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Research Article
Parvitermes (Isoptera, Termitidae, Nasutitermitinae) in Central America: Two new termite species and reassignment of Nasutitermes mexicanus
expand article infoRudolf Scheffrahn
‡ University of Florida, Davie, United States of America
Open Access

Abstract

The termite genus Parvitermes is now recognized on the Central American mainland to include P. mexicanus, new combination (previously in Nasutitermes) and two new species, P. mesoamericanus sp. n. and P. yucatanus sp. n., herein described from soldiers and workers. These three species, nine West Indian Parvitermes, and Antillitermes subtilis all share characteristic enteric valve spines that orientate against intestinal flow. All species are subterranean nesters and cellulose feeders. Evidence is mounting that generic-level endemicity may be completely absent among the West Indian nasutitermitine fauna and that its origins stem from Central America.

Keywords

Neotropics, soldier key, enteric valve armature, new combination, taxonomy

Introduction

Emerson (1949) erected the genus Parvitermes to accommodate six small nasutiform termites: Nasutitermes brooksi Snyder, 1925 from Cuba, Constrictotermes discolor Banks, 1919 and Nasutitermes wolcotti Snyder, 1924a from Puerto Rico, Constrictotermes flaveolus Banks, 1919 and C. pallidiceps Banks, 1919 from Hispaniola, and Nasutitermes laticephalus Snyder, 1926 from Bolivia. Three additional Parvitermes species from Hispaniola were later added (P. subtilis Scheffrahn & Krecek, 1993, P. collinsae Scheffrahn & Roisin, 1995, and P. dominicanae Scheffrahn et al., 1998). The distribution of P. brooksi and P. wolcotti was expanded to include the central Bahamas (Scheffrahn et al. 2006) and the British and U.S. Virgin Islands (Scheffrahn et al. 2003), respectively.

Roisin et al. (1996) revised the small nasutes of the West Indies based mainly on worker morphology. The following taxa were reassigned to Parvitermes: Constrictotermes toussainti Banks, 1919, Nasutitermes aequalis Snyder, 1924b, and Eutermes antillarum Holmgren, 1910. Furthermore, P. discolor was placed in a new genus, Caribitermes Roisin, 1996, and P. subtilis was placed in another new genus, Antillitermes Roisin, 1996. The removal of Parvitermes as a Neotropical mainland genus was completed by Roisin et al. (1996), who transferred P. laticephalus to Velocitermes, and by Cuezzo and Cancello (2009), who showed that the Brazilian P. bacchanalis Mathews, 1977 should also be excluded from Parvitermes.

In the present paper, Parvitermes is shown to be a widespread endemic genus of the Central American mainland as Parvitermes mexicanus (Light, 1933), comb. n. and as two new Central American species, P. mesoamericanus and P. yucatanus. All three species are described mainly by the shape of soldier nasus and their enteric valve armature.

Materials and methods

All material is from the University of Florida Termite Collection (UF) at the author’s address. Photographs (Figs 1A, 35) were taken as multi-layer montages using a Leica M205C stereomicroscope controlled by Leica Application Suite version 3 software. Preserved specimens were taken from 85% ethanol and suspended in a pool of Purell® Hand Sanitizer to position the specimens over a transparent Petri dish background. Microphotographs (Figs 1B, 1C, 2, 6) were taken from slide mounts in PVA medium (BioQuip, Rancho Dominquez, CA) using a Leica CTR 5500 compound microscope with bright field lighting. The distribution map (Fig. 7) was created using ArcGIS Desktop ver. 10.3 (ESRI, Redlands, CA).

Figure 1.

Parvitermes brooksi. A Dorsal left view of gut: M = mesenteron, P1-P4 = proctodeal segments 1-4 (limits of P2 highlighted) B Whole mount of P2 with musculature removed. Posterior (end attached to P3) at top of image C P2 splayed open; bacterial pellet attached to spines of the central pad.

Figure 2.

Enteric valve armatures. Parvitermes mexicanus comb. n., A worker B soldier. Parvitermes mesoamericanus sp. n. C worker D soldier. Parvitermes yucatanus sp. n. E worker F soldier. Posterior (end attached to P3) at top of each image.

Figure 3.

Parvitermes soldier head capsule. Parvitermes mexicanus comb. n., A dorsal B lateral. Parvitermes mesoamericanus sp. n. C dorsal D lateral. Parvitermes yucatanus sp. n. E dorsal F lateral.

