Taxonomy, Distribution, and Notes On The Termites (Isoptera: Kalotermitidae, Rhinotermitidae, Termitidae) of Puerto Rico and The U.S. Virgin Islands

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 SU1
Ann. Entomol. Soc. Am. 96(3): 181201 (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 otsam 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, identication 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 33314. 2 Ohio State University, 102 Extension Entomology Building, 1991 Kenny Road, Columbus, OH 43210. 3 Terminix International, 1235B Eagles Landing Pkwy., Stockbridge, GA 30281. 4 Terminix International, 9390 North Florida Ave., Suite B, Tampa, FL 33612.
The rst 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 Snyders (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 dene some termites of the region. To unify these works with more recent ndings and to add substantially toward an understanding of termites of the entire West Indies, we
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surveyed termites on eight islands of Puerto Rico and the U.S. Virgin Islands. Based on this survey, we provide keys for the identication 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 2002. 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 tted with an ocular micrometer. Measurements were partly adopted from Roonwal (1970). The color scheme of Sands (1965) was used with relevant modications. 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 identication 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 25. For example, the formula 2 3 4 5 indicates that the second article is longer than the third and the third through fth 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 1,1,1,3,3,3hexamethyldisilazane (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 ArcView GIS version 3.0a software [Environmental Systems Research Institute (ESRI), Redlands, CA]. Co-
ordinates 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 inated and greater in median circumference than middle or hind tibiae; postclypeus greatly inated; 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 inated; nonsubterranean or soildwelling, if latter, often constructing foraging tubes above ground . . . . . . . . . . . . . . . 2 2. Nasutiform, i.e., head distinctly modied by long frontal projection with terminal exocrine orice; mandibles reduced to nonfunctional stubs (Figs. 25) . . . . . . . . . . . . . . . . . . 3 Head not distinctly modied by long frontal projection with terminal glandular orice; mandibles well developed, functional (Figs. 6 22) . . . . . . . . . . . . . . . . . . . . . . . . . 6 3. 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) . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 4. 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 1314-segmented (Fig. 4) . . . . . . . . . . . . . . . . . . . Parvitermes wolcotti
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Fig. 112. Scanning electron micrographs of termites from Puerto Rico and the U.S. Virgin Islands. Lateral view of Anoplotermes n. sp. with arrows indicating inated 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. 718); nonsubterranean . . . . . . . . . 7
Left mandible without marginal teeth in distal half (Figs. 19 22); subterranean or aboveground nests . . . . . . . . . . . . . . . . . . . 18 7. Head blackish, strongly phragmotic, truncate, cuboidal; mandibles not projecting much beyond labrum (Figs. 6 9) . . . . . . . . . . . . . 8
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Fig. 1322. 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.
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Head ferruginous orange, rectangular; mandibles projecting far beyond labrum (Figs. 10 18) . . . . . . . . . . . . . . . . . . . . . . . . . 11 8. Rugosity of head capsule faint or absent (Figs. 6 and 7) . . . . . . . . . . . . . . . . . . . . . . . . 9 Rugosity of head capsule conspicuous (Figs. 8 and 9) . . . . . . . . . . . . . . . . . . . . . . . 10 9. 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) . . . . . . . . . . . . . . . . 12 Soldier smaller, maximum head width 1.9 mm; mandible base glabrous or with few short setae near dorsal mandibular condyle (Figs. 1218) . . . . . . . . . . . . . . . . . . . . . . . 13 12. 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, ange, or ridge (Figs. 12 and 13) . . . . . . . . . . . . . . . . 14 Head capsule without frontal modications (Figs. 14 18) . . . . . . . . . . . . . . . . . . 15 14. 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) . . . . . . . . . . . . . . . . . 16 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 attened, more elevated . . . . . . . . . . . . . . . . . . 17 17. Mandibles with almost all edges nely, 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 and 18) . . . . . . . . . . . . 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. 20 22); nests subterranean or hidden in wood on ground . . . . . . . . . . . . . . . . . 19 19. 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 . . . . . . 20 20. 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. 2325) . . . . . 2 Two pigmented and sclerotized veins in costal margin of wing (Figs. 26 and 27) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13 2. 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. 24 and 25) . . . . . . . . . . . . . . . . . . . . . . 7 3. Arolia between tarsal claws absent . . . . . . . . . . . . . . . . . . . . . . . . . . Cryptotermes brevis Arolia between tarsal claws present . . . . . 4 4. 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 . . . . . . . . . . . . . . . . . 5 5. 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
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Fig. 2327. 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.
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Fig. 2831. 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) . . . . . . . . 9 8. 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 scle-
rite width; surface of head and pronotum densely textured by ne granulation . . . . . . . . . . . . . . . . . . . Glyptotermes liberatus 9. Media pigmented and sclerotized; large species, head width at eyes 1.4 mm . . . . . . . . . . 10 Media unpigmented and unsclerotized, frons convex and smooth; smaller species, head width at eyes 1.4 mm . . . . . . . . . . . . . 11 10. Frons attened 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;
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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.
12.
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wing membrane subhyaline with very faint nodulations . . . . . . . . . . . . . . . . . . . . 12 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 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) . . . . . . . . . . . . . . . . 16 Forewing membrane smooth; costal margin not convex in middle; median vein distinct, especially near suture . . . . . . . . . . . . . . 15 Forewing membrane reticulate; costal margin convex in middle; median vein absent . . . . . . . . . . . . . . . . . . . Prorhinotermes simplex 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 0.2 mm . . . . 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 0.4 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . Coptotermes havilandi Third antennal article much smaller than fourth . . . . . . . . . . . . . . . . . . . . . . . 17 Third and fourth antennal articles subequal . . . . . . . . . . . . . . . . . . . . . . . . 18 Head and pronotum pale brownish, wings very pale orange brown; postclypeus distinctly inated, 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 . . . . . . 19 General coloration pale dirty or orange brown; head noticeably darker than pronotum; ocelli distinctly protruding . . . . . . . . . . . . . . 20 19. 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 ights 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 species strict association with moisture-protected structural lumber. Cryptotermes havilandi (Sjo stedt) (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
