Taxonomy, Distribution, and Notes on the Termites (Isoptera: Kalotermitidae, Rhinotermitidae, Termitidae) of Puerto Rico and the U.S.

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1 SYSTEMATICS Taxonomy, Distribution, and Notes on the Termites (Isoptera: Kalotermitidae, Rhinotermitidae, Termitidae) of Puerto Rico and the U.S. Virgin Islands RUDOLF H. SCHEFFRAHN, 1 SUSAN C. JONES, 2 JAN KRECEK, 1 JAMES A. CHASE, 3 JOHN R. MANGOLD, 4 AND NAN-YAO SU 1 Ann. Entomol. Soc. Am. 96(3): 181Ð201 (2003) ABSTRACT Termite surveys from Buck, Culebra, Mona, Puerto Rico, St. Croix, St. John, St. Thomas, and Vieques islands yielded 1,564 colony samples from 274 sites. Twenty-one species were recorded including: Cryptotermes brevis, Cryptotermes havilandi, Cryptotermes rotundiceps, Cryptotermes undulans, Glyptotermes liberatus, Glyptotermes pubescens, Incisitermes bequaerti, Incisitermes furvus, Incisitermes incisus, Neotermes intracaulis n. sp., Neotermes mona, and Procryptotermes corniceps (Kalotermitidae); Coptotermes havilandi, Heterotermes sp., and Prorhinotermes simplex (Rhinotermitidae); and Anoplotermes n. sp. undescribed, Caribitermes discolor, Nasutitermes acajutlae, Nasutitermes costalis, Parvitermes wolcotti, and Termes hispaniolae (Termitidae). Of these, twelve are West Indian endemics, six have endemic ranges that include the tropical American mainland, and three are nonendemic pests. Distribution maps and keys based on the soldier caste and the winged imago are provided. Neotermes intracaulis, a new species from St. Croix, is described from the imago and soldier. Additional descriptions or redescriptions are given for G. liberatus (imago), I. incisus (imago and soldier), Ca. discolor (imago and soldier), and Pa. wolcotti (imago). An understanding of West Indian termite biogeography relies on current faunal distributions and the few fossil termites available from Hispaniola. Over-water dispersal of termites on ßotsam is the most plausible mechanism for contemporary distributions, however, vicariate speciation cannot be discounted for species that are poor dispersalists. Faunal composition of Puerto Rican and the Virgin Islands termites suggest a biogeographical origin that is derived from both western and southern mainland sources. KEY WORDS survey, identiþcation key, Caribbean, new species, biogeography PUERTO RICO, ITS OUTLYING islands, and the United States Virgin Islands are located near the eastern end of the Greater Antilles island chain. With the exception of tiny Navassa Island west of Hispaniola, Puerto Rico and the U.S. Virgin Islands constitute the only United States dependencies in the Caribbean region. Recently, there has been a renewed interest in documenting the rich diversity of termites in the West Indies (Collins et al. 1997, Darlington 1992, Genet et al. 2000, Jones et al. 1995, Jones and Nalepa 2002, Scheffrahn et al. 1990), but to date, detailed deliberate surveys have been inclusive of only relatively small land areas. 1 Fort Lauderdale Research and Education Center, University of Florida, Institute of Food and Agricultural Sciences, 3205 College Ave., Fort Lauderdale, FL Ohio State University, 102 Extension Entomology Building, 1991 Kenny Road, Columbus, OH Terminix International, 1235B Eagles Landing Pkwy., Stockbridge, GA Terminix International, 9390 North Florida Ave., Suite B, Tampa, FL The Þrst taxonomic summary of West Indian termites included nine species from Puerto Rico and the U.S. Virgin Islands (Banks 1919). Snyder (1956) compiled a list of 20 species from Culebra, Mona, Puerto Rico, St. Croix, St. Thomas, and Vieques based on reports by Burmeister (1839), Banks (1919), Martorell (1945), Ramos (1946), Wolcott (1948), unpublished records, and museum specimens. Scheffrahn et al. (1994) partially revised SnyderÕs (1956) list and added some island records based on newer unpublished and published records. Many of the species reported from the region have been plagued by descriptions that lack detail, lack imago caste descriptions, or have erroneous nomenclature. Many type specimens are lost or damaged. Recent taxonomic revisions (Krecek et al. 2000; Roisin et al. 1996; Scheffrahn and Krecek 1999, 2001; Scheffrahn and Roisin 1995; Scheffrahn et al. 1994, 2000b; Thorne et al. 1994) have helped deþne some termites of the region. To unify these works with more recent Þndings and to add substantially toward an understanding of termites of the entire West Indies, we /03/0181Ð0201$04.00/ Entomological Society of America

2 182 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 96, no. 3 surveyed termites on eight islands of Puerto Rico and the U.S. Virgin Islands. Based on this survey, we provide keys for the identiþcation of soldiers and imagos, describe one new species, redescribe four species, and discuss the distribution, biology, and biogeography of these termites. Materials and Methods The current survey is based on examination of 1,564 colony samples, including six samples from Buck Island, 63 from Culebra, 48 from Mona (including those reported by Jones et al. 1995), 739 from Puerto Rico, 318 from St. Croix, 92 from St. John, 74 from St. Thomas, 213 from Vieques, and 11 from miscellaneous locations within this archipelago. Samples from 274 sites were collected between June 1984 and February Collection sites were selected based on geography, habitat type, and roadside accessibility. Termite microhabitats within walking distance at each site were exposed by hatchet, shovel, or hand including standing dead (dry, damp, decayed) and living wood, nests, foraging structures, and the under-surfaces of objects on soil, especially wood, rocks, and aged livestock dung. Records of endemic species in Hispaniola and Guadeloupe (see Table 10) are from our unpublished data. Morphometric data from 