Figure 4.

Parvitermes worker head and thorax, lateral views. A P. mexicanus comb. n. B P. mesoamericanus sp. n. C P. yucatanus sp. n.

Figure 5.

Parvitermes worker bodies with P1 highlighted. A P. mexicanus comb. n., dorsolateral view B P. mesoamericanus sp. n., ventrolateral view C P. yucatanus sp. n., ventrolateral view.

Figure 6.

Parvitermes ventral views of left worker mandibles. A P. mexicanus comb. n. B P. mesoamericanus sp. n. C P. yucatanus sp. n.

Figure 7.

Collection localities of three Parvitermes species in the UF Termite collection. The far western sample of P. mexicanus comb. n. was taken near the type locality.

Taxonomy

Key to soldiers of Central American species of Parvitermes

1 Head capsule widest near posterior third (Fig. 3A), nasus angled slightly above plane of vertex (Fig. 3B) P. mexicanus
Head capsule widest near middle (Fig. 3C, E), nasus angled in line or below plane of vertex (Fig. 3D, F) 2
2(1) Nasus, in lateral view, nearly cylindrical in apical 2/3 (Fig. 3D) P. mesoamericanus sp. n.
Nasus, in lateral view, conical (Fig. 3F) P. yucatanus sp. n.

Parvitermes Emerson, 1949

Type species

Nasutitermes brooksi Snyder, 1925. Type: soldier; Cuba, Cienfuegos, Soledad.

Remarks

The nomenclatural summary for Parvitermes is provided by Krishna et al. (2013). The generic redescription of Parvitermes by Roisin et al. (1996) is relevant to all three Parvitermes species described herein with the exception that P. mexicanus comb. n. has a shorter first proctodeal segment (P1) compared to all others.

Diagnosis

The spine arrangement and counter-current orientation of the Parvitermes enteric valve armature (EVA), with the exception of Antillitermes, is unique among all termite genera. In addition to Parvitermes, only three other nasutitermitine genera are found from Mexico to Nicaragua, including Nasutitermes, Subulitermes, and Tenuirostritermes (Atlantitermes from Nicaragua in Scheffrahn et al. (2005) is an error). Compared to mainland Parvitermes, head capsules of Nasutitermes soldiers are larger and darker (with the exception of N. glabritergus Snyder & Emerson, 1949 from Honduras, unpubl. record), those of Subulitermes are much smaller with much narrower cylindrical nasi, and the head capsules of Tenuirostritermes are very constricted near their middle.

Workers and soldiers

The EVA arises within the second proctodeal segment (P2) which forms a swelling at the terminus of a very long (shorter and thicker in P. mexicanus), U-shaped P1. The P2 constricts somewhat at its attachment near the dorsal surface of the third proctodeal segment (P3 or paunch) to form a pear-shaped segment (Fig. 1A). The posterior EV ring (sensu Noirot 2001) of both workers and soldiers is uniquely composed of three keel-shaped pads covered with about 7-15 long spines directed into the P2 lumen (Fig. 1B). The spines are curved or angled counter to the direction of the food flow. The spiny pads are separated with or without additional patches of tiny conical teeth (Figs 1C, 2). In preserved specimens, the Parvitermes spines of each pad are imbedded into a congealed pellet of presumed bacterial cells (Fig. 1C).

Parvitermes mexicanus (Light, 1933), comb. n.

Nasutitermes mexicanus Nickle & Collins, 1988 (soldier). Type localities: MEXICO: Colima: Colima, Jala, and Madrid.

Material examined

MEXICO, 76 km S. Oaxaca, 16.49, -96.74, 11 Jan 1997, T.G. Myles & D.A. Muruvanda, UF no. MX23; Chamela, 19.5314, -105.0832, 1 Apr 1996, G. Thompson, MX99; Aguaje de la Anona, 15.7731, -96.2168 , 27 May 2006, T. Atkinson, MX572. Soldiers of these three colony samples were identified based on the following: congruence with both dorsal and lateral head capsule photographs from the original description (Figs L, O, Light 1933), Light’s 1933 rostrum (nasus) diagnosis stating that it is “slightly uplifted distally”, proximity of the Chamela sample to the type localities, and Light’s 1933 measurements. Furthermore, scanning electron micrographs of a P. mexicanus soldier from Chamela (figs 15D, H, Nickle and Collins 1988) agree perfectly with the examined soldiers.