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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 formers 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 conned 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 nely 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 at, 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 1213 articles, usually 13; formula 2 3 4 5. 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) Head length with labrum Head width at eyes, maximum Head height without postmentum Eye diameter, maximum Eye to head base, minimum Ocellus diameter, maximum Eye sclerite to ocellus, minimum Pronotum length, maximum Pronotum width, maximum Total length with wings Total length without wings Fore wing length from suture Fore wing width, maximum Hind tibia length Range 1.281.49 1.081.21 0.670.75 0.290.33 0.160.20 0.220.27 0.070.10 0.640.75 0.961.13 8.9110.64 5.456.78 6.387.58 1.682.04 0.961.14 Mean SD 1.34 0.063 1.12 0.040 0.71 0.022 0.30 0.012 0.18 0.015 0.24 0.014 0.09 0.0076 0.67 0.035 1.02 0.045 9.34 0.57 5.99 0.42 6.75 0.42 1.82 0.13 1.02 0.054
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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-fth 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), Hagens type of Cr. posticus was destroyed. Without an available type, we suspect that Hagens 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 (500 1000 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; 18.30N, 65.78W; RS, JC, JM, JR; 31-V-93; 2 alates (PR141); Guavate Forest picnic area on hwy 184; 18.10N, 66.05W; RS, JC, JM, JR; 1-VI-93; 2 alates (PR197); Hwy 143 between hwy 139 and 140; 18.18N, 66.48W; RS, JC, JM, JR; 2-VI-93; 2 alates (PR224), two alates (PR226); Laguillo Natl. Forest, hwy 191 (Quebrada Caimitillo trail); 18.30N, 65.78W; Susan C. Jones, Christine A. Nalepa, and Elizabeth A. McMahan; 20-III-93; 1 alate. Distribution. Martorell (1973) rst reported this species from Puerto Rico. This species is also found on Cuba, Hispaniola, and Jamaica (Fig. 33).
Table 2. Measurements of Incisitermes incisus imago, n 5 males, 5 females from 6 colonies Measurement (mm) Head length with labrum Head width at eyes, maximum Head height without postmentum Eye diameter, maximum Eye to head base, minimum Ocellus diameter, maximum Eye sclerite to ocellus, minimum Pronotum length, maximum Pronotum width, maximum Total length with wings Total length without wings Fore wing length from suture Fore wing width, maximum Hind tibia length Range 1.311.52 1.131.32 0.640.77 0.340.38 0.130.19 0.160.19 00.02 0.831.01 1.161.41 9.4412.64 5.596.12 7.1810.11 2.342.80 1.011.24 Mean SD 1.41 0.061 1.21 0.077 0.71 0.046 0.37 0.012 0.16 0.019 0.18 0.0088 0.01 0.0057 0.89 0.068 1.26 0.091 10.84 1.23 5.81 0.22 8.50 1.18 2.53 0.19 1.85 0.094
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, gs.] 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 attened, 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 1518 articles, usually 17; anten-
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).
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Table 3. Measurements of Incisitermes incisus small soldier, n 9 from 7 colonies Measurement (mm) Head length to tip of mandibles Head length to postclypeus Head width, maximum Head height, without postmentum Left mandible length, tip to articulation of ventral condyle Postmentum, length in middle Postmentum width, maximum Postmentum width, minimum Pronotum width, maximum Pronotum length, maximum Hind tibia length Total length Range 2.823.71 1.832.57 1.311.68 1.001.34 1.361.65 1.291.81 0.520.65 0.210.31 1.231.73 0.941.26 0.951.31 5.597.85 Mean SD 3.14 0.29 2.13 0.23 1.48 0.13 1.15 0.10 1.49 0.093 1.56 0.20 0.56 0.041 0.26 0.038 1.49 0.11 1.10 0.11 1.14 0.13 6.83 0.89
nal formula 2 3 4 5. 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) Head length to tip of mandibles Head length to postclypeus Head width, maximum Head height, without postmentum Left mandible length, tip to articulation of ventral condyle Postmentum, length in middle Postmentum width, maximum Postmentum width, minimum Pronotum width, maximum Pronotum length, maximum Hind tibia length Total length Mean SD 3.964.65 2.773.47 1.531.90 1.281.55 1.521.79 2.112.51 0.600.72 0.210.31 1.581.93 1.301.50 1.231.55 8.119.71 Range 4.27 0.27 3.11 0.26 1.76 0.12 1.41 0.10 1.67 0.095 2.33 0.16 0.66 0.041 0.28 0.033 1.82 0.11 1.39 0.079 1.41 0.10 9.16 0.60
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 attened. 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 1315 articles, usually 14 15; in large morph with 1215, usually 14. Antennal formula 2 3 4 5. 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 inated than those of I. snyderi. Material Examined and Measured. U.S. Territory. Puerto Rico: Two km S Caguas on hwy 1; 18.20 N, 66.05 W; 29-V-93; 2 alates, one small, three large soldiers (PR6); Bosque de Aguirre; 17.93 N, 66.15 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; 17.98 N, 65.92 W,1-VI-93; 2 small, two large soldiers (PR202); Wildlife Stn. on hwy three NE Humacao; 18.17 N, 65.75 W; 01-VI-93; 1 alate, one small soldier (PR208), one alate, one small soldier (PR214); Monte de Estado camp on hwy 366; 18.15 N, 66.97 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 misidentication 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
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Fig. 34. Localities and distribution of Procryptotermes corniceps and Incisitermes incisus in Puerto Rico and the U.S. Virgin Islands.
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 attened, 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-fths lengths of ocelli. Antennae with 18 19 articles, usually 19; antennal formula 2 3 4 5. 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 4 6 branches; branching begins approximately at twofths 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) Head length with labrum Head width at eyes, maximum Head height without postmentum Eye diameter, maximum Eye to head base, minimum Ocellus diameter, maximum Eye sclerite to ocellus, minimum Pronotum length, maximum Pronotum width, maximum Total length with wings Total length without wings Fore wing length from suture Fore wing width, maximum Hind tibia length Range 1.551.73 1.421.55 0.830.92 0.560.61 0.180.23 0.250.28 0 Mean SD Holotype 1.67 0.049 1.46 0.041 0.87 0.022 0.58 0.019 0.20 0.017 0.26 0.010 0 1.67 1.44 0.87 0.56 0.20 0.26 0 0.93 1.57 13.70 7.98 10.24 2.93 1.28
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 at, 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 3 4 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. AnTable 6. Measurements of Neotermes intracaulis soldier, n 10 from 9 colonies Measurement (mm) Head length to tip of mandibles Head width, maximum Head height, without postmentum Labrum, maximum width Left mandible length, tip to articulation of ventral condyle Postmentum, length in middle Postmentum width, maximum Postmentum width, minimum Pronotum width, maximum Pronotum length, maximum Hind tibia length Total length Mean SD 4.165.05 1.832.15 1.361.65 0.440.51 1.831.99 2.142.70 0.620.80 0.280.41 1.882.32 0.961.18 1.211.41 8.5110.11 Range 4.68 0.32 2.02 0.10 1.53 0.094 0.47 0.021 1.93 0.051 2.45 0.18 0.73 0.059 0.33 0.042 2.09 0.018 1.09 0.082 1.30 0.067 9.26 0