85% alcohol-preserved specimens were obtained using a stereomicroscope Þtted with an ocular micrometer. Measurements were partly adopted from Roonwal (1970). The color scheme of Sands (1965) was used with relevant modiþcations. The terms small and large soldiers (Krishna 1961) are equal to the terms short-headed and long-headed soldiers (Banks and Snyder 1920), respectively. These terms are used to separate the two common size morphs of soldiers occurring within some kalotermitid species. Structures useful in the identiþcation of Cryptotermes spp. soldiers include two pairs of protuberances: one dorsal pair in front of the antennal fossae, and one ventral pair projecting forward from the genae. These are called the frontal and genal horns, respectively (Scheffrahn and Krecek 1999). Formulae are given for relative lengths of antennal articles 2Ð5. For example, the formula indicates that the second article is longer than the third and the third through Þfth are subequal in length. Scanning electron micrographs were taken with a Hitachi S530 instrument at 20 kv of specimens that were dehydrated in absolute ethanol and 1,1,1,3,3,3- hexamethyldisilazane (Nation 1983) and then sputtercoated with platinum or gold. Scanning electron micrographs were digitized at 600 dpi, the resultant image outline traced using photograph-enhancing software (Adobe Photoshop Elements, Adobe Systems, San Jose, CA), the background converted to black, and the scale bar redrawn. Collection localities were mapped from geographic coordinates using Arc- View GIS version 3.0a software [Environmental Systems Research Institute (ESRI), Redlands, CA]. Coordinates were obtained from either GPS receiver readings at collection sites or by transposing sites from local hardcopy maps to Digital Map of the World version 1.0 (ESRI). The following names of collectors are abbreviated in the text as follows: Paul Ban (PB), James A. Chase (JC), Jan Krecek (JK), Boudanath Maharajh (BM), John R. Mangold (JM), Julian de la Rosa (JR), and Rudolf H. Scheffrahn (RS). Type and voucher material will be deposited at the Fort Lauderdale R.E.C. termite collection and at American Museum of Natural History, New York; National Museum of Natural History (Smithsonian Institution), Washington, D.C.; and the Florida State Collection of Arthropods, Florida Department of Agriculture and Consumer Services, Division of Plant Industry, Gainesville, FL. Keys to the Isoptera of Puerto Rico and U.S. Virgin Islands Based on the Soldier Caste and the Winged Imago Soldier Caste 1. Soldierless; worker fore tibiae inßated and greater in median circumference than middle or hind tibiae; postclypeus greatly inßated; soil-dwelling but do not build foraging tubes above ground (Fig. 1)... Anoplotermes n. sp. Soldiers present in colonies; workers with all tibiae equal in circumference; postclypeus not greatly inßated; nonsubterranean or soildwelling, if latter, often constructing foraging tubes above ground Nasutiform, i.e., head distinctly modiþed by long frontal projection with terminal exocrine ori- Þce; mandibles reduced to nonfunctional stubs (Figs. 2Ð5)... 3 Head not distinctly modiþed by long frontal projection with terminal glandular oriþce; mandibles well developed, functional (Figs. 6Ð22) Head capsule dark brown; nasus distinctly conical (Figs. 2 and 3); on or above-ground carton nests... 4 Head capsule pale brown; nasus slightly conical or almost cylindrical; nest in soil (Figs. 4 and 5) Head covered with dozens of erect setae (Fig. 2)... Nasutitermes acajutlae Head with six evenly-spaced setae (Fig. 3)..... Nasutitermes costalis 5. Head noticeably bilobed near middle; anterior lobe slightly narrower than posterior lobe; nasus much darker apically than at base, antennae 13Ð14-segmented (Fig. 4)..... Parvitermes wolcotti

3 May 2003 SCHEFFRAHN ET AL.: TERMITES OF PUERTO RICO 183 Fig Scanning electron micrographs of termites from Puerto Rico and the U.S. Virgin Islands. Lateral view of Anoplotermes n. sp. with arrows indicating inßated postclypeus and fore tibia (1); dorsal views of soldier head capsules of Nasutitermes acajutlae (2), Na. costalis (3), Parvitermes wolcotti (4), Caribitermes discolor (5), Cryptotermes rotundiceps with arrows indicating frontal and genal horns (6), Cr. havilandi with arrows indicating frontal and genal horns (7), Cr. brevis (8), Cr. undulans (9), Neotermes intracaulis (10), Ne. mona (11), and Glyptotermes pubescens (12). Antennae partially or completely removed for clarity. Scale bars equal 1 mm. Head faintly constricted anteriorly, not bilobed; head anterior much narrower than posterior; nasus almost concolorous throughout; antennae 12-segmented (Fig. 5) Caribitermes discolor 6. Left mandible with marginal teeth in distal half (Figs. 7Ð18); nonsubterranean... 7 Left mandible without marginal teeth in distal half (Figs. 19Ð22); subterranean or aboveground nests Head blackish, strongly phragmotic, truncate, cuboidal; mandibles not projecting much beyond labrum (Figs. 6Ð9)... 8

4 184 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 96, no. 3 Fig Scanning electron micrographs of termite soldiers from Puerto Rico and the U.S. Virgin Islands. Dorsal view of soldier head capsule of Procryptotermes corniceps (13); lateral views of soldier head capsules and G. liberatus (14) and Incisitermes bequaerti (15); and dorsal views of soldier head capsules of I. furvus (16A, right mandible 16B), I. incisus minor soldier (17) and major soldier (18), Termes hispaniolae (19), Heterotermes sp. with arrow indicating fontanelle (20), Coptotermes havilandi with arrow indicating fontanelle (21), and Prorhinotermes simplex with arrow indicating fontanelle (22). Antennae partially or completely removed for clarity. Scale bars equal 1 mm with the exception of Fig. 16B, which equals 0.5 mm.