Comparisons

See below under P. mesoamericanus sp. n.

Alate

Unknown.

Soldier

(Table 1, Fig. 2B, 3A, B). Monomorphic; however some rare divergent size morphs reported by Light 1933. Head capsule and pronotum light orange-brown; nasus darker. In dorsal view, cephalic gland duct partially or completely visible from nasus to reservoir. Many small and a few longer setae scattered on head; setae on nasus very small and numerous.

Measurements (mm) of Parvitermes mexicanus comb. n. soldiers.

Colony Head length to end of nasus Head width (max.) Pronotal width Hind tibia length
MX23 (n=3) 1.44–1.52 0.80–0.82 0.42–0.46 0.94–0.96
MX99 (n=2) 1.50–1.52 0.88–0.92 0.44–0.45 0.92–0.93
MX572 (n=8) 1.34–1.46 0.72–0.80 0.44–0.46 0.89–0.96
Range 1.34–1.52 0.72–0.92 0.42–0.46 0.89–0.96
Mean 1.43 0.80 0.45 0.92

In dorsal view, head capsule outline, without nasus, subtrapazoidal; nasus about 2/3 as long as rest of head capsule; head capsule slightly constricted behind antennal sockets; widest at posterior 1/3. In lateral view, vertex with slight concavities near midpoint; second slight concavity at base of nasus; plane of vertex parallel with ventral margin of head capsule. In dorsal view nasus is narrowly conical, about twice its width at base compared to midpoint. In lateral view nasus narrowly conical; angled ca. 5°above plane of vertex. Mandibles without points. Antennal with 13 articles (1>2<3>4). Hind tibia longer than head width. Pronotum with scattered microscopic setae (0.03 mm); anterior lobe evenly convex and ca. 90° from plane of posterior lobe, posterior lobe more blunt. Each tergite with 3-4 long (0.1 mm) setae and dozens of microscopic (0.03 mm) setae. EVA consists of three irregular rows of sharp, narrow, and down-curved spines; a few small scale-like spines in the anterior ring.

Worker

(Table 2, Figs 2A, 4A, 5A, 6A). Monomorphic. Head capsule very pale yellow with two slightly darker yellowish-orange dorso-lateral patches; pronotum very pale yellow; body, antennae, and legs hyaline. Antennal with 12 articles. Postclypeus considerably inflated in lateral view; scattered short and medium setae on head capsule. Abdomen with many short and a few scattered longer setae. Mandibles with about eight ridges on molar plate, molar plate with distinct dorsal notch; apical and first marginal teeth of similar shape and projection; third marginal smaller, separated from first by slightly concave cutting edge. Gut with P1 U-shaped turn near midpoint, bottom of turn extending only to dorso-lateral aspect of abdomen. EVA consists of three irregular rows of about 10-12 sharp, narrow, and down-curved spines; anterior ring with three patches of small scale-like spines.

Measurements (mm) of Parvitermes mexicanus comb. n. workers.

Colony Head length to end of postclypeus Postclypeal length Head width Pronotal width Hind tibia length
MX23 (n=3) 0.84–1.06 0.21–0.22 0.88–1.06 0.44–0.48 0.72–0.84
MX99 (n=1) 0.92 0.23 0.96 0.42 0.74
MX572 (n=11) 0.80–1.01 0.21–0.26 0.90–1.04 0.40–0.55 0.74–0.94
Range 0.80–1.06 0.21–0.26 0.88–1.06 0.40–0.55 0.72–0.94
Mean 0.93 0.23 0.96 0.44 0.78

Distribution

Tropical Pacific slope of Mexico (Fig. 7).

Parvitermes mesoamericanus sp. n.

Type-locality

Honduras, S. Pinalillo, 15.0860, -88.2160, 144 m elev.

Holotype

Soldier, 2 Jun 2007, Scheffrahn et al. cols., UF no. HN822 (in microvial).

Paratypes

GUATEMALA: Salama, 15.1055, -90.3261 , 28 May 2006, Scheffrahn et al., GUA16; Road to Rabinal, 15.1045, -90.3722, 28 May 2006, Scheffrahn et al., GUA33; HONDURAS: Coyolito, 13.3149, -87.6227, 31 May 2007, Scheffrahn et al., HN431; NICARAGUA: Los Cardones, 12.8851, -86.0534, 30 May 2004, Scheffrahn et al., NI114.

Imago

Unknown.