0.921.05 0.96 0.039 1.421.60 1.50 0.055 13.1713.97 13.57 0.25 7.058.11 7.51 0.37 9.9810.51 10.24 0.18 2.873.13 1.231.34 3.02 0.081 1.27 0.030
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tennae with rst 3 4 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 attened. 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 1217 articles, usually 1314; antennal formula 2 3 4 5. 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 2 4 are subequal in Ne. castaneus. The inner margin of right apical tooth is serrate and the basal humps of the mandible have 12 dozens setae in Ne. intracaulis. In N. castaneus, the inner margin of right apical tooth is even, and 510 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; 17.730N, 64.740W; 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; 17.736N, 64.837W; 8-XI-2000; 1 soldier (VI214); Hwy 80 near Salt River Road; 17.775N, 64.763W; 8-XI-2000; 1 alate and one soldier (VI232); Hwy 751 end; 17.768N, 64.743W; 8-XI-2000; 1 soldier (VI247); Hwy 66 0.5 mi. E hwy 663; 17.723N, 64.763W; 9-XI-2000; 1 alate, one soldier (VI332); Intersection of hwys 81 and 74; 17.750N, 64.743W; 11-XI-2000; 1 soldier (VI440); Hwy 669 at hwy 72, Holy Cross Church; 17.733N, 64.807W; 11-XI-2000; 1 alate, one soldier (VI475); Little Princess; 17.755N, 64.732W; 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. Barthe lemy (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 conned 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-
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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 ights 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 signicant 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 ats, often nests in and under
damp or rotting logs. Galleries are lined with mudlike fecal packing. Dispersal ights 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 Silvestris 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 heterospecic 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, gs.] Parvitermes discolor; Emerson in Snyder 1949: 305, 376 [n. gen., n. comb., catal.]
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Fig. 36. Localities and distribution of Anoplotermes n. sp., Caribitermes discolor, Parvitermes wolcotti, and Termes hispaniolae in Puerto Rico and the U.S. Virgin Islands.
Caribitermes discolor; Roisin et al. 1996: 782 [n. gen., n. comb.] Imago. (Figs. 27 and 30; Table 7). Previously undescribed. In dorsal view, head capsule, pronotum, and abdominal sternites almost concolorous, dark sepia brown; antennae appear even darker; several paler, sepia brown or brownish structures on head capsule, i.e., crescent-shaped mark on each anterolateral corner, and anterior bifurcation of often faint fontanelle which continues longitudinally in median area of vertex; anterior cranial elements, mandibles pale brown; postclypeus brown, labrum and anteclypeus pale yellow-brown with whitish area. Epicranial suture reduced to incomplete and faint posterior line. Comparatively distinct T-shaped mark on pronotum, while midline faint. Tibiae brownish; remainder of legs pale brownish. Head capsule and pronotum densely punctated by contrasting pale setal follicles. Brown chevron pattern on pterothorax rather inconspicuous, because poor contrast from underlying mesonotum. Wings with two sclerotized brown veins on anterior edge of both pair of wings and media, and cubitus; their branches dark sepia brown, particularly those proximal likewise posterior fringed narrow obscurations of radial sectors; wing membranes duskily sepia brownish. Practically all structures, except for anteTable 7. Measurements of Caribitermes discolor imago, n 5 males, 5 females from 5 colonies Measurement (mm) Head length with labrum Head length to postclypeus Head width at eyes, maximum Head height without postmentum Eye diameter, maximum Eye to head base, minimum Ocellus diameter, maximum Eye sclerite to ocellus, minimum Pronotum length, maximum Pronotum width, maximum Total length with wings Total length without wings Fore wing length from suture Fore wing width, maximum Hind tibia length Range 1.131.26 0.670.78 0.961.05 0.440.52 0.280.31 0.070.09 0.080.11 0.050.08 0.430.52 0.740.88 9.3111.31 5.055.99 7.458.91 1.982.57 1.151.24 Mean SD 1.20 0.043 0.72 0.039 1.01 0.023 0.48 0.028 0.30 0.010 0.08 0.0084 0.10 0.0093 0.07 0.011 0.47 0.033 0.79 0.049 10.36 0.61 5.56 0.29 8.30 0.47 2.31 0.17 1.19 0.029
clypeus and thoracic intersegmental membranes, densely, regularly, and conspicuously pilous with medium-length, subequal setae. Pronotum adorned with distinctly dark median setae on interior disc, versus longer and faintly pigmented setae on its perimeter. Head longitudinally suboval. Mandibles with pair of medium-sized setae near ventral articulations. Antennae with 15 articles; relative length formula 2 3 4 5. Compound eyes medium-sized and subcircular, semiglobular in dorsal view. Ocelli white, obliquely suboval, and rather protruding; ocelli remote from eyes by distance equal to minimum diameter of ocelli. Pronotum trapeziform; pronotum near and parallel with anterior margin proled by grooved depression, continuous with elevation of anterior margin; pronotum anterior margin straight and sides anteriorly rectate, angled at one-thirds into truncate posterior part, and with posterior margin distinctly biconvexly emarginate. Comparisons. Among darkly pigmented imagos of the region, Caribitermes discolor resembles Nasutitermes costalis and Termes hispaniolae. Termes hispaniolae is signicantly smaller than Ca. discolor, its third antennal article is much shorter than second. In Ca. discolor, the third antennal article is subequal to the second in length. The postclypeus of Ca. discolor is 3 wider than long, whereas in T. hispaniolae it is about twice as wide as long. Nasutitermes costalis is distinctly larger than Ca. discolor, exemplied by head length with labrum with 1.5 mm in the former and 1.26 mm in the latter, by head width at eyes with 1.5 mm and 1.05 mm, and total length with wings with 14 mm in the former, and 12 mm. The postclypeus, antennae, and head capsule are concolorous. Soldier. (Fig. 5; Table 8). Monomorphic. In dorsal view, head capsule and nasus pale brown with ochraceous tinge, anterior third of head with slightly darker transversal band, posterior of head paler brown; antennae dark brown, except for much paler two rst articles. Complete abdominal tergites and perimeter of all thoracic tergites brown. In dorsal view, head capsule pyriform, faintly constricted at anterior third. Nasus rather long, slightly conical. Mandibles vestigial, but with apical points. Head capsule with not too numerous medium-sized
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Table 8. Measurements of Caribitermes discolor soldier, n 14 from 9 colonies Measurement, mm Head length with nasus Head length without nasus Head width, maximum Nasus width at base Nasus width at middle Head height, without postmentum Pronotum, maximum width Pronotum, maximum length Hind tibia length Total length Range 1.271.42 0.760.84 0.730.81 0.150.16 0.090.10 0.490.55 0.360.40 0.140.15 0.780.88 2.803.30 Mean SD 1.35 0.045 0.80 0.024 0.77 0.026 0.16 0.0062 0.09 0.0052 0.51 0.021 0.38 0.012 0.14 0.0059 0.82 0.032 3.09 0.15