5 May 2003 SCHEFFRAHN ET AL.: TERMITES OF PUERTO RICO 185 Head ferruginous orange, rectangular; mandibles projecting far beyond labrum (Figs. 10Ð 18) Rugosity of head capsule faint or absent (Figs. 6 and7)... 9 Rugosity of head capsule conspicuous (Figs. 8 and9) Rugosity of head capsule faint; frontal horns much larger than genal horns (Fig. 6) Cryptotermes rotundiceps Head capsule smooth; frontal horns much smaller than genal horns (Fig. 7) Cryptotermes havilandi 10. Rugosity of head capsule robust (Fig. 8); length of left apical tooth of mandible equal to basal width; maximum head width 1.19 mm (in structural wood only).. Cryptotermes brevis Rugosity of head capsule undulating (Fig. 9), wrinkled; left apical tooth of mandible attenuate, 2 longer than basal width; maximum head width 1.16 mm Cryptotermes undulans 11. Soldier large, maximum head width 2.0 mm; mandible base with dense lateral pilosity (Figs. 10 and 11) Soldier smaller, maximum head width 1.9 mm; mandible base glabrous or with few short setae near dorsal mandibular condyle (Figs. 12Ð18) Left mandible hooked, mandible tip curves 90 ; anterior and posterior margin of pronotum subparallel; eye spots unpigmented; setae sparse on head capsule (Fig. 10)...Neotermes intracaulis Left mandible not hooked, mandible tip curves 90 ; anterior margin of pronotum deeply concave, posterior margin subrectate; eye spots dark; setae dense on head capsule (Fig. 11)... Neotermes mona 13. Head capsule with frontal cleft, ßange, or ridge (Figs. 12 and 13) Head capsule without frontal modiþcations (Figs. 14Ð18) Head capsule with frons distinctly bilobed (Fig. 12); frontal ridge absent; frons slopes from vertex 45 ; genae without horns..... Glyptotermes pubescens Head capsule with frons not bilobed; frontal ridge distinct (Fig. 13); frons slopes from vertex 45 ; genal horns acute Procryptotermes corniceps 15. Third antennal articles elongated and enlarged (e.g., Fig. 16A) Third antennal articles similar to second and fourth in length and width (Fig. 14) Glyptotermes liberatus 16. Head capsule strongly compressed dorsoventrally (Fig. 15)... Incisitermes bequaerti Head capsule not so dorsoventrally ßattened, more elevated Mandibles with almost all edges Þnely, distinctly, and somewhat irregularly serrate; femora only moderately swollen, even in large soldiers; mandible bases hairless; smaller species, maximum head width 1.3 mm (Fig. 16A and B)... Incisitermes furvus Edges of mandibles usually even; when serrated, then limited to right apical tooth; femora, particularly of hind legs, and particularly in large soldiers very swollen; few short setae near dorsal mandibular condyle, particularly in large soldiers; larger species, maximum head width 1.3 mm (Figs. 17 and18)...incisitermes incisus 18. Head capsule with horn-like frontal projection; mandibles stick-like and weakly asymmetrical (Fig. 19); exposed black carton nests in tree crotches... Termes hispaniolae Head capsule without frontal projection; mandibles sickle-shaped and symmetrical (Figs. 20Ð22); nests subterranean or hidden in wood on ground Head capsule subrectangular; fontanelle faint or absent, opening dorsally; mandibles straight except for 30 curvature in distal one-fourth (Fig. 20)... Heterotermes sp. Head capsule pyriform; fontanelle distinct, opening to anterior or dorsum; mandibles curved 70 in distal one-fourth Fontanelle large, opens to anterior above labrum (Fig. 21)... Coptotermes havilandi Fontanelle minute, opens to dorsum on vertex (Fig. 22)... Prorhinotermes simplex Winged Imagos 1. Three or four pigmented and sclerotized veins in costal margin of fore wing (Figs. 23Ð25) Two pigmented and sclerotized veins in costal margin of wing (Figs. 26 and 27) Median vein of anterior wing recurved toward and intersecting radial sector at about half of wing length (Fig. 23)... 3 Median vein of anterior wing extending to near wing apex, not intersecting radial sector (Figs. 24and25) Arolia between tarsal claws absent......cryptotermes brevis Arolia between tarsal claws present Smaller species; head length with labrum 1.14 mm, pronotum width 0.82 mm, and hind tibia length 0.69 mm... Cryptotermes undulans Larger species; head length with labrum 1.18 mm, pronotum width 0.88 mm, and hind tibia length 0.75 mm With dark band between compound eyes, and with distinct V -shaped mark on frons Procryptotermes corniceps Without dark band between compound eyes, and with faint V -shaped mark on frons.. 6

6 186 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 96, no. 3 Fig Scanning electron micrographs of termite imagos from Puerto Rico and the U.S. Virgin Islands. Dorsal views of fore wings of Cryptotermes undulans (23), Incisitermes incisus (24), Glyptotermes pubescens (25), Prorhinotermes simplex (26), and Caribitermes discolor (27). Arrows indicate positions of median veins. Scale bars equal 1 mm.

7 May 2003 SCHEFFRAHN ET AL.: TERMITES OF PUERTO RICO 187 Fig Scanning electron micrographs of termite imagos from Puerto Rico and the U.S. Virgin Islands. Lateral view of head and pronotum of Neotermes intracaulis (28); dorsal view of head, prothorax, and wing scales of dealated imagos of Heterotermes sp. (29), Caribitermes discolor (30), and Parvitermes wolcotti (31). Arrows indicate positions of fore and hind wing scales. Scale bars equal 1 mm. 6. Head brownish; smaller species; maximum head width 1.00 mm and fore wing length from suture 7.81 mm... Cryptotermes havilandi Head yellowish; larger species; maximum head width 1.01 mm and fore wing length from suture 8.38 mm.. Cryptotermes rotundiceps 7. Media running very close and parallel to unbranched radial sector (Fig. 25)... 8 Media running parallel, but not very close to branched radial sector (Fig. 24) Head capsule and pronotum reddish-brown; wing membrane and nodulations hyaline, ocelli maximum diameter 0.13 mm, and ocelli separated from ocular sclerite by distance shorter than ocular sclerite width; surface of head and pronotum smooth Glyptotermes pubescens Head capsule and pronotum blackish; wing membrane semiopaque with pigmented tips of nodulations, ocelli maximum diameter 0.09 mm, and ocelli separated from ocular sclerite by distance larger than ocular sclerite width; surface of head and pronotum densely textured by Þne granulation Glyptotermes liberatus 9. Media pigmented and sclerotized; large species, head width at eyes 1.4mm...10 Media unpigmented and unsclerotized, frons convex and smooth; smaller species, head width at eyes 1.4mm Frons ßattened and rather rugose; ocelli rather indistinct, subhyaline; pronotum almost concolorous with head; pronotum deeply concave...neotermes mona Frons smooth; ocelli very distinctly pigmented, white; pronotum distinctly paler than head; pronotum shallowly concave (Fig. 28) Neotermes intracaulis 11. General dorsal pigmentation dark castaneous; pterothorax chevron pattern absent; wing membranes blackish with noticeable nodulations Incisitermes furvus General dorsal pigmentation yellow-orange or brown; pterothorax chevron pattern distinct;

