The most diverse and best-preserved paleofauna of the higher termites heretofore known, all found in Miocene amber of the Dominican Republic, is described. The imago of Coptotermes priscus Emerson is redescribed, and the soldier of C. priscus, the first known fossil soldier of this genus, is described. The fauna includes the following 29 new species, all in existing genera, with Krishna and Grimaldi as authors of each: in the Rhinotermitidae, two new species based on imagoes of each—Coptotermes hirsutus and C. paleodominicanus; in the Termitidae, 23 new species based on imagoes—Amitermes lucidus, Anoplotermes bohio, A. cacique, A. carib, A. maboya, A. naboria, A. nitaino, A. quisqueya, A. taino, Atlantitermes antillea, A. caribea, A. magnoculus, Microcerotermes insulanus, M. setosus, Nasutitermes amplioculatus, N. incisus, N. magnocellus, N. medioculatus, N. pilosus, N. seminudus, Subulitermes hispaniola, S. insularis, and Termes primitivus; in the Nasutitermitinae four new species based on nasute soldiers—Caribitermes hispaniola, Nasutitermes rotundicephalus, Parvitermes longinasus, and Velocitermes bulbus. This brings the total termite fauna in Dominican amber to four families, 17 genera, and 39 species, a number that exceeds that of the present-day fauna of Hispaniola. Biogeographical, paleoecological, and phylogenetic implications of the Dominican amber termites are discussed.
Introduction
This is the last and largest report in a series of six papers on the fossil record of the order Isoptera. Prior papers have dealt with Cretaceous taxa (Engel et al., 2007a; Grimaldi et al., 2008), Eocene taxa in Baltic amber (Engel et al., 2007b; Engel, 2008), and Kalotermitidae in Miocene amber from the Dominican Republic (Engel and Krishna, 2007). Here, we describe the most diverse and best-preserved paleofauna, also in Dominican amber, of the so-called “higher” termites in the large family Termitidae, totaling 27 new species in 10 extant genera. In addition, two new species of Rhinotermitidae are described. The total termite fauna in Dominican amber thus consists of four families, 17 genera, and 39 species, which exceeds that of the present fauna of Hispaniola, with three families, 15 genera, and 31 species (table 1). Later in this paper we discuss the biogeographic implications of this difference. The sum of fossil termite taxa from all six recent papers is 47 newly described species and a further 17 revised, which largely redefines what had previously been known (Nel and Paicheler, 1993; Thorne et al., 2000) of the fossil record of this ecologically fundamental group of insects. The purpose of these studies on termite fossils is to construct a phylogeny of the major lineages of termites, both living and extinct.
Table 1
Hispaniolan Termites
The main phylogenetic pattern emerging thus far for termites is that basal families (like Mastotermitidae, Hodotermitidae, and Termopsidae) and stem groups predominated in the Cretaceous. By the Eocene (i.e., Baltic amber), Kalotermitidae and Rhinotermitidae were diverse (Engel et al., 2007b). Interestingly, definitive fossils of the Termitidae—a family that comprises more than 75% of all termite species—did not appear until the Late Oligocene (Martins-Neto and Pesenti, 2006; Nel and Paicheler, 1993). Unfortunately, those records are preserved as compressions and impressions in rocks, which seriously compromises the interpretation of Isoptera, a group where taxonomically important structures are usually minute and subtle. As we report here, by the Miocene ca. 20 mya the Termitidae had evolved into essentially modern tropical groups, indicating that the radiation of the higher termites was relatively recent as well as rapid. Indeed, this radiation appears to be the most significant one among all insects that took place in the mid- to Late Tertiary (Grimaldi and Engel, 2005).
Methods
Dominican amber is Miocene in age, approximately 17 myo (Iturralde-Vinent and MacPhee, 1996), and material in the AMNH collection derived from dealers in the Dominican Republic, as is all commercially available amber from that country. All of the amber comes from mines in the Cordillera Septentrional approximately 10 to 30 km NE of Santiago (Grimaldi, 1995), but the exact source (i.e., the mine) is generally impossible to determine for commercial amber. With the obvious exception of copal, there is little or no variation in composition or age among main amber mines of the Dominican Republic (Grimaldi, 1995; Iturralde-Vinent and MacPhee, 1996). Copal is immediately recognizable: it is lighter in color, softer, crazes readily, and the surface becomes tacky when a drop of solvent is applied. Recent 14C dating found that Dominican copal is merely several hundred years old (Grimaldi, unpubl.). Thus, there is no doubt that all the material we studied is definitively Miocene in age. Unless the provenance of an amber specimen is specifically known and stated, it should be assumed that the locus typicus of all new species is the following: Hispaniola (Dominican Republic), Cordillera Septentrional near Santiago, specific provenance (mine) unknown, El Mamey Formation, Early to mid-Miocene.