ests. Foragers build dark external tunnels to food sources and tend to infest decayed wood on the ground. Alates, present in decaying wood, were collected in May and June. A population of Ca. discolor also occurs in the drier littoral forests of southwest Puerto Rico (Genet et al. 2000). Distribution. Puerto Rico and Hispaniola (Dominican Republic). Two additional records, Cuba (Snyder 1956) and Culebra (Banks 1919), are apparently based on misidentication (see Table 11). Nasutitermes acajutlae (Holmgren) (Fig. 2) Remarks. A builder of large, brownish black arboreal carton nests with strong and resistant outer walls. The nests are connected to logs, buildings, and other foraging sites by conspicuous foraging tubes. It is abundant and widespread in drier coastal habitats. Workers are apt to bite the neck and shoulder skin of a collector who stands under the nest as it is being opened. Distribution. Within the survey area, this species is found in littoral forests. It has been collected from all islands except Mona (see Table 10). In Puerto Rico, Na. acajutlae is less common than Na. costalis and the former is more restricted to coastal areas (Fig. 37). Martorell (1971) reports Na. acajutlae on Culebra and the small cays off the east coast of Puerto Rico including Luis Pen a, Diablo, Lobos, Hicacos, Obispo, and Islas Pin eros and Ramos. Na. acajutlae is common on B.V.I. (Collins et al. 1997), and it occurs on Antigua, Montserrat, Trinidad (Thorne et al. 1994), and Guyana (Emerson 1925). Nasutitermes costalis (Holmgren) (Fig. 3) Remarks. This species constructs rather fragile arboreal or epigeal carton nests that are smaller than those of Na. acajutlae. Dark brown foraging tubes radiating from nests are commonly seen on structural walls, trees, and other vertical surfaces. Nasutitermes costalis is a sound, usually dry wood feeder that attacks wood on or above ground. This species is particularly common in submontane areas of Puerto Rico. On Puerto Rico, Na. costalis is noticeably more common
setae and few longer setae; nasus covered with short, dense, and anterolaterally projected setae. Antennae with 12 articles; rst 25 subequal in length. Comparisons. Caribitermes discolor soldiers differ from Parvitermes wolcotti by being larger (maximum head width is 0.73 0.81 mm versus 0.51 0.65 mm, respectively) and the posterior lobe of the Ca. discolor head capsule is dominant, whereas in Pa. wolcotti, the cephalic lobes are subequal. The nasus of Ca. discolor is slightly conical and concolorous, while in Pa. wolcotti it is almost cylindrical and with a darker tip. Material Examined and Measured. U.S. Territory, Puerto Rico: El Yunque Park near entrance on Hwy 191; 18.34 N, 65.77 W; 31-V-93; 2 alates (PR127); El Yunque Park, Sierra Palm trail; 18.30 N, 65.78 W; 31-V-93; 2 alates (PR140); Campo Elisa Coldberg on hwy 186; 18.33 N, 65.82 W; 31-V-93; 2 alates (PR153); Guavate Forest picnic area on hwy 184; 18.10 N, 66.05 W; 1-VI-93; 2 alates (PR192); De Alta Vega State Park; 18.42 N, 66.37 W; 4-VI-93; 2 alates (PR330); Maricao State Forest; 18.20 N, 66.98 W; 19-V-92; 2 soldiers each; N Las Marias, road 119; 18.28 N, 67.00 W; 21-V-92; 2 soldiers; S Las Marias, road 120; 18.26 N, 66.99 W; 21-V-92; 2 soldiers each; N Maricao, road 120; 18.18 N, 66.96 W; 21-V-92; 1 soldier; N. Sabana Grande, road 120; 18.17 N, 66.93 W; 1 soldier; Luquillo Natl. Park, hwys 191 and 930; 18.30 N, 65.78 W; 21-V-92; 4-XI-92; 1 soldier. Colony samples PR127330 were collected by RS, JC, JM, and JR; unnumbered samples were collected by S. Jones. Remarks. This species is a common subterranean inhabitant of moister montane and submontane for-
Fig. 37. Localities and distribution of Nasutitermes acajutlae and Na. costalis in Puerto Rico and the U.S. Virgin Islands.
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Table 9. Measurements of Parvitermes wolcotti imago, n 1 male, 7 females from 3 colonies Measurement (mm) Head length with labrum Head length to postclypeus Head width at eyes, maximum Head height without postmentum Eye diameter, maximum Eye to head base, minimum Ocellus diameter, maximum Eye sclerite to ocellus, minimum Pronotum length, maximum Pronotum width, maximum Total length with wings Total length without wings Fore wing length from suture Fore wing width, maximum Hind tibia length Range 1.521.62 0.820.90 1.181.24 0.510.57 0.380.43 0.070.09 0.160.20 0.030.05 0.770.83 1.161.28 14.1015.16 7.858.65 12.2412.50 3.173.43 1.421.50 Mean SD 1.57 0.038 0.84 0.026 1.23 0.023 0.55 0.025 0.40 0.012 0.08 0.075 0.19 0.011 0.04 0.0052 0.81 0.025 1.23 0.036 14.85 0.38 8.28 0.26 12.37 0.094 3.31 0.12 1.45 0.027
and widespread than Na. acajutlae, whereas on islands in the study area where both species occur, Na. acajutlae is more common (Fig. 37). Distribution. On smaller islands within the studied area, Na. costalis is uncommon (Vieques and St. Croix) or apparently absent (Mona Is., Culebra, St. Thomas, and St. John). Nasutitermes costalis is the most common and widespread termitid of the West Indies, occurring on almost all islands except for the Bahamas and a few remote islands of the northern Lesser Antilles. Parvitermes wolcotti (Snyder) (Figs. 4 and 31) Nasutitermes (Tenuirostritermes) wolcotti Snyder 1924a: 131 [soldier, gs.] Parvitermes wolcotti; Emerson in Snyder 1949: 305, 376 [n.gen., n. com., catal.] Scheffrahn and Roisin 1995: 595 [major, minor soldier, worker, gs.] Imago. (Fig. 31; Table 9). Previously undescribed. In dorsal view, head capsule dark orange brown with slightly paler anterolateral areas, fontanelle, and at insertions of densely spaced setae. Postclypeus orange-yellow; anteclypeus subhyaline. Antennae and mandibles ferruginous, except for much darker dentition. Pronotum pale ochraceous; posterior areas of pterothorax and abdominal tergites brown. Wing membranes smoky ochraceous, venation brown. In lateral view, head capsule moderately attened dorsoventrally; in dorsal view, head capsule ovoid because of elongated mandibles; eyes and ocelli large, both oval, and narrowly separated by distance shorter than half of ocellus minimum diameter; ocelli protruding, hyaline with white bottom. Head capsule, postclypeus, and labrum pilous; pronotum, legs, and all abdominal tergites and sternites covered with long setae. Antennae with 16 articles; formula variable but usually 2 3 4 5. Comparisons. The Pa. wolcotti imago differs from two other described congeners, Pa. dominicanae and Pa. tousainti, as follows: the Pa. tousainti head is a
much darker, sepia brown, its pronotum is almost concolorous with head capsule, and its ocelli are much smaller, less protruded, and removed by one ocellar diameter from the eyes. In Pa. wolcotti, the head is orange-brown and its pronotum is paler, pale ochraceous, its ocelli are very large, protruding, and removed from the eyes by a distance shorter than half of its minimum diameter. Parvitermes dominicanae is similar to Pa. wolcotti with the exception that Pa. dominicanae possesses subcircular eyes, its ocelli are smaller and less protruding, and are separated from the eyes by more than one-half of the minimal ocellar diameter. In Pa. wolcotti the eyes are suboval and the ocelli are very large, protruding, and separated from eyes by a distance shorter than half of ocellus minimum diameter. Material Examined and Measured. U.S. Territory, Puerto Rico: Guanica Commonwealth Forest; 17.97 N, 66.87 W; 7-VII-97; 2 alates (PR834), two alates (PR835); Guanica Commonwealth Forest; 18.01 N, 66.87 W; 5-VII-98; 4 alates (PR836). The colony samples collected by J. Genet. Remarks. This species builds light brown foraging tubes and sheets. They nest in soil or under rocks. Parvitermes wolcotti occurs sparsely in dry forests. On Vieques, Pa. wolcotti foragers were collected under a log within 7 m of the shoreline and under a rock at the relatively lush summit of Mt. Pirata at 300m. Dispersal ights commence at different seasons on different islands depending on climatic conditions. JK observed night ights during rain at October 2200 hours, whereas on Puerto Rico the