8 188 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 96, no. 3 Fig. 32. Localities and distribution of Cryptotermes brevis, Cr. rotundiceps, Cr. undulans, Incisitermes bequaerti, and I. furvus in Puerto Rico and the U.S. Virgin Islands. wing membrane subhyaline with very faint nodulations Smaller species with brown dorsum; head width at eyes and pronotum width 1.0 mm; head distinctly darker anteriorly than posteriorly; vertex and occiput with two longitudinal stripes on each side.. Incisitermes bequaerti Larger species with yellow-orange dorsum; head width at eyes 1.3 mm, and pronotum width 1.4 mm; head nearly concolorous; vertex and occiput without lateral stripes.... Incisitermes incisus 13. Scales of anterior wing noticeably larger than, and overlapping, posterior scales (Fig. 29).. 14 Scales on anterior and posterior wings subequal; anterior scale not overlapping posterior scale (Figs. 30 and 31) Forewing membrane smooth; costal margin not convex in middle; median vein distinct, especially near suture Forewing membrane reticulate; costal margin convex in middle; median vein absent Prorhinotermes simplex 15. Wing membrane with few hairs; smaller species, head width at eyes 0.9 mm, pronotum width 0.8 mm, and maximum diameter of compound eyes 0.2mm... Heterotermes sp. Wing membrane with many hairs; larger species, head width at eyes 1.4 mm, pronotum width 1.3 mm, and maximum diameter of compound eyes 0.4mm Coptotermes havilandi 16. Third antennal article much smaller than fourth...17 Third and fourth antennal articles subequal Head and pronotum pale brownish, wings very pale orange brown; postclypeus distinctly inßated, its length two-thirds of its width; fontanelle almost invisible Anoplotermes n. sp. Head, pronotum, and wings dark sepia brown; postclypeus only slightly convex, its length one-half of its width; fontanelle distinct.... Termes hispaniolae 18. General coloration dark brown to blackish; head concolorous or only slightly darker than pronotum; ocelli faintly elevated General coloration pale dirty or orange brown; head noticeably darker than pronotum; ocelli distinctly protruding Larger species; head length with labrum 1.5 mm; head width at eyes 1.5 mm; postclypeus much paler than head capsule Nasutitermes costalis Smaller species; head length with labrum 1.26 mm; head width at eyes 1.05 mm; postclypeus slightly paler than head capsule (Fig. 30)... Caribitermes discolor 20. Head width at eyes 1.5 mm; postclypeus subcircular; distances between ocelli and antennal fossae larger than maximum ocellar diameter...nasutitermes acajutlae Head width at eyes 1.2 mm; postclypeus subtruncate; distances between ocelli and antennal fossae smaller than maximum ocellar diameter (Fig. 31)... Parvitermes wolcotti Kalotermitidae Cryptotermes brevis (Walker) (Fig. 8) Remarks. Cryptotermes brevis is the most destructive drywood termite species in the region and worldwide. Dispersal ßights of Cr. brevis are nocturnal and peak in late spring. Distribution. Although Cr. brevis was described from Jamaica (Walker 1853), its endemic origin is not know and it has never been collected from nonstructural wood in the West Indies. This pest is likely established on every inhabited island in the West Indies. Our records for the region (Fig. 32) are limited because of this speciesõ strict association with moisture-protected structural lumber. Cryptotermes havilandi (Sjöstedt) (Fig. 7) Remarks. This nonendemic Cryptotermes is usually sympatric with other regional kalotermitids infesting solid dead wood in wild and domestic habitats including dead branches on living trees and shrubs as well as

9 May 2003 SCHEFFRAHN ET AL.: TERMITES OF PUERTO RICO 189 Fig. 33. Localities and distribution of Cryptotermes havilandi, Glyptotermes liberatus, G. pubescens, Neotermes intracaulis, and Ne. mona in Puerto Rico and the U.S. Virgin Islands. structural lumber. Cryptotermes havilandi apparently has a greater moisture requirement than Cr. brevis limiting the formerõs ability to colonize wood under xeric conditions. Distribution. Cryptotermes havilandi is apparently native to tropical Africa (Gay 1967) and has been introduced to the Lesser Antilles. The collections of Cr. havilandi on St. Croix and St. Thomas (Fig. 33) are the westernmost on record for the region. Cryptotermes rotundiceps Scheffrahn and Krecek (Fig. 6) Remarks. Cryptotermes rotundiceps is a recently described species (Scheffrahn and Krecek 1999) that is an inhabitant of rather xeric localities. This species usually infests dry solid hardwoods. In Puerto Rico, it is known from two coastal localities in the southeast. Distribution. Cryptotermes rotundiceps is conþned to southern Hispaniola and southeastern coastal Puerto Rico (Fig. 32). Cryptotermes undulans Scheffrahn and Krecek (Figs. 9 and 23) Remarks. Cryptotermes undulans is an uncommon inhabitant of coastal and inland forests where it infests dead limbs and tree trunks. Wolcott (1948) reports the occurrence of this species (referred to as Cr. cavifrons, see Table 11) in structures. Distribution. This species was recently described from Puerto Rico (Scheffrahn and Krecek 1999). New records include Vieques and St. Croix (Fig. 32). Glyptotermes liberatus (Snyder) (Fig. 14) Kalotermes posticus (Hagen); Banks 1919: 478 [soldier] Kalotermes (Kalotermes) liberatus Snyder 1929: 81 [soldier] Glyptotermes liberatus; Snyder 1949: 49 [catal.] Imago. (Table 1). Previously undescribed. In dorsal view, head, pronotum, antennae, and mandible dentition dark chestnut brown; wing scales, sclerotized veins, and thoracic and abdominal tergites chestnut brown; anterior frons, labrum, and mandibles except for dentition, ferruginous. Epicranial suture reduced to short posterior line. V-shaped