Careful study of a termite in amber requires meticulous preparation. Particularly for Rhinotermitidae and Termitidae, identifying imagoes to genus and distinguishing among species depends upon proportions and shapes of the head, eyes, ocelli, clypeus, and pronotum. Details such as the position and structure of the fontanelle and the shape and setation of the head and pronotum require very close views of these structures. Thus, for this study, amber pieces were trimmed and polished for a full frontal view of the head and full dorsal view of the pronotum, where possible. Mandibular dentition is also significant, and fortuitously the labrum of some specimens was clear enough to observe the mandibles beneath. Where possible, a full view of the wings was also prepared so as to document venation. Amber pieces were trimmed with a small water-fed diamond saw (thickness ca. 1 mm), and the faceted surfaces were ground and polished with a water-fed lapidary wheel using emory papers of successively finer grits (first 400, then 600, 2400, and finally 4000). For optimal observation the amber specimen was mounted on a small piece of plasticene or dental wax applied to a glass slide, in the desired orientation, and a small drop of glycerine and a coverslip were applied to the surface of the amber. This last step obscures fine scratches on the surface that reduce visibility of minute structures. Microscope study used fiber optic illumination and either a Zeiss SV8 stereoscope (up to 64×) or a Leitz Wetzlar stereoscope (up to 144×); measurements were made with an ocular reticle on the Leitz stereoscope. Photomicrography used a Nikon D1X digital camera attached to either Infinity® lenses or a Nikon compound microscope, with fiber-optic flash illumination provided by a MicrOptics® system.
Systematics
FAMILY RHINOTERMITIDAE Froggatt, 1897
subfamily coptotermitinae Holmgren, 1910
Genus Coptotermes Wasmann, 1896
The genus Coptotermes is found in all tropical areas of Earth. Many species of this genus have been introduced in various parts of the world, and some have become serious pests, most notoriously Coptotermes formosanus Shiraki. Of the total number of 75 living and two fossil species of the genus hitherto described, only four living (one introduced) and two fossil (one Dominican Republic amber and one Mexican amber) are found in the Neotropical Region (all others are in southeast Asia and Africa). In this paper we are adding two new species and describing the first soldier of the genus Coptotermes in the fossil record.
Coptotermes hirsutus, new species
Figure 1
Photomicrographs of Coptotermes species in Dominican amber. A. Coptotermes hirsutus, n. sp., AMNH DR-PB278. B. Coptotermes hirsutus AMNH DR10-1656. C. Coptotermes paleodomincanus, n. sp., AMNH DR10-1513. D. Coptotermes priscus Emerson, AMNH DR-PB279.
Figure 2
Drawings of head and portions of thoraces of Coptotermes species in Dominican amber, dorsal views. A. Coptotermes hirsutus, n. sp., AMNH DR10-1656 (head), DR10-1698 (pronotum). B. Coptotermes paleodominicanus, n. sp., AMNH DR10-1513 (slightly oblique view).
Figure 3
Wings of Coptotermes species in Dominican amber. A, B. Coptotermes hirsutus, n. sp., AMNH PB278. A. forewing; B. hindwing. C, D. Coptotermes paleodominicus, n. sp., AMNH DR10-1513.
Table 2
Measurements (mm) of imagoes of three species of Coptotermes
Diagnosis
The head, pronotum, and wing scale of the imago of this new species are densely covered with setae of various lengths, in contrast to C. priscus Emerson and C. paleodominicanus, n. sp., in which setae are sparse. It differs from C. priscus in having larger eyes and a wider head and from C. paleodominicanus in having a smaller head and smaller eyes.
Description
Imago: Head dark brown; pronotum, antennae, and legs lighter than head; wing scale brownish, about same color as pronotum; wings brownish yellow, membrane transparent. Head, pronotum, and wing scale densely covered with long bristles, longest about 0.19 mm; costal margin of forewing with several short hairs; wing membrane with fine short hairs. Head semicircular. Eyes medium sized, somewhat oval. Ocelli oval, very close to eyes. Fontanelle dot shaped; situated in middle of head, about 0.49 mm from posterior margin of head (in distorted specimen appearing closer to posterior margin). Postclypeus narrow, faintly arched; width four times length. Antennae with 18–19 articles; third shortest; second longer than fourth. Pronotum slightly narrower than head; front margin broadly concave; posterolateral corners broadly rounded; posterior margin emarginate, faintly angular. Forewing with costal margin heavily sclerotized; radius equally sclerotized, running parallel to costa; median weak, emerging separately from scale, running close to cubitus, with two or three distal subbranches; cubitus as weak as median, with about 11 subbranches to posterior border of wing. Hind wing with costal margin and radius as in forewing; median arising from radius outside of the scale; median and cubitus as in forewing. Tibial spurs 3∶2∶2.