alates were collected under rocks in May. Rare major soldiers of this species (Scheffrahn and Roisin 1995) lack a head constriction. Distribution. Previous records include only Puerto Rico (Snyder 1924a), St. John (Jones and Nalepa 2002), and four British Virgin Is. (Scheffrahn et al. 1994). Three new island records include Vieques, Culebra, and St. Thomas (Fig. 36). Termes hispaniolae (Banks) (Fig. 19) Remarks. A common and widespread arboreal species on many islands in the West Indies. T. hispaniolae builds diffuse nests in solid hard wood, tree hollows or crotches, especially near sandy beaches. Nests are made of thick, black, and solid carton. Foragers are often active in more decayed wood in scares and crevices. Distribution. Within the studied area, this species occurs only on Puerto Rico where it is uncommon along western coasts, St. Croix, where it is common, and on St. Thomas, where is uncommon. (Fig. 36). Summary Table 10 includes the species by island records for Puerto Rico and the U.S. Virgin Islands based on this survey and our recognition of doubtful or erroneous records (Table 11). Of the 21 species occurring in Puerto Rico and the Virgin Islands, 12 are West Indian
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Table 10. Termites of Puerto Rico and the U.S. Virgin Islands based on current survey. Listings for Hispaniola and Guadeloupe show only those species in common with Puerto Rico and the U.S. Virgin Islands. Islands listed from largest to smallest Species Kalotermitidae Cryptotermes brevisa Cr. havilandia Cr. rotundicepsb Cr. undulansc Glyptotermes liberatusb G. pubescensc Incisitermes bequaertib I. furvusc I. incisusd Neotermes intracaulisb Ne. monab Procryptotermes cornicepsb Rhinotermitidae Coptotermes havilandia Heterotermes sp.d Prorhinotermes simplexd Termitidae Anoplotermes n. sp.c Caribitermes discolorb Nasutitermes acajutlaed Na. costalisd Parvitermes wolcottic Termes hispaniolaed Total no. endemic speciese
a c b
Hispaniola Puerto Rico Guadeloupe St. Croix Vieques St. Thomas St. John Mona Culebra Buck 17
30
14
6 4
Recent immigrant pest. Endemic to parts of West Indies only. Endemic to Puerto Rico and/or Virgin Islands only. d Endemic to West Indies and New World mainland. e Unpublished survey data.
endemics, six also occur on the Central and South American mainland (Constantino 1998), and three are nonendemic pests. Four species found in Puerto Rico and the U.S. Virgin Islands also occur on the U.S. mainland (Florida) as introduced pests of unknown origins. The latter include Co. havilandi (Su et al. 1997), Cr. brevis (Scheffrahn et al. 2000a), Heterotermes sp. (Scheffrahn and Su 1995), and Na. costalis (Scheffrahn et al. 2002). With the exception of Na. costalis on Tortola and Guana, the termite fauna of all the larger British Virgin Islands (Scheffrahn et al. 1994, Collins et al. 1997) constitute the same six species endemic to St. John. We suspect that Cr. brevis occurs on all inhabited islands (Scheffrahn and Krecek 1999).
Table 11. Island Culebra Mona Puerto Rico
Discussion Two principle models have been proposed to explain the biogeography of West Indian vertebrates, probably the most studied of all animal groups in the region (Hedges 2001). These include: (1) vicariance (i.e., allopatric speciation or fragmentation of a previously unfragmented range) of fauna on Protoantillean land masses between North and South America during the late Cretaceous (Rosen 1975), and (2) over-water dispersal of fauna from South America during the Cenozoic (Hedges 1996). A recent hypothesis (Itirralde-Vincent and MacPhee 1999) suggests that dispersal was facilitated by a land bridge between
Doubtful or erroneous termite species records for Puerto Rico and the U.S. Virgin Islands Record in doubt Ca. discolor I. snyderi (Light) I. milleri (Emerson) I. schwarzi (Banks) I. snyderi Cr. cavifrons Banks Ne. castaneus (Burmeister) Microcerotermes arboreus Emerson A. meridianus Emerson Cr. longicollis Banks I. snyderi Cr. cavifrons Termes panamensis (Snyder) Na. ephratae (Holmgren) G. posticus (Hagen) Reference Banks 1919 Ramos 1946 Luykx et al. 1990 Araujo 1977 Martorell 1945 Snyder 1956 Wolcott 1948 Burmeister 1839 Wolcott 1948 Snyder 1949 Scheffrahn et al. 1994 Snyder 1956 Snyder 1956 Snyder 1956 Snyder 1956 Banks 1919 Conclusion Mistaken for Pa. wolcotti Mistaken for I. incisus or I. bequaerti Mistaken for I. bequaerti Mistaken for I. incisus or I. bequaerti Mistaken for I. incisus or I. bequaerti Mistaken for I. incisus or I. bequaerti Mistaken for a n. sp., Cr. undulans Easternmost range Hispaniola Nearest congener Cayman Is., Trinidad, or mainland Argentina sp., not conspecic with Puerto Rico sp. Mistaken for a n. sp., Cr. rotundiceps Mistaken for I. incisus Mistaken for a new sp., Cr. undulans Mistaken for T. hispaniolae Mistaken for Na. costalis Nomen dubium
St. Croix
St. Thomas
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the Greater Antilles and South America via an emergent Aves Ridge during the mid-Cenozoic. Because of the complex and uncertain geologic history of the West Indies, especially the location and form of dry land (Donovan and Jackson 1994), a unifying model of the regions biogeography is still outstanding. Therefore, conclusions on West Indian termite biogeography must rely on current faunal distributions and the few fossil records available. It does appear certain that the biogeographical clock for the Protoantilles was reset at the dawn of the Cenozoic some 65 mya as the result of a cataclysmic extinction of most, if not all, life from a bolide impact in the western Caribbean (Hedges et al. 1992). Dominican amber, aged at 1520 million years old (Itirralde-Vincent and MacPhee 1996), constitutes the only fossil records of termites in the West Indies. These include the primitive genus Mastotermes now restricted to the Australian Region (Poinar and Poinar 1994) and the Neotropical genera Glyptotermes (Scheffrahn unpubl.), Cryptotermes (Krishna and Bacchus 1987), Constrictotermes, and Nasutitermes (Krishna 1996). These discoveries show that the termite fauna of Hispaniola during the upper Tertiary was a synthesis of extant and extinct higher taxa. Current faunal similarities between Hispaniola and Puerto Rico hint that the paleofauna of Puerto Rico underwent a similar evolution. Tectonic reconstructions (Ross and Scotese 1988) even suggest that Puerto Rico and Hispaniola had an Oligocene to Early Miocene connection. Over-water colonization ights by termite alates over distances greater than hundreds of meters are implausible. Not only are termites weak iers and have dehiscent wings, but also more importantly, mating occurs after wings are shed. This precludes aerial dispersal by an inseminated female termite but allows the same with a volant solitary insect group such as butteries (Miller and Miller 2001). Long-distance dispersal of termites in otsam, however, is altogether likely. One could easily envision storm-dislodged otsam such as a log colonized by drywood or dampwood termites or an uprooted tree carrying an arboreal Nasutitermes or Termes nest rafting in oceanic currents for hundreds of kilometers before making landfall. Heatwole and Levins (1972) found hundreds of live termites (species not reported) in driftwood otsam at sea off Puerto Rico and Wheeler (1916) reported a live Pheidole ant colony in beached otsam in Brazil. A modern account even documents the rafting of iguanas on hurricane-dislodged otsam from Guadeloupe to Anguilla (Censky et al. 1998). Several times we have observed live termite colonies in beached driftwood but could not determine whether the wood was colonized before or after landfall. Because of the apparent dispersalist abilities of some termites in otsam, colonization of new land masses could be the result of ancient as well as much more recent events. Puerto Rico, and the West Indies in general, is depauperate of termite species that are poor overwater dispersalists, especially soil-dwelling termitids, which constitute the bulk of termite diversity in the
