2urn:lsid:arphahub.com:pub:45048D35-BB1D-5CE8-9668-537E44BD4C7Eurn:lsid:zoobank.org:pub:91BD42D4-90F1-4B45-9350-EEF175B1727AZooKeysZK1313-29891313-2970Pensoft Publishers10.3897/zookeys.617.1004010040Research ArticleAnimaliaArthropodaHexapodaInsectaInvertebrataIsopteraTermitidaeFaunistics & DistributionIdentification keySystematicsTaxonomyCenozoicNeogeneAmericasParvitermes (Isoptera, Termitidae, Nasutitermitinae) in Central America: Two new termite species and reassignment of NasutitermesmexicanusScheffrahnRudolf H.1Fort Lauderdale Research and Education Center, University of Florida, Institute of Food and Agricultural Sciences, 3205 College Avenue, Davie, Florida, 33314, USAUniversity of FloridaDavieUnited States of America
Corresponding author: Rudolf H. Scheffrahn (rhsc@ufl.edu)
Academic editor: E. Cancello
20161509201661747630F66E60F-A663-1826-9A7E-FFF7736B1B4ECCDBFC5F-FBFB-41C9-967E-C12024280F241541042907201609092016Rudolf ScheffrahnThis 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.http://zoobank.org/CCDBFC5F-FBFB-41C9-967E-C12024280F24
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.mesoamericanussp. n. and P.yucatanussp. n., herein described from soldiers and workers. These three species, nine West Indian Parvitermes, and Antillitermessubtilis 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.
Neotropicssoldier keyenteric valve armaturenew combinationtaxonomyUniversity of Florida100007698http://doi.org/10.13039/100007698Citation
Scheffrahn RH (2016) Parvitermes (Isoptera, Termitidae, Nasutitermitinae) in Central America: Two new termite species and reassignment of Nasutitermes mexicanus. ZooKeys 617: 47–63. doi: 10.3897/zookeys.617.10040
Introduction
Emerson (1949) erected the genus Parvitermes to accommodate six small nasutiform termites: Nasutitermesbrooksi Snyder, 1925 from Cuba, Constrictotermesdiscolor Banks, 1919 and Nasutitermeswolcotti Snyder, 1924a from Puerto Rico, Constrictotermesflaveolus Banks, 1919 and C.pallidiceps Banks, 1919 from Hispaniola, and Nasutitermeslaticephalus 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.dominicanaeScheffrahn 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: Constrictotermestoussainti Banks, 1919, Nasutitermesaequalis Snyder, 1924b, and Eutermesantillarum 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 Parvitermesmexicanus (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, 3–5) 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).
Parvitermesbrooksi. 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 CP2 splayed open; bacterial pellet attached to spines of the central pad.
Enteric valve armatures. Parvitermesmexicanus comb. n., A worker B soldier. Parvitermesmesoamericanus sp. n. C worker D soldier. Parvitermesyucatanus sp. n. E worker F soldier. Posterior (end attached to P3) at top of each image.
Parvitermes soldier head capsule. Parvitermesmexicanus comb. n., A dorsal B lateral. Parvitermesmesoamericanus sp. n. C dorsal D lateral. Parvitermesyucatanus sp. n. E dorsal F lateral.
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.
https://binary.pensoft.net/fig/103161TaxonomyKey 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)
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 (sensuNoirot 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).
AnimaliaIsopteraTermitidae8B2A47B1-404E-5EC7-9138-248F8FAF158DParvitermesmexicanus(Light, 1933)comb. n.NasutitermesmexicanusNickle & 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 Parvitermesmexicanus 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 Parvitermesmexicanus comb. n. workers.
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 Parvitermesmesoamericanus sp. n. soldiers.
Field photograph of P.mesoamericanus sp. n. foragers feeding within a crevice of damp wood (Coyolito, Honduras; paratype locality, HN431).
https://binary.pensoft.net/fig/103162Worker
(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 Parvitermesmesoamericanus 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.
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 Parvitermesyucatanus 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 Parvitermesyucatanus 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 AntillitermessubtilisEVA closely resembles that of Parvitermes s. str. (Roisin et al. 1996) and suggests that the EVA is a mainland synapomorphy of both genera. Antillitermessubtilis may very well be a species of Parvitermes. Caribitermesdiscolor 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 Constrictotermesguantanamensis 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.