Soldier

(Table 3, Figs 2D, 3CD, 8). Monomorphic. Head capsule and pronotum light orange-brown; nasus darker. In dorsal view, cephalic gland duct partially or completely visible from nasus to reservoir. Many small, some medium, and a few longer setae scattered on head; setae on nasus small and numerous. In dorsal view, head capsule outline, without nasus, ovoid; nasus about 2/3 as long as rest of head capsule; head capsule barely constricted behind antennal sockets; widest in middle. In lateral view, vertex nearly in a flat plane; vertex and ventral margin of head capsule converge to front. In dorsal view nasus is narrowly conical, about thrice its width at base compared to midpoint; In lateral view nasus conical; projecting directly anterior below plane of vertex. Mandibles with short, very narrow, points. Antennal with 12 articles (1>2<3=4). Hind tibia about as long as or shorter than maximum head width. Pronotum with scattered microscopic setae (0.05 mm); anterior lobe evenly convex and ca. 90° from plane of posterior lobe, posterior lobe more blunt. Each tergite with 3-4 long (0.13 mm) setae and dozens of microscopic (0.05 mm) setae. EVA consists of three irregular rows of about 8-12 long subtriangulate, and very slightly down-curved spines; a few small scale-like spines in the anterior ring.

Measurements (mm) of Parvitermes mesoamericanus sp. n. soldiers.

Colony Head length to end of nasus Head width (max.) Pronotal width Hind tibia length
GUA16 (n=12) 1.38–1.50 0.76–0.84 0.36–0.41 0.66–0.78
GUA33 (n=12) 1.48–1.63 0.83–0.91 0.42–0.48 0.76–0.84
HN431 (n=12) 1.37–1.49 0.76–0.82 0.41–0.43 0.68–0.76
HN822 (n=10) 1.37–1.46 0.71–0.80 0.36–0.39 0.64–0.75
NI114 (n=12) 1.30–1.43 0.69–0.75 0.40–0.42 0.64–0.75
Range 1.30–1.63 0.69–0.91 0.36–0.48 0.64–0.84
Mean 1.44 0.78 0.40 0.72
Figure 8.

Field photograph of P. mesoamericanus sp. n. foragers feeding within a crevice of damp wood (Coyolito, Honduras; paratype locality, HN431).

Worker

(Table 4, Figs 2C, 4B, 5B, 6B, 8). Monomorphic. Head capsule very pale yellow with two slightly darker yellowish-orange dorso-lateral patches; pronotum very pale yellow; body, antennae, and legs hyaline. Antennal with 12-13 articles. Postclypeus considerably inflated in lateral view; scattered short, medium, and a few longer setae on head capsule. Abdomen with many short and longer setae. Mandibles with about eight ridges on molar plate, molar plate with slight dorsal notch; apical and first marginal teeth of similar shape and projection; third marginal smaller, separated from first by slightly emarginate cutting edge. Gut with very long P1; U-shaped turn near midpoint, bottom of turn extending to ventro-lateral aspect of abdomen. EVA consists of three irregular rows of about 8-12 long subtriangulate, and very slightly down-curved spines; three patches with small scale-like spines in the anterior ring.

Measurements (mm) of Parvitermes mesoamericanus sp. n. workers.

Colony Head length to end of postclypeus Postclypeal length Head width Pronotal width Hind tibia length
GUA16 (n=12) 0.74–0.92 0.17–0.23 0.80–0.92 0.39–0.52 0.54–0.75
GUA33 (n=12) 0.85–0.98 0.20–0.23 0.87–0.92 0.44–0.54 0.63–0.80
HN431 (n=12) 0.70–0.88 0.17–0.20 0.77–0.90 0.37–0.48 0.53–0.70
HN822 (n=10) 0.76–0.86 0.17–0.20 0.76–0.86 0.36–0.46 0.58–0.74
NI114 (n=12) 0.76–0.85 0.18–0.20 0.77–0.86 0.40–0.46 0.60–0.75
Range 0.70–0.98 0.17–0.23 0.76–0.92 0.36–0.54 0.53–0.80
Mean 0.83 0.20 0.84 0.45 0.66

Etymology and distribution

Named for Middle America which encompasses Guatemala, Honduras, and Nicaragua; the known range of this termite. The distribution habitat of P. mesoamericanus (Fig. 7) is more xeric than adjacent regions lacking this termite.