mainland tropics. It is difcult to envision over-water dispersal for the semiobligate or obligate soil nesting species including Anoplotermes sp., Ca. discolor, or Pa. wolcotti. Some vicariate origins may be more plausible for these, however, the recent discovery of a rainforest-inhabiting Anoplotermes on Guadeloupe (R.H.S. et al., unpublished) suggests that over-water dispersal for even soil-dwelling species cannot be fully discounted. As with other West Indian insects (Liebherr 1988) and vertebrates (Hedges 1996), there is a moderate proportion of regional or island endemism among Puerto Rican termites. Of Puerto Ricos 17 endemic species, ten are shared with Hispaniola, three with the Virgin Islands only, three are restricted solely to the island of Puerto Rico, and one, Pror. simplex, is shared with Cuba and Florida (Table 10). Only three of Puerto Ricos endemic species are widespread and found on the mainland and the Greater and the Lesser Antilles, including the highly vagile species I. incisus, T. hispaniolae, and Na. costalis (Constantino 1998, Issa 2000). The termite fauna of the Puerto Rico and the Virgin Islands suggests a biogeographical origin that is derived from both western and southeastern sources. Specically, the fauna of Puerto Rico and the Virgin Islands encompass the eastern-most distributions of ve Antillean kalotermitids, including Cr. rotundiceps, G. liberatus, I. bequaerti, Ne. mona, and Proc. corniceps, the rhinotermitid, Pror. simplex, and the small nasutitermitine, Ca. discolor. A western afnity is also suspected for ancestral stock of some apparently autochthonous species including Pa. wolcotti, whose congeners occur only on Hispaniola, Cuba, and the Bahamas; Incisitermes furvus which resembles its western congeners such as I. milleri (Emerson) and I. rhyzophorae Herna ndez; and Cr. undulans which is closest to the Hispaniolan Cr. mangoldi Scheffrahn and Krecek. The Anoplotermes n. sp. from Puerto Rico is also more closely aligned morphologically with congeners from Hispaniola and Cuba than with the Anoplotermes sp. on Guadeloupe (T.G. Myles, personal communication, R.H.S. unpubl.). No mainland distribution has been identied for any of these species. Current distributions of some endemic species, which are all good candidates for dispersal, suggest an origin in mainland South America. The ancestral origin of the island endemic, G. pubescens, points toward northern South America with a dispersal track along the Lesser Antilles, where numerous congeners reside. Nasutitermes acajutlae has a disjunct distribution that includes the Puerto Rican Bank, Trinidad, and Venezuela, but is not found on the intervening Lesser Antilles. Nasutitermes costalis ranges broadly from South America through the Lesser Antilles, Greater Antilles, and the Turks and Caicos Islands. Likewise, I. incisus occurs from Venezuela to the Turks and Caicos, but is not found in Jamaica. Termes hispaniolae also occurs throughout the West Indies as well as Brazil, Guyana, Panama, and Venezuela.
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The termite fauna of St. Croix reects the islands remoteness from the Puerto Rican Bank and its proximity to islands east of the Anegada passage. Of the regional endemics, it lacks Pa. wolcotti, but supports Cr. undulans. Furthermore, St. Croix shares the distribution of Ne. intracaulis with St. Barthe lemy. St. Croix shares T. hispaniolae with much of the West Indies, but not with the other Virgin Islands. The remoteness of Mona island is reected in its depauperate, dispersalist fauna that includes four kalotermitids found also on Hispaniola as well as Puerto Rico. Based on an almost complete survey of West Indian termites, the Anegada Passage, a wide deep-water channel between the Greater Antilles (including the Virgin Islands) and the Lesser Antilles to the southeast, demarcates two major termite faunal zones of the region. To the west of this passage are found the nasutitermitine genera of Puerto Rico, Hispaniola, and Cuba including Antillitermes, Caribitermes, Constrictotermes, and Parvitermes and the termitid genera Amitermes and Anoplotermes. To the east and south of the Anegada Passage are found the kalotermitid genera Calcaritermes and Comatermes. With the exception of Anoplotermes in Guadeloupe and Rhinotermes in Hispaniola, distributions of these two genera are also limited to the Greater Antilles or Lesser Antilles, respectively. Acknowledgments
The authors thank Paul Ban, John Genet, Peter Luykx, Elizabeth McMahan, Christine Nalepa, Juan Rivera, Julian de la Rosa, Juan Torres, Mark Weinberg, and Abbie Zeltzer for additional valuable collections and assistance. We are grateful to Diann Achor at the University of Florida, Lake Alfred Citrus Research and Education Center, for her assistance with scanning electron microscopy; and W. H. Kern, Jr. and B. J. Cabrera for their critical reviews. We thank the United States Navy for permission to collect on their property in Vieques; Rafe Boulon, U.S. Department of the Interior, for permission to collect in Virgin Islands National Park, St. John; and Miguel Canals, Jose L. Chabert, Gerardo Hernandez, Guillermo Riera, and Jose Silva, Departamento de Recursos Naturales y Ambientales, for assistance in the Commonwealth parklands of Puerto Rico. Florida Agricultural Experiment Station, Journal Series No. R-08395.
References Cited
Araujo, R. L. 1977. Cata logo dos Isoptera do Novo Mundo. Academia Brasileira de Cie ncias, Rio de Janeiro, Brazil. Banks, N. 1919. Antillean Isoptera. Bull. Mus. Comp. Zool. 26: 475 489. Banks, N., and T. E. Snyder. 1920. A revision of the Nearctic termites. Bull. United States Natl. Mus. 108: 1228. Burmeister, H. 1839. Handbuch der Entomologie, pp. 758 768. Reimer, Berlin. Censky, E. J., K. Hodge, and J. Dudley. 1998. Over-water dispersal of lizards due to hurricanes. Nature (Lond.) 395: 556. Collins, M. S., M. I. Haverty, and B. L. Thorne. 1997. The termites (Isoptera: Kalotermitidae, Rhinotermitidae, Termitidae) of the Br. Virgin Islands: distribution, moisture relations, and cuticular hydrocarbons. Sociobiology 30: 6376.
Constantino, R. 1998. Catalog of the living termites of the New World (Insecta: Isoptera). Arq. Zool. (Sa o Paulo) 35: 135231. Emerson, A. E. 1925. The termites from Kartabo, Bartica District, Guyana. Zoologica 6: 291 459. Darlington, J.P.E.C. 1992. Survey of termites in Guadeloupe, Lesser Antilles (Isoptera: Kalotermitidae, Rhinotermitidae, and Termitidae). Florida Entomol. 75: 104 109. Donovan, S. K., and T. A. Jackson [eds.]. 1994. Caribbean geology: an introduction. Univ. West Indies Publ. Assoc., Kingston, Jamaica. Gay, F. J. 1967. A world review of introduced species of termites. Commonwealth Scientic and Industrial Res. Organization, Melbourne, Australia, Bull. 286: 1 88. Genet, J. A., K. S. Genet, T. M. Burton, and P. G. Murphy. 2000. Quantitative characterization of a subtropical dry forest termite community (Isoptera: Kalotermitidae, Rhinotermitidae, and Termitidae). Trop. Ecol. 41: 119 122. Hagen, H. A. 1858. Specielle Monographie der Termiten. Linnea Entomologica 12: 4 342. Heatwole, H., and R. Levins. 