mark on frons faint; T-shaped mark on pronotum very distinct. Head capsule cuticle Þnely granulate causing dull appearance. Two intermediate abdominal sternites much paler than remaining sternites. Wing membranes with noticeable smoky golden brown tinge and iridescent. Head capsule with scarce medium setae and a few longer setae. Pronotum perimeter with alternating medium and long setae. Frons and median vertex rather ßat, and with weak slope between planes of frons vertex. Compound eyes small, suboval, and with rectate margins along of antennal sockets, and with distinct ocular sclerite. Ocelli white, very small, very near eyes; slightly pointed anterodorsally. Antennae with 12Ð13 articles, usually 13; formula Pronotum with anterior margin moderately concave, lateral margins distinctly convex, posterolateral corners subtruncate, and posterior rectate. Wing membranes with dense, distinct, and pointed nodulations; points darkly pigmented. Comparisons. The G. liberatus imago generally differs from that of the sympatric G. pubescens by the former having both a darker body and wing mem- Table 1. Measurements of Glyptotermes liberatus imago, n 4 males, 5 females from 5 colonies Measurement (mm) Range Mean SD Head length with labrum 1.28Ð Head width at eyes, maximum 1.08Ð Head height without 0.67Ð postmentum Eye diameter, maximum 0.29Ð Eye to head base, minimum 0.16Ð Ocellus diameter, maximum 0.22Ð Eye sclerite to ocellus, minimum 0.07Ð Pronotum length, maximum 0.64Ð Pronotum width, maximum 0.96Ð Total length with wings 8.91Ð Total length without wings 5.45Ð Fore wing length from suture 6.38Ð Fore wing width, maximum 1.68Ð Hind tibia length 0.96Ð

10 190 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 96, no. 3 branes. The general coloration of G. liberatus is dark brown and its wing membranes are distinctly tinged with golden brown, while G. pubescens is pale brown with hyaline membranes. The ocellar diameter of G. pubescens is approximately one-third the diameter of its eyes, whereas in G. liberatus the ocellar diameter is approximately one-þfth of the eye diameter. Remarks. Snyder (1929) gave the soldier that Banks (1919) described as Kalotermes ( Glyptotermes) posticus (Hagen) the new name, Kalotermes ( Glyptotermes) liberatus. This sample was collected from the Blue Mountains in Cinchona, Jamaica. Snyder (1929) reasoned that the imago described from St. Thomas as Calotermes posticus by Hagen (1858) was most likely a Cryptotermes because Hagen (1858) stated that his posticus is near Cr. brevis. According to Araujo (1977), HagenÕs type of Cr. posticus was destroyed. Without an available type, we suspect that HagenÕs posticus imago was probably Cr. brevis or one of more recently described kalotermitids found on St. Thomas. Because it is unlikely that a new Kalotermitid will be found on St. Thomas, we propose that posticus be relegated to nomen dubium status. Glyptotermes liberatus is a montane (500Ð1000 m) forest species that colonizes damp or wet wood of trees and palms. Alates have been collected from colonies in May and June. Material Examined and Measured. U.S. Territory. Puerto Rico: El Yunque Park, Sierra Palm trail; N, W; RS, JC, JM, JR; 31-V-93; 2 alates (PR141); Guavate Forest picnic area on hwy 184; N, W; RS, JC, JM, JR; 1-VI-93; 2 alates (PR197); Hwy 143 between hwy 139 and 140; N, W; RS, JC, JM, JR; 2-VI-93; 2 alates (PR224), two alates (PR226); Laguillo Natl. Forest, hwy 191 (Quebrada Caimitillo trail); N, W; Susan C. Jones, Christine A. Nalepa, and Elizabeth A. McMahan; 20-III-93; 1 alate. Distribution. Martorell (1973) Þrst reported this species from Puerto Rico. This species is also found on Cuba, Hispaniola, and Jamaica (Fig. 33). Glyptotermes pubescens Snyder (Figs. 12 and 25) Remarks. Glyptotermes pubescens is a widespread inland forest and lower montane inhabitant. Alates have been collected from roadside tree stumps and logs in May and June. Wolcott (1948) reports the occurrence of this species in structures. Distribution. Puerto Rico only (Fig. 33). Incisitermes bequaerti (Snyder) (Fig. 15) Remarks. Incisitermes bequaerti typically inhabits dry littoral forests. Distribution. The species is widespread from Cuba and the Bahamas to Culebra, but within the survey area, it is common only on Mona Island and the dry southwest of Puerto Rico (Fig. 32). Table 2. Measurements of Incisitermes incisus imago, n 5 males, 5 females from 6 colonies Measurement (mm) Range Mean SD Head length with labrum 1.31Ð Head width at eyes, maximum 1.13Ð Head height without 0.64Ð postmentum Eye diameter, maximum 0.34Ð Eye to head base, minimum 0.13Ð Ocellus diameter, maximum 0.16Ð Eye sclerite to ocellus, minimum 0Ð Pronotum length, maximum 0.83Ð Pronotum width, maximum 1.16Ð Total length with wings 9.44Ð Total length without wings 5.59Ð Fore wing length from suture 7.18Ð Fore wing width, maximum 2.34Ð Hind tibia length 1.01Ð Incisitermes furvus Scheffrahn (Fig. 16A and B) Remarks. Incisitermes furvus is a local and less common species in submontane dry forests and in dispersed trees of savannas. Its striking black alates have been collected from wood in May and June (Scheffrahn 1994). Distribution. This species is endemic to northwestern Puerto Rico where it is known from four localities (Fig. 32). Incisitermes incisus (Silvestri) Calotermes incisus Silvestri 1901: 2 [imago]; Silvestri 1903: 27 [imago, soldier, Þgs.] Kalotermes incisus; Snyder 1949: 16 [catal.] Incisitermes incisus; Krishna 1961: 356 [n. gen., new comb.] Imago. (Fig. 24; Table 2). In dorsal view, head capsule, pronotum, and abdominal sternites almost concolorous, ochraceous, except for subhyaline anteclypeus, darker tips of mandibles, and ferruginous stripe along posterior margins of all abdominal plates. Certain phenotypes with pale chestnut brown frons and mandibles, slightly darker transverse band beyond ocelli and brownish darkening of posterolateral areas of pronotum. Epicranial suture complete, but very faint. V-shaped pattern on frons and T-shaped mark on pronotum almost absent. Chevron pattern on pterothorax slight to moderate. Sclerotized wing venation ferruginous; wing membranes subhyaline with faint ochraceous tinge. Head capsule with scarce medium-sized setae; periphery of pronotum with alternating long and medium-sized setae and dense very short setae; pronotum interior glabrous. Anterior of vertex, and part of frons ßattened, and with small angle between planes of frons and vertex. Compound eyes moderately sized, roundly subtriangular, and rectate in front of antennal sockets. Ocelli white, large, oblique and abutted to eyes. Antennae with 15Ð18 articles, usually 17; anten-