Comparisons

The soldier of P. mesoamericanus has the nasus directed forward, the head capsule widest in the middle, a few scattered long setae on the vertex, and points on the mandibular stubs while in P. mexicanus, the nasus is slightly upturned, the head is widest in the posterior third, the vertex lacks scattered long setae, and the mandibular stubs have points. The worker of P. mesoamericanus has a much longer and more ventrally positioned P1, stouter and less curved EVA spines, and longer setae on the vertex, while in P. mexicanus the P1 is shorter and more dorsal, the EVA spines are thinner and more curved, and the setae on the vertex are shorter. The P. mesoamericanus worker is proportionally smaller to its soldier as compared P. mexicanus. Both castes of P. mexicanus have longer hind tibia than P. mesoamericanus.

Parvitermes yucatanus sp. n.

Type-locality

Mexico, 0.9 km N. gate of Punta Sam, 21.2423, -86.8056, 2 m elev.

Holotype

Soldier. 9 Dec 1997, J. Chase, J. Mangold cols., UF col. no. MX161 (in microvial).

Paratypes

GUATEMALA: P. N. Tikal, 17.1371, -89.6803, 30 May 2006, Scheffrahn et al., GUA222; MEXICO: Hwy 307, 1 km S Marine, 20.5803, -87.1424, 8 Dec 1997, J. Chase, J. Mangold, MX148; same data, MX152; Chicana Ecovillage, 18.5178, -89.4846, 21 Jan 2001, MX281; 10.5 km W Coba toward Chemax, 20.5514, -87.8049, 22 Jan 2003, J. Chase, J. Mangold, MX492.

Alate

Unknown.

Soldier

(Table 5, Figs 2F, 3E, F). In all respects, similar to P. mesoamericanus except for the following: In dorsal view nasus conical, about 1.6x its width at base compared to midpoint; in lateral view nasus broadly conical. Mandibles with short, very narrow, points. Antennal with 11-12 articles (1>2<3=4). Hind tibia usually shorter than maximum head width. Pronotum with a few longer setae (0.10 mm) along margin of anterior lobe.

Measurements (mm) of Parvitermes yucatanus sp. n. soldiers.

Colony Head length to end of nasus Head width (max.) Pronotal width Hind tibia length
GUA222 (n=12) 1.36–1.45 0.70–0.78 0.36–0.40 0.65–0.70
MX148 (n=2) 1.28–1.29 0.66 0.36 0.58
MX152 (n=12) 1.34–1.42 0.68–0.74 0.34–0.40 0.64–0.70
MX161 (n=12) 1.36–1.46 0.74–0.78 0.38–0.44 0.66–0.74
MX281 (n=12) 1.38–1.45 0.72–0.78 0.34–0.39 0.64–0.76
MX492 (n=12) 1.32–1.42 0.72–0.78 0.34–0.38 0.64–0.70
Range 1.28–1.46 0.66–0.78 0.34–0.44 0.58–0.76
Mean 1.38 0.72 0.37 0.67

Worker

(Table 6, Figs 2E, 4C, 5C, 6C). In all respects, similar to P. mesoamericanus except for the following: Mandibles with about seven ridges on molar plate, molar plate without dorsal notch; apical and first marginal teeth of similar shape and projection; third marginal smaller, separated from first by slightly concave cutting edge. EVA consists of three irregular rows of about 7–12 long, narrow, subtriangulate, and slightly down-curved spines.

Measurements (mm) of Parvitermes yucatanus sp. n. workers.

Colony Head length to end of postclypeus Postclypeal length Head width Pronotal width Hind tibia length
GUA222 (n=12) 0.73–0.85 0.17–0.20 0.76–0.86 0.36–0.46 0.54–0.67
MX148 (n=4) 0.73–0.80 0.17–0.19 0.73–0.80 0.36–0.39 0.51–0.58
MX152 (n=12) 0.68–0.84 0.18–0.23 0.71–0.85 0.32–0.44 0.54–0.70
MX161 (n=12) 0.78–0.87 0.17–0.19 0.77–0.84 0.40–0.53 0.56–0.74
MX281 (n=12) 0.76–0.84 0.17–0.19 0.78–0.84 0.36–0.44 0.60–0.74
MX492 (n=12) 0.75–0.85 0.17–0.19 0.74–0.82 0.36–0.40 0.54–0.67
Range 0.68–0.87 0.17–0.23 0.71–0.86 0.32–0.53 0.51–0.74
Mean 0.78 0.18 0.79 0.38 0.61

Etymology and distribution

Named for the Yucatan Peninsula which encompasses Belize, Mexico, and Guatemala; the known range of P. yucatanus (Fig. 7). This region has a pronounced dry winter season.