1972. Biogeography of the Puerto Rican Bank: otsam transport of terrestrial animals. Ecology 53: 112117. Hedges, S. B. 1996. Historical biogeography of West Indian vertebrates. Annu. Rev. Ecol. Syst. 27: 163196. Hedges, S. B. 2001. Biogeography of the West Indies: An overview, pp. 1533. In C. A. Woods and F. E. Sergile [eds.], Biogeography of the West Indies, patterns and perspectives, 2nd ed. CRC, Boca Raton, FL. Hedges, S. B., C. A. Hass, and L. R. Maxon. 1992. Caribbean biogeography: molecular evidence of dispersal in West Indian terrestrial vertebrates. Proc. Natl. Acad. Sci. U.S.A. 89: 1909 1913. Issa, S. 2000. A checklist of the termites from Venezuela (Isoptera: Kalotermitidae, Rhinotermitidae, Termitidae). Florida Entomol. 83: 379 382. Itirralde-Vincent, M. A., and R.D.E. MacPhee. 1996. Age and paleogeographical origin of Dominican amber. Science 273: 1850 1852. Itirralde-Vincent, M. A., and R.D.E. MacPhee. 1999. Paleogeography of the Caribbean region: implications for Cenozoic biogeography. Bull. Am. Mus. Nat. Hist. 238: 195. Jones, S. C., and C. A. Nalepa. 2002. Survey of the termites (Isoptera: Kalotermitidae, Rhinotermitidae, Termitidae) of St. John, U. S. Virgin Islands. Sociobiology 39: 155163. Jones, S. C., C. A. Nalepa, E. A. McMahan, and J. A. Torres. 1995. Survey and ecological studies of the termites (Isoptera: Kalotermitidae) of Mona Island. Florida Entomol. 78: 305313. Krecek, J., N.-Y. Su, and R. H. Scheffrahn. 2000. Redescription of Neotermes mona, a dampwood termite (Isoptera, Kalotermitidae) from the central West Indies. Florida Entomol. 83: 268 275. Krishna, K. 1961. A generic revision and phylogenetic study of the family Kalotermitidae (Isoptera). Bull. Am. Mus. Nat. Hist. 122: 303 408. Krishna, K. 1996. New fossil species of termites of the subfamily Nasutitermitinae from Dominican and Mexican Amber (Isoptera, Kalotermitidae). Am. Mus. Nov. no. 3176: 113. Krishna, K., and S. Bacchus. 1987. A new fossil species of termite from Dominican Amber, Cryptotermes yamini (Isoptera, Kalotermitidae). Am. Mus. Nov. no. 2884: 15. Liebherr, J. K. (ed.). 1988. Zoogeography of Caribbean insects. Cornell University Press, Ithaca, NY.
May 2003
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Martorell, L. F. 1945. A survey of the forest insects of Puerto Rico. J. Agric. Univ. Puerto Rico 29: 69 608. Martorell, L. F. 1971. New locality and host records for two species of Nasutitermes (Isoptera: Termitidae). J. Agric. Univ. Puerto Rico 55: 135137. Martorell, L. F. 1973. Glyptotermes liberatus (Snyder) (Isoptera: Kalotermitidae): a new termite record for Puerto Rico. J. Agric. Univ. Puerto Rico 57: 355356. Miller, J. Y., and L. D. Miller. 2001. The biogeography of the West Indian Butteries (Lepidoptera): an application of a vicariance/dispersalist model, pp. 127155. In C. A. Woods and F. E. Sergile [eds.], Biogeography of the West Indies, patterns and perspectives, 2nd ed. CRC, Boca Raton, FL. Nation, J. A. 1983. A new method using hexamethyldisilazane for the preparation of soft insect tissue for scanning electron microscopy. Stain Technol. 55: 347352. Poinar, G., and R. Poinar. 1994. The quest for life in amber. Addison-Wesley, Reading, MA. Ramos, J. A. 1946. The insects of Mona Island (West Indies). J. Agric. Univ. Puerto Rico 30:174 2 plates. Roisin, Y., R. H. Scheffrahn, and J. Krecek. 1996. Generic revision of the smaller nasute termites of the Greater Antilles (Isoptera, Termitidae, Nasutitermitinae). Ann. Entomol. Soc. Am. 89: 775787. Roonwal, M. L. 1970. Measurements of termites (Isoptera) for taxonomic purposes. J. Zool. Soc. India 21: 9 66. Rosen, D. E. 1975. A vicariance model of Caribbean biogeography. Syst. Zool. 24: 431 464. Ross, M. I., and C. R. Scotese. 1988. A hierarchical tectonic model of the Gulf of Mexico and Caribbean region. Tectonophysics 155: 139 168. Sands, W. A. 1965. A revision of the termite subfamily Nasutitermitinae (Isoptera, Termitidae) from the Ethiopian Region. Bull. Br. Mus. Nat. Hist., Entomol. Suppl. 4: 1172. Scheffrahn, R. H. 1994. Incisitermes furvus, a new drywood termite (Isoptera: Kalotermitidae) from Puerto Rico. Florida Entomol. 77: 365372. Scheffrahn, R. H. 2001. A rst for Florida. Pest Control Technol. 29: 54. 56: 65 66. Scheffrahn, R. H., and J. Krecek. 1999. Termites of the genus Cryptotermes Banks (Isoptera: Kalotermitidae) from the West Indies. Insecta Mundi 13: 111171. Scheffrahn, R. H., and J. Krecek. 2001. New World Revision of the Termite Genus Procryptotermes (Isoptera: Kalotermitidae). Ann. Entomol. Soc. Am. 94: 530 539. Scheffrahn, R. H., and Y. Roisin. 1995. Antillean Nasutitermitinae (Isoptera: Termitidae): Parvitermes collinsae, a new subterranean termite from Hispaniola and redescription of P. pallidiceps and P. wolcotti. Florida Entomol. 78: 585 600. Scheffrahn, R. H., and N.-Y. Su. 1995. A new subterranean termite introduced to Florida: Heterotermes Froggatt (Rhinotermitidae: Heterotermitinae) established in Miami. Florida Entomol. 78: 623 627. Scheffrahn, R. H., N.-Y. Su, Y., and B. Diehl. 1990. Native, introduced, and structure-infesting termites of the Turks
and Caicos Islands, B.W.I. (Isoptera: Kalotermitidae, Rhinotermitidae, Termitidae). Florida Entomol. 73: 622 627. Scheffrahn, R. H., J.P.E. C. Darlington, M. S. Collins, J. Krecek, and N.-Y. Su. 1994. Termites (Isoptera: Kalotermitidae, Rhinotermitidae, Termitidae) of the West Indies. Sociobiology 24: 213238. Scheffrahn, R. H., N.-Y. Su, J. A. Chase, J. R. Mangold, J. K. Grace, and J. R. Yates. 2000a. First record of Cryptotermes cynocephalus Light (Isoptera: Kalotermitidae) and natural woodland infestations of C. brevis (Walker) on Oahu, Hawaiian Islands. Proc. Hawaiian Entomol. Soc. 34: 141145. Scheffrahn, R. H., J. Krecek, and N.-Y. Su. 2000b. Redescriptions of the Dampwood Termites, Neotermes jouteli and N. luykxi (Isoptera: Kalotermitidae) from Florida, Cuba, Bahamas, and Turks and Caicos. Ann. Entomol. Soc. Am. 93: 785794. Scheffrahn, R. H., B. J. Cabrera, W. H. Kern Jr., and N.-Y. Su. 2002. Nasutitermes costalis (Isoptera: Termitidae) in Florida: rst record of a non-endemic establishment by a higher termite. Florida Entomol. 85: 273275. Silvestri, F. 1901. Nota preliminare sui termitidi Sud-Americani. Bollettino dei Musei di Zoologia e Anatomia Comparata della Universitdi Torino 16(389): 1 8. Silvestri, F. 1903. Contribuzione alla conoscenza dei Termiti e Termitoli dellAmerica Meridionale. Redia 1: 1234. Snyder, T. E. 1924a. Description of a new termite from Porto Rico Proc. Entomol. Soc. Washington 26: 131132. Snyder, T. E. 1924b. Descriptions of new species and hitherto unknown castes of termites from America and Hawaii. Proc. United States National Museum 64: 1 45. Snyder, T. E. 1929. New termites from the Antilles and Middle America. Proc. Entomol. Soc. Washington 31: 79 87. Snyder, T. E. 1949. Catalog of the termites (Isoptera) of the world. Smithsonian Misc. Coll. 112: 1 490. Snyder, T. E. 1956. Termites of the West Indies, the Bahamas and Bermuda. J. Agric. Univ. Puerto Rico 40: 189 202. Su, N.-Y., R. H. Scheffrahn, and T. Weissling. 1997. A new introduction of a subterranean termite, Coptotermes havilandi Holmgren (Isoptera: Rhinotermitidae) in Miami, Florida. Florida Entomol. 80: 408 411. Thorne, B. L., M. I. Haverty, and M. S. Collins. 1994. Taxonomy and biogeography of Nasutitermes acajutlae and N. nigriceps (Isoptera: Termitidae) in the Caribbean and Central America. Ann. Entomol. Soc. Am. 87: 762770. Walker, F. 1853. List of the specimens of neuropterous insects in the collections of the British Museum, pp. 501 529. British Museum (Nat. Hist.) Wheeler, W. M. 1916. Ants carried in a oating log from the Brazilian mainland to San Sebastian Island. Psyche 23: 180 183. Wolcott, G. N. 1948. The insects of Puerto Rico, Isoptera: Termites. J. Agric. Univ. Puerto Rico 32: 6274. Received for publication 4 October 2002; accepted 3 February 2003.