11 May 2003 SCHEFFRAHN ET AL.: TERMITES OF PUERTO RICO 191 Table 3. Measurements of Incisitermes incisus small soldier, n 9 from 7 colonies Measurement (mm) Range Mean SD Head length to tip of mandibles 2.82Ð Head length to postclypeus 1.83Ð Head width, maximum 1.31Ð Head height, without postmentum 1.00Ð Left mandible length, tip to articulation of 1.36Ð ventral condyle Postmentum, length in middle 1.29Ð Postmentum width, maximum 0.52Ð Postmentum width, minimum 0.21Ð Pronotum width, maximum 1.23Ð Pronotum length, maximum 0.94Ð Hind tibia length 0.95Ð Total length 5.59Ð nal formula Pronotum with anterior margin slightly concave; sides divergent anteriorly; widely rounded posteriorly; and with posterior margin slightly emarginate. Anterior wing veins sclerotized including 6-branched radial sector; media unsclerotized; 15 arched and faintly sclerotized and pigmented subveins between media and radial sector. Wing membranes faintly nodulose. Arolia present. Comparisons. Incisitermes incisus can be confused with I. snyderi. Incisitermes incisus is distinguishable by the following characters: general body coloration in I. incisus alate is ochraceous, in I. snyderi it is pale orange. Abdominal sternites of I. incisus have a darker, ferruginous posterior, whereas those of I. snyderi are pale orange throughout. In I. incisus, the sclerotized wing veins are ferruginous without distal fading, whereas in I. snyderi they are pale ferruginous orange with distinct distal fading. Incisitermes snyderi has a southeastern Nearctic distribution with a Neotropical extension into Cuba whereas I. incisus is allopatrically distributed from Hispaniola to Venezuela. Soldier. (Figs. 17 and 18; Tables 3 and 4). Dimorphic to polymorphic as some forms are intermediate between large and small morph. In dorsal view, head capsule and labrum ferruginous to pale chestnut brown, except for darker, chestnut brown anterior frons. Mandibles almost black, in some specimens with dark chestnut base. Thoracic nota, abdominal tergites and sternites, and legs yellowish. Table 4. Measurements of Incisitermes incisus large soldier, n 8 from 4 colonies Measurement (mm) Mean SD Range Head length to tip of mandibles 3.96Ð Head length to postclypeus 2.77Ð Head width, maximum 1.53Ð Head height, without postmentum 1.28Ð Left mandible length, tip to articulation 1.52Ð of ventral condyle Postmentum, length in middle 2.11Ð Postmentum width, maximum 0.60Ð Postmentum width, minimum 0.21Ð Pronotum width, maximum 1.58Ð Pronotum length, maximum 1.30Ð Hind tibia length 1.23Ð Total length 8.11Ð In dorsal view, head distinctly elongate (large morph) or moderately elongate (smaller morphs), subrectangular, with sides parallel or faintly convex. In lateral view, head capsule moderately dorsoventraly ßattened. Head capsule with scarce medium-sized setae; periphery of pronotum with not very spaced medium-sized setae. Mandibles with distinct dentition; edge of right apical tooth serrate. Antennae in small morph with 13Ð15 articles, usually 14Ð15; in large morph with 12Ð15, usually 14. Antennal formula Eye spots whitish, small, and slightly oblique. Anterior margin of pronotum distinctly incised and serrate; sides of pronotum convergent, posterolateral corners rounded or subtruncate, and posterior margin slightly or faintly emarginate. Comparisons. Incisitermes incisus is traditionally confused with I. snyderi in the West Indies, although I. incisus is distinguishable by the following characters: the head capsule of I. incisus is more pigmented, particularly on the anterior frons, which is chestnut brown, compared with ferruginous in I. snyderi. In large soldiers, the hind tibiae of I. incisus are more inßated than those of I. snyderi. Material Examined and Measured. U.S. Territory. Puerto Rico: Two km S Caguas on hwy 1; N, W; 29-V-93; 2 alates, one small, three large soldiers (PR6); Bosque de Aguirre; N, W; 1-VI-93; 2 alates, one small, one large soldier (PR168), two alates, one small soldier (PR180); two km E Recio on hwy 3; N, W,1-VI-93; 2 small, two large soldiers (PR202); Wildlife Stn. on hwy three NE Humacao; N, W; 01-VI-93; 1 alate, one small soldier (PR208), one alate, one small soldier (PR214); Monte de Estado camp on hwy 366; N, W; 2-VI-93; 2 alates, two small, two large soldiers (PR247). All colony samples collected by RS, JC, JM, and JR. Remarks. Incisitermes incisus is the most common and widespread drywood termite species in most of the West Indies and will infest structural lumber. It was recently recorded as the most collected kalotermitid on St. John (Jones and Nalepa 2002). Snyder (1956) had recorded it only from Barbados, Mona Is., and St. Croix, owing to its misidentiþcation as I. snyderi in other locations (see Table 11). Distribution. Incisitermes incisus is the only termite species that was recorded on all of eight principle islands of the survey area (see Table 10; Fig. 34) and, as noted above, is widespread throughout the West Indies. Neotermes intracaulis n. sp. Scheffrahn and Krecek Imago. (Fig. 28; Table 5). In dorsal view, head capsule chestnut brown; pronotum, labrum, and mandible bases, ferruginous to ferruginous orange; anteclypeus yellowish, abdominal tergites ferruginous orange. Epicranial sutures distinct, lateral branches weakly sinuous. V-shaped pattern on frons distinct and compressed laterally. T-shaped mark on pronotum moderate. Chevron pattern on pterothorax distinct, pale chestnut brown. Femora yellow-white, tibiae and