Comparisons

The soldiers of P. yucatanus and P. mesoamericanus are very similar with the following exception: the nasus of P. yucatanus, in lateral view, is more conical and broader at the base than that of P. mesoamericanus. The workers of P. yucatanus and P. mesoamericanus are indistinguishable. The distributions of P. yucatanus and P. mesoamericanus appear to be allopatric (Fig. 7) with the latter species occupying a more arid zone.

Biology

The Central American Parvitermes are wood-surface feeders. They typically attack wood in contact with the ground where they encase their surroundings with dark carton material (Fig. 8) reminiscent of Amitermes and build narrow foraging galleries to above-ground feeding sites (Light 1933, Weesner 1970 for P. mexicanus). Colonies nest in the soil underneath rocks and logs where brood and larvae have been found in weak cells of thin dark carton. In the West Indies, Parvitermes are often collected in hollowed-out stems of woody herbaceous plants (P. brooksi and P. wolcotti). In the arid lands of the Dominican Republic, P. flaveolus attacks wooden fence posts, and after rains, will feed on dried grass bunches that they cover with a thin arcade of soil.

Discussion

The current study reveals that Parvitermes is no longer a genus exclusive to the West Indies (Roisin et al. 1996) but has a widespread mainland complement of three species. This leaves only the monospecific genera Antillitermes and Caribitermes as the remaining endemics of all termite genera in the West Indies (excluding the continental islands of Trinidad and Tobago). The gestalt of the Antillitermes subtilis EVA closely resembles that of Parvitermes s. str. (Roisin et al. 1996) and suggests that the EVA is a mainland synapomorphy of both genera. Antillitermes subtilis may very well be a species of Parvitermes. Caribitermes discolor may also have a mainland lineage as it resembles an undescribed species from Panama (PN1315, Scheffrahn unpubl.). Therefore, it is quite possible, with the exception of the relict Constrictotermes guantanamensis from Cuba (Krěcěk et al. 1996), that all West Indian termites share congeneric species on the Central American mainland and that the West Indian fauna arose from Pleistocene/Miocene (Krishna and Grimaldi 2009) overwater dispersal events from Central America (Darlington 1938, Hedges 1996) or the more recent late Pleistocene land connections (Scheffrahn et al. 2006).

Acknowledgements

I thank my fellow collectors on both the Guatemala and Honduras expeditions (Scheffrahn et al. in “materials examined sections above): Brian Bahder, Jim Chase, Jan Krecek, Vinda Maharajh, John Mangold, Tim Myles, Tom Nishimura, and Bob Setter. In Nicaragua, I was accompanied by JC, JK, VM, JM, Bayardo Herrera, and Jorge Moreno. I thank Liam Lynch for his able assistance with montage photography, Tony Postle for morphometrics, and Ben Gillenwaters and John Warner for review of this manuscript. Much of this research was funded by Terminix International Co. L.P.