Taxonomy, Distribution, and Notes On The Termites (Isoptera: Kalotermitidae, Rhinotermitidae, Termitidae) of Puerto Rico and The U.S. Virgin Islands

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Taxonomy, Distribution, and Notes On The Termites (Isoptera: Kalotermitidae, Rhinotermitidae, Termitidae) of Puerto Rico and The U.S. Virgin Islands

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Ann. Entomol. Soc. Am. 96(3): 181201 (2003)

0013-8746/03/01810201$04.00/0 2003 Entomological Society of America

ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA

Vol. 96, no. 3

SCHEFFRAHN ET AL.: TERMITES OF PUERTO RICO

ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA

Vol. 96, no. 3

SCHEFFRAHN ET AL.: TERMITES OF PUERTO RICO

ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA

Vol. 96, no. 3

SCHEFFRAHN ET AL.: TERMITES OF PUERTO RICO

ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA

Vol. 96, no. 3

SCHEFFRAHN ET AL.: TERMITES OF PUERTO RICO

ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA

Vol. 96, no. 3

SCHEFFRAHN ET AL.: TERMITES OF PUERTO RICO

ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA

Vol. 96, no. 3

SCHEFFRAHN ET AL.: TERMITES OF PUERTO RICO

ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA

Vol. 96, no. 3

SCHEFFRAHN ET AL.: TERMITES OF PUERTO RICO

ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA

Vol. 96, no. 3

SCHEFFRAHN ET AL.: TERMITES OF PUERTO RICO

ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA

Vol. 96, no. 3

SCHEFFRAHN ET AL.: TERMITES OF PUERTO RICO

ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA

Vol. 96, no. 3

SCHEFFRAHN ET AL.: TERMITES OF PUERTO RICO

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