombianus
— Frontal horns visible (Fig. 3D) C. contognathus
5 Vertex excavated; with deeply folding rugosity (e.g. Fig. 3H) 6
— Vertex not excavated; rugosity more shallow (e.g. Fig. 3M) 10
6 Head constricted behind frontal flange (Fig. 3E, F) 7
— Head not constricted behind frontal flange (Fig. 3G–I) 8
7 Genal horns visible from above, mandibles with lateral humps; Gran Chaco region (Fig. 3E) C. chacoensis *
— Genal horns not visible from above, mandibles without lateral humps; widespread (Fig. 3F) C. brevis *
8 Frontal horns barely extend beyond anterolateral margin of frontal flange (Fig. 3G) C. cubicoceps
— Frontal horns extend well beyond anterolateral margin of frontal flange (Fig. 3H, I) 9
9 Anterior margin of postclypeus linear; outer span of mandibles <1/2 width of head (Fig. 3H) C. aequacornis
— Anterior margin of postclypeus rounded; outer span of mandibles >1/2 width of head (Fig. 3I) C. pugnus sp. nov. *
10 In lateral view, frontal flange emerges above vertex as a rounded mound (Fig. 3J, K) 11
— In lateral view, frontal flange forms angular intersection with vertex (Fig. 3L–N) 12
11 Frontal flange semicircular; humid Chaco (Fig. 3J) C. camelus
— Frontal flag quadrant (Fig. 3K) C. verruculosus
12 Lateral margin of vertex linear in lateral view (Fig. 3L) C. cylindroceps
— Lateral margin of vertex concave (Fig. 3M–O) 13
13 Flange without elevated rim (Fig. 3M) C. rhicnocephalus
— Flange with elevated rim (Fig. 3N, O) 14
14 Frontal horn not projecting beyond frontal flange (Fig. 3N) C. fatulus
– Frontal horn projects beyond frontal flange (Fig. 3O) C. mangoldi

Discussion

Cryptotermes pugnus is the second species of the genus described from Brazil and the first from the Caatinga dry forest, with records for two ecoregions, which have different geomorphological history and climatic parameters, “Planalto da Borborema” (São José dos Cordeiros, Paraíba State) and “Depressão Sertaneja Meridional” (Curaçá, Bahia State) (Silva et al. 2018). There is also a record of C. havilandi from the Caatinga dry forest (Vasconcellos unpublished data), an exotic species which probably originated in tropical West Africa (Scheffrahn et al. 2003). There are no records of C. pugnus infestations in buildings, either in urban or agricultural environments. Other kalotermitids reported from the Caatinga include two undescribed species of Glyptotermes Froggatt, 1897, Rugitermes cf. niger Oliveira, 1979, an undescribed species of Rugitermes Holmgren, 1911 (Bandeira et al. 2003), and Tauritermes bandeirai Scheffrahn & Vasconcellos, 2022 (Scheffrahn and Vasconcellos 2020).

Small colonies of C. pugnus were found on adult individuals of Cenostigma nordestinum E. Gagnon & G.P. Lewis, an endemic tree of the Caatinga dry forest, which presents hard, highly dense (>0.84 g/cm3) wood and individuals that can exceed 10 m in height (Silva et al. 2009). Due to the hardness of the wood, access to C. pugnus colonies is difficult, requiring the use of an ax and/or chainsaw. Possibly because of this, its colonies are rarely found. At the type locality, there are records of C. pugnus alate flights from late December to early February (Lucena et al. 2022).

Acknowledgements

We thank Reginaldo Constantino for image of C. contognathus (Fig. 3D).

Additional information

Conflict of interest

The authors have declared that no competing interests exist.

Ethical statement

No ethical statement was reported.

Funding

This study was supported by the University of Florida. The second coauthor thanks CNPq for the research grant (proc.309820/2020-0).

Author contributions

Scheffrahn wrote first draft. Vasconcellos did field work and inproved first draft.

Author ORCIDs

Alexandre Vasconcellos https://orcid.org/0000-0001-7211-7097

Data availability

All of the data that support the findings of this study are available in the main text.

References

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1 Imago without arolia (C. camelus imago unknown).
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