References

  • Banks N (1919) Antillean Isoptera. Bulletin of the Museum of Comparative Zoology 62: 475–489.
  • Cuezzo C, Cancello EM (2009) A new species of Obtusitermes (Isoptera, Termitidae, Nasutitermitinae) from South America. Zootaxa 1993: 61–68.
  • Darlington PJ (1938) The origin of the fauna of the Greater Antilles, with discussion of dispersal of animals over water and through the air. The Quarterly Review of Biology 13: 274–300. doi: 10.1086/394561
  • Emerson AE (1949) Descriptions of new genera. In: Snyder TE (Ed.) Catalog of the termites of the world. Smithsonian Miscellaneous Collections 112: 374–377.
  • Hedges SB (1996) Historical biogeography of West Indian vertebrates. Annual Review of Ecology and Systematics 27: 163–196. doi: 10.1146/annurev.ecolsys.27.1.163
  • Holmgren N (1910) Versuch einer Monographie der amerikanischen Eutermes-Arten. Mitteilungen aus dem Naturhistorischen Museum (Hamburg) 27: 171–325.
  • Krěcěk J, Scheffrahn RH, Roisin Y (1996) Greater Antillean Nasutitermitinae (Isoptera: Termitidae): Constrictotermes guantanamensis, a new subterranean termite from eastern Cuba. Florida Entomologist 79: 180–187. doi: 10.2307/3495815
  • Krishna K, Grimaldi DA (2009) Diverse Rhinotermitidae and Termitidae (Isoptera) in Dominican amber. American Museum Novitates 3640: 1–48. doi: 10.1206/633.1
  • Krishna K, Grimaldi DA, Krishna V, Engel MS (2013) Treatise on the Isoptera of the world: Volume 5 Termitidae (part two). Bulletin of the American Museum of Natural History 377: 1496–1987. doi: 10.1206/377.5
  • Light SF (1933) Termites of western Mexico. University of California Publications in Entomology 6: 79–164.
  • Mathews AGA (1977) Studies on Termites from the Mato Grosso State, Brazil. Academia Brasileira de Ciencias, Rio de Janeiro, 267 pp.
  • Nickle DA, Collins MS (1988) The termite fauna (Isoptera) in the vicinity of Chamela, State of Jalisco, Mexico. Folia Entomologica Mexicana 77: 85–122.
  • Noirot C (2001) The gut of termites (Isoptera). Comparative anatomy, systematics, phylogeny. II. Higher termites (Termitidae). Annales de la Societe Entomologique de France 37: 431–471.
  • Roisin Y, Scheffrahn RH, Křeček J (1996) Generic revision of the smaller nasute termites of the Greater Antilles (Isoptera, Termitidae, Nasutitermitinae). Annals of the Entomological Society of America 89: 775–787. doi: 10.1093/aesa/89.6.775
  • Scheffrahn RH, Křeček J (1993) Parvitermes subtilis, a new subterranean termite (Isoptera: Termitidae) from Cuba and the Dominican Republic. Florida Entomologist 76: 603–607. doi: 10.2307/3495793
  • Scheffrahn RH, Jones SC, Křeček J, Chase JA, Mangold JR, Su N-Y (2003) Taxonomy, distribution, and notes on the termites (Isoptera: Kalotermitidae, Rhinotermitidae, Termitidae) of Puerto Rico and the U.S. Virgin Islands. Annals of the Entomological Society of America 96: 181–201. doi: 10.1603/0013-8746(2003)096[0181:TDANOT]2.0.CO;2
  • Scheffrahn RH, Krecek J, Maharajh B, Chase JA, Mangold JR, Moreno J, Bayardo H (2005) Survey of the termites (Isoptera: Kalotermitidae, Rhinotermitidae, Termitidae) of Nicaragua. Florida Entomologist 88: 549–552. doi: 10.1653/0015-4040(2005)88[549:SOTTIK]2.0.CO;2
  • Scheffrahn RH, Křeček J, Chase JA, Maharajh B, Mangold JR (2006) Taxonomy, biogeography, and notes on termites (Isoptera: Kalotermitidae, Rhinotermitidae, Termitidae) of the Bahamas and Turks and Caicos Islands. Annals of the Entomological Society of America 99: 463–486. doi: 10.1603/0013-8746(2006)99[463:TBANOT]2.0.CO;2
  • Scheffrahn RH, Roisin Y (1995) Antillean Nasutitermitinae (Isoptera: Termitidae): Parvitermes collinsae, a new subterranean termite from Hispaniola and redescription of P. pallidiceps and P. wolcotti. Florida Entomologist 78: 585–600. doi: 10.2307/3496044
  • Scheffrahn RH, Roisin Y, Su N-Y (1998) Greater Antillean Nasutitermitinae (Isoptera: Termitidae): Parvitermes dominicanae, a new subterranean termite from Hispaniola. Florida Entomologist 81: 179–187. doi: 10.2307/3496084
  • Snyder TE (1924b) A new subgenus of Nasutitermes Banks (Isop.). Proceedings of the Entomological Society of Washington 26: 20–24.
  • Snyder TE (1924a) Description of a new termite from Porto Rico. Proceedings of the Entomological Society of Washington 26: 131–132.
  • Snyder TE (1925) A new Cuban termite. Proceedings of the Entomological Society of Washington 27: 105–106.
  • Snyder TE (1926) Termites collected on the Mulford Biological Exploration to the Amazon Basin, 1921–1922. Proceedings of the United States National Museum 68: 1–76.
  • Weesner FM (1970) Termites of the Nearctic region. In: Krishna K, Weesner FM (Eds) Biology of termites. Vol. 2. Academic Press, New York, 477–525.