Specimens
Holotype (imago) AMNH No. PB-278. Paratypes (imagoes) AMNH nos. DR8-343, DR10-1263, DR10-1518, DR10-1535, DR10-1561, DR10-1578, DR10-1643, DR10-1648, DR10-1656, PB-277.
Etymology
This species is named after the Latin term hirsutus, “hairy”, and refers to the characteristic dense covering of the head and pronotum.
Coptotermes paleodominicanus, new species
Diagnosis
The imago of this new fossil species has a distinctly longer and wider head, larger eyes and ocelli, and a longer and wider pronotum than the other two fossil species of this genus: C. priscus Emerson and C. hirsutus, n. sp.
Description
Imago: Head rusty brown; postclypeus and labrum brownish, lighter than frons; pronotum brown, lighter than head; legs and antennae light brown; wings brownish yellow. Head with a few short and long bristles; pronotum surface and margins moderately covered with short and long setae, longest about 0.13 mm; sternites, tergites, tibiae, and tarsi covered with short and long bristles; forewing scale with several long bristles, longest about 0.15 mm; costal margin of forewing with several short hairs; wing surface with scattered short bristles. Head much larger than those of C. priscus and C. hirsutus. Eyes large. Ocelli long, very close (0.02 mm) to eye; outer margin indented when viewed dorsally. Fontanelle small, dotlike; about 0.61 mm from posterior margin of head. Postclypeus short, faintly arched; width 3.8 times length. Antennae with 21 articles (only 10 articles visible in holotype); third article shortest; fourth subequal to second. Pronotum narrower than head; anterior margin broadly indented in middle; posterolateral margins broadly rounded; posterior margin emarginate. Forewing with costal margin heavily sclerotized; radius equally sclerotized, running parallel to costa; median weak, emerging separately from scale, running close to cubitus, with two or three distal subbranches; cubitus as weak as median, with about 15 subbranches to posterior border of wing. Hind wing with costal margin and radius as in forewing; median arising from radius outside of the scale; median and cubitus as in forewing. Tibial spurs 3∶2∶2.
Etymology
The species name is a combination of paleo-, “ancient”, and dominicanus, for the Dominican Republic, the general locality where the amber fossil was collected.
Coptotermes priscus Emerson
Figure 4
Coptotermes priscus Emerson in Dominican amber, heads. A, B. Soldier, in ventrolateral view (A) and dorsal view (B, slightly oblique). AMNH DR10-1248. C. Alate, slightly oblique dorsal view, AMNH DR-PB277.
Figure 5
A swarm of Coptotermes priscus Emerson in Dominican amber. This is the most common species of termite in Dominican amber, and is frequently preserved as swarms. AMNH DR-PB279.
Table 3
Measurements (mm) of three soldiers of Coptotermes priscus Emerson
Redescription (based on New Material)
Imago: Head dark brown; pronotum, antennae, and legs lighter than head; wing scale and costal margin brown; wing membrane transparent. Head with many short hairs and several long bristles; pronotum surface moderately covered with short and medium-sized bristles, longer bristles along margin; wing scale with longer bristles than head and surface of pronotum; anterior margin of fore- and hind wing with a row of very short hairs; wing membrane moderately covered with very short hairs. Head circular. Eyes small, rounded, slightly protruded. Ocelli oval, oblique, very close to eyes. Fontanelle dotlike; in a slight depression in middle of head. Postclypeus short, slightly arched; four times as wide as long. Mandible dentition typical of Coptotermes: left mandible with an apical tooth and three marginal teeth; first marginal tooth shorter than both apical and second marginal tooth; second marginal tooth subequal to apical. Antennae with 21 articles; third shortest; second longer than fourth. Pronotum as wide as head; anterior margin broadly concave; lateral margins rounded; posterior margin emarginate. Forewing with costal margin heavily sclerotized; radius equally sclerotized, running parallel to costa; median weak, emerging separately from scale, running close to cubitus, with two or three subbranches in distal one-third; cubitus as weak as