ReferencesBanksN (1919) Antillean Isoptera.62: 475–489.CuezzoCCancelloEM (2009) A new species of Obtusitermes (Isoptera, Termitidae, Nasutitermitinae) from South America.1993: 61–68.DarlingtonPJ (1938) The origin of the fauna of the Greater Antilles, with discussion of dispersal of animals over water and through the air.13: 274–300. doi: 10.1086/394561EmersonAE (1949) Descriptions of new genera. In: SnyderTE (Ed.) Catalog of the termites of the world.112: 374–377.HedgesSB (1996) Historical biogeography of West Indian vertebrates.27: 163–196. doi: 10.1146/annurev.ecolsys.27.1.163HolmgrenN (1910) Versuch einer Monographie der amerikanischen Eutermes-Arten.27: 171–325.KrěcěkJScheffrahnRHRoisinY (1996) Greater Antillean Nasutitermitinae (Isoptera: Termitidae): Constrictotermesguantanamensis, a new subterranean termite from eastern Cuba.79: 180–187. doi: 10.2307/3495815KrishnaKGrimaldiDA (2009) Diverse Rhinotermitidae and Termitidae (Isoptera) in Dominican amber.3640: 1–48. doi: 10.1206/633.1KrishnaKGrimaldiDAKrishnaVEngelMS (2013) Treatise on the Isoptera of the world: Volume 5 Termitidae (part two).377: 1496–1987. doi: 10.1206/377.5LightSF (1933) Termites of western Mexico.6: 79–164.MathewsAGA (1977) . Academia Brasileira de Ciencias, Rio de Janeiro, 267 pp.NickleDACollinsMS (1988) The termite fauna (Isoptera) in the vicinity of Chamela, State of Jalisco, Mexico.77: 85–122.NoirotC (2001) The gut of termites (Isoptera). Comparative anatomy, systematics, phylogeny. II. Higher termites (Termitidae).37: 431–471.RoisinYScheffrahnRHKřečekJ (1996) Generic revision of the smaller nasute termites of the Greater Antilles (Isoptera, Termitidae, Nasutitermitinae).89: 775–787. doi: 10.1093/aesa/89.6.775ScheffrahnRHKřečekJ (1993) Parvitermessubtilis, a new subterranean termite (Isoptera: Termitidae) from Cuba and the Dominican Republic.76: 603–607. doi: 10.2307/3495793ScheffrahnRHJonesSCKřečekJChaseJAMangoldJRSuN-Y (2003) Taxonomy, distribution, and notes on the termites (Isoptera: Kalotermitidae, Rhinotermitidae, Termitidae) of Puerto Rico and the U.S. Virgin Islands.96: 181–201. doi: 10.1603/0013-8746(2003)096[0181:TDANOT]2.0.CO;2ScheffrahnRHKrecekJMaharajhBChaseJAMangoldJRMorenoJBayardoH (2005) Survey of the termites (Isoptera: Kalotermitidae, Rhinotermitidae, Termitidae) of Nicaragua.88: 549–552. doi: 10.1653/0015-4040(2005)88[549:SOTTIK]2.0.CO;2ScheffrahnRHKřečekJChaseJAMaharajhBMangoldJR (2006) Taxonomy, biogeography, and notes on termites (Isoptera: Kalotermitidae, Rhinotermitidae, Termitidae) of the Bahamas and Turks and Caicos Islands.99: 463–486. doi: 10.1603/0013-8746(2006)99[463:TBANOT]2.0.CO;2ScheffrahnRHRoisinY (1995) Antillean Nasutitermitinae (Isoptera: Termitidae): Parvitermescollinsae, a new subterranean termite from Hispaniola and redescription of P.pallidiceps and P.wolcotti.78: 585–600. doi: 10.2307/3496044ScheffrahnRHRoisinYSuN-Y (1998) Greater Antillean Nasutitermitinae (Isoptera: Termitidae): Parvitermesdominicanae, a new subterranean termite from Hispaniola.81: 179–187. doi: 10.2307/3496084SnyderTE (1924b) A new subgenus of Nasutitermes Banks (Isop.).26: 20–24.SnyderTE (1924a) Description of a new termite from Porto Rico.26: 131–132.SnyderTE (1925) A new Cuban termite.27: 105–106.SnyderTE (1926) Termites collected on the Mulford Biological Exploration to the Amazon Basin, 1921–1922.68: 1–76.WeesnerFM (1970) Termites of the Nearctic region. In: KrishnaKWeesnerFM (Eds) . Vol. 2. Academic Press, New York, 477–525.