12 192 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 96, no. 3 Fig. 34. Islands. Localities and distribution of Procryptotermes corniceps and Incisitermes incisus in Puerto Rico and the U.S. Virgin tarsi pale chestnut brown. Sclerotized wing venation pale chestnut brown; wing membranes smoky with pale brown tinge. Head capsule short, compact, and oval. Head capsule and pronotum with several dozen long and erect setae and scattered shorter setae. Vertex anterior and frons ßattened, and slightly angled from each other. Compound eyes large, oval, and with rectate margins along antennal sockets and ocelli. Ocelli white, very large, oblique and bordering eyes along three-þfths lengths of ocelli. Antennae with 18Ð19 articles, usually 19; antennal formula Pronotum with anterior margin weakly concave; lateral margins convex; posterolateral corners subtruncate; and posterior margin slightly concave or rectate. Anterior wing veins including media sclerotized; radial sector with 4Ð6 branches; branching begins approximately at two- Þfths of wing length from suture. Arolia present. Tibial spines long. Wing membranes faintly nodulose. Comparisons. The strikingly dark head of Ne. intracaulis compared with lighter body pigmentation, is unique among Neotermes of the region. Compound eyes of Ne. intracaulis are proportionally 2 larger in an area, than in Ne. castaneus. Also ocelli of Ne. intracaulis are distinctly larger than those of Table 5. Measurements of Neotermes intracaulis imago, n 3 males, 7 females from 6 colonies Measurement (mm) Range Mean SD Holotype Head length with labrum 1.55Ð Head width at eyes, 1.42Ð maximum Head height without 0.83Ð postmentum Eye diameter, maximum 0.56Ð Eye to head base, minimum 0.18Ð Ocellus diameter, maximum 0.25Ð Eye sclerite to ocellus, minimum Pronotum length, maximum 0.92Ð Pronotum width, maximum 1.42Ð Total length with wings 13.17Ð Total length without wings 7.05Ð Fore wing length from 9.98Ð suture Fore wing width, maximum 2.87Ð Hind tibia length 1.23Ð Ne. castaneus. Pilosity of Ne. castaneus pronotum is denser than that of Ne. intracaulis. Legs of Ne. intracaulis are bicolored, while they are monocolored in Ne. castaneus. Neotermes intracaulis is much smaller than Ne. mona in all measurements, except for maximum eye and ocellar diameters. The frons of Ne. intracaulis is ßat, whereas in Ne. mona it is concave. The compound eyes of Ne. intracaulis are oval with margins shortly rectate along antennal sockets, whereas those of Ne. mona are subcircular and broadly rectate along antennal sockets. The Ne. intracaulis imago is generally darker than that of Ne. holmgreni. The head capsule of former is chestnut brown, while pale chestnut brown and with ferruginous anterior frons in the latter; the V-shaped pattern on the frons of Ne. intracaulis is distinct and compressed laterally, compared with indistinct and not compressed in Ne. holmgreni; and the tibiae of the former are chestnut brown, ferruginous in the latter. The head capsule of Ne. intracaulis is covered with 0.2-mm-long setae, the setae of Ne. holmgreni are 3Ð4 shorter. The distinct diagonal subvenation interconnecting the Ne. holmgreni cubitus and media is absent in Ne. intracaulis. Soldier. (Fig. 10; Table 6). Monomorphic. In dorsal view, head capsule and labrum pale orange, except for darker, ferruginous orange anterior frons, frontal carinae, and antennal sockets. Anteclypeus pale yellow. Mandibles pitch black, except for chestnut bases. An- Table 6. Measurements of Neotermes intracaulis soldier, n 10 from 9 colonies Measurement (mm) Mean SD Range Head length to tip of mandibles 4.16Ð Head width, maximum 1.83Ð Head height, without postmentum 1.36Ð Labrum, maximum width 0.44Ð Left mandible length, tip to articulation 1.83Ð of ventral condyle Postmentum, length in middle 2.14Ð Postmentum width, maximum 0.62Ð Postmentum width, minimum 0.28Ð Pronotum width, maximum 1.88Ð Pronotum length, maximum 0.96Ð Hind tibia length 1.21Ð Total length 8.51Ð

13 May 2003 SCHEFFRAHN ET AL.: TERMITES OF PUERTO RICO 193 tennae with Þrst 3Ð4 articles ferruginous orange, paling distally. Thoracic nota, abdominal tergites and tibiae of fore legs yellowish. Sternites, and remainder of legs yellow-white. In dorsal view, head distinctly elongate, subrectangular, with sides parallel or slightly convergent posteriorly; posterior margin of head capsule emarginate. In lateral view, head capsule slightly dorsoventraly ßattened. Head capsule with scarce long setae; head generally with dispersed medium-sized setae; pronotum and posterior dorsum with scarce, medium-sized setae. Mandibles evenly sinuous in lateral outline, apically hooked 90, corpulent, and with compressed dentition; interior margin of apical tooth with wavy serration. Mandibular bases weakly rugose; each with a few dozen evenly spaced short setae. Labrum linguiform with slight median point. Antennae with 12Ð17 articles, usually 13Ð14; antennal formula Eye spots yellow-white, elongate, slightly oblique, inset, and surrounded by narrow, lighter area of cuticle. Anterior margin of pronotum evenly and shallowly concave; lateral sides of pronotum evenly convex; posterior margin of pronotum weakly convex, subparallel with anterior margin, and slightly emarginate in middle. Comparisons. The soldier of Ne. intracaulis differs from Ne. castaneus in that the former has a distinctly more elongate and narrower head than that of the latter and the third antennal article of Ne. intracaulis is almost 2 longer than second or fourth, while articles 2Ð4 are subequal in Ne. castaneus. The inner margin of right apical tooth is serrate and the basal humps of the mandible have 1Ð2 dozens setae in Ne. intracaulis. In N. castaneus, the inner margin of right apical tooth is even, and 5Ð10 setae are present on each hump. The large soldiers of Ne. mona exceed all measurements of Ne. intracaulis and the small soldiers of N. mona are greater for maximum head width ( 2.31 mm versus 2.15 mm), maximum labrum width ( 0.64 mm versus 0.51 mm), and maximum pronotum width ( 2.73 mm and 2.32 mm). The eye spots of N. mona are almost black, whereas colorless in N. intracaulis. The head of N. mona is subsquare and densely pilous, whereas subrectangular and scarcely adorned with setae in N. intracaulis. The head capsule of Ne. intracaulis is pale orange, whereas it is ferruginous orange in N. holmgreni. The mandibles of the former are evenly sinuous in lateral outline, whereas basally humped in the latter. Mandibular dentition of Ne. intracaulis soldiers is reduced, whereas in Ne. holmgreni it is conspicuous. The posterior margin of Ne. intracaulis pronotum is convex and parallel with the anterior margin of pronotum, in Ne. holmgreni, it is subrectate and not parallel with the anterior margin. Etymology. The species name originates from the Latin intra, for within, and caulis, for stalk or stem, describing colony loci in living stems of Leucaena glauca (L.) Benth. Type Material Measured. U.S. Virgin Islands. St. Croix: Sion Hill; N, W; PB; 3-III-1999; 4 alates, including HOLOTYPE, two soldiers (VI58), one soldier (VI141); two alates (VI143); The following samples collected by JK, BM, and JM. Mahogany Road E of hwy 765; N, W; 8-XI-2000; 1 soldier (VI214); Hwy 80 near Salt River Road; N, W; 8-XI-2000; 1 alate and one soldier (VI232); Hwy 751 end; N, W; 8-XI-2000; 1 soldier (VI247); Hwy mi. E hwy 663; N, W; 9-XI-2000; 1 alate, one soldier (VI332); Intersection of hwys 81 and 74; N, W; 11-XI-2000; 1 soldier (VI440); Hwy 669 at hwy 72, Holy Cross Church; N, W; 11-XI-2000; 1 alate, one soldier (VI475); Little Princess; N, W; 11-XI.2000; 1 alate (VI495), one soldier (VI496). Remarks. Neotermes intracaulis is unique among termites of the region in that it almost exclusively colonizes the living stems of the ubiquitous shrub, L. glauca (Mimosaceae). Colonies often occupy the entire stem of these weedy legumes creating a single large gallery in the sapwood. In comparison, Neotermes castaneus (Burmeister), found from Florida to Hispaniola, infests a wider host range of live trees and shrubs, and will more frequently infest dead wood than Ne. intracaulis. Other regional congeners infest primarily dead wood. Distribution. Neotermes intracaulis is known only from St. Croix (Fig. 33) and St. Barthélemy (French Antilles). Neotermes mona (Banks) (Fig. 11) Remarks. Neotermes mona is a localized and uncommon dampwood species inhabiting dry littoral forests and mangrove swamps. The colonies sometimes invade the xylem of living trees and shrubs. Distribution. In the study area, Ne. mona is conþned to the southern coast of Puerto Rico and the islands of Mona, Vieques, St. John, and St. Croix (new record, Fig. 33). The species also occurs also on Guana Islands (B.V.I.), Hispaniola, and the Turks and Caicos Islands (Scheffrahn et al. 1990, Krecek et al. 2000). Procryptotermes corniceps (Snyder) (Fig. 13) Remarks. A common and widespread species, occurring more often in coastal than inland habitats, and tending to occupy dry littoral shrub forests (Scheffrahn and Krecek 2001). After I. incisus, Proc. corniceps is the second most common kalotermitid in the studied area. Distribution. In the study area, Proc. corniceps was collected on all islands except for Buck Is. (Fig. 34). This species also occurs in the Dominican Republic, Jamaica, Turks and Caicos Islands, and the southeastern Bahamas (Scheffrahn and Krecek 2001). Rhinotermitidae Coptotermes havilandi Holmgren (Fig. 21) Remarks. This nonendemic species is associated with structural infestations and heavy damage. Cop-

14 194 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 96, no. 3 Fig. 35. Localities and distribution of Coptotermes havilandi, Heterotermes sp., and Prorhinotermes simplex in Puerto Rico and the U.S. Virgin Islands. totermes havilandi constructs carton nests in structural voids and tree trunks. Crepuscular dispersal ßights occur February through April. Distribution. Within the study area, it has been collected only in San Juan (Fig. 35). Coptotermes havilandi is a southeast Asian species that has been brought to parts of the West Indies, Brazil, and Florida (Su et al. 1997, Scheffrahn 2001). Heterotermes sp. (Figs. 20 and 29) Remarks. Descriptions of winged imagos of West Indian Heterotermes spp. are based, for the most part, on location of the median vein, number of wing hairs, and shape of notal margins (Snyder 1924b). Soldiers are separated mainly by relative abundance of head capsule hair. Snyder (1924b) suggested the existence of a single extremely variable species or, alternatively, a complex series of very closely related species, there being two extremes tenuis Hagen and convexinotatus Snyder with intergrading connecting species, which display characters of either extreme (Snyder 1924b). Recent collections of Heterotermes from the West Indies and Florida (Scheffrahn et al. 1994, Scheffrahn and Su 1995) seem to support the hypothesis of a single variable species, however, an extensive taxonomic investigation of Heterotermes has yet to be conducted. The genus Heterotermes represents a subterranean termite with wide environmental tolerance and signiþcant economic importance; no doubt the greatest of all subterranean termites in the West Indies. Heterotermes sp. forages on all wood and cellulosic debris in contact with the ground. Wolcott (1948) reported serious damage to military buildings by Heterotermes in Puerto Rico. Distribution. Heterotermes sp. inhabits all islands in this study except for Mona (Fig. 35) and all surveyed islands in the West Indies, except for Barbuda, Saba, and the northern Bahamas. Prorhinotermes simplex (Hagen) Remarks. This characteristic species, found near tidal zones and costal ßats, often nests in and under damp or rotting logs. Galleries are lined with mudlike fecal packing. Dispersal ßights in Florida occur primarily in December and January. Distribution. Prorhinotermes simplex occurs in southeastern Florida, western Cuba, and Puerto Rico. Termitidae Anoplotermes n. sp. (Fig. 1) Anoplotermes meridianus Emerson 1925: 421 [new name] Anoplotermes sp.; Wolcott 1948: 74 [record] Puerto Rico Remarks. This soldierless termite is an uncommon island endemic. The species lives in drier areas, and almost nothing is known about its ecology. Alates were collected under a rock in May. To avoid apparent synonymy with an uncertain West Indian species that Latreille (1805) named Termes morio and a termite from Argentina, Uruguay, and Paraguay that Silvestri (1903) originally named A. morio, Emerson (1925) renamed SilvestriÕs species as A. meridianus. Snyder (1949, 1956), unaware of this nomenclatural change, erroneously listed A. meridianus as occurring in Puerto Rico, Martinique, and Hispaniola. The Anoplotermes sp. in our Puerto Rican material is heterospeciþc with Anoplotermes spp. we have from Cuba, Bahamas, Hispaniola, and Guadeloupe, and is undoubtedly a new species. Revision of the soldierless termites of the New World may result in reassigning this and other West Indies species to new genera. Distribution. Our collections are all from coastal southern and western Puerto Rico (Fig. 36). Wolcott (1948) reported an Anoplotermes sp. from Pueblo Viejo, Puerto Rico (location unknown). Caribitermes discolor (Banks) Constrictotermes discolor Banks 1919: 489 [soldier, Þgs.] Parvitermes discolor; Emerson in Snyder 1949: 305, 376 [n. gen., n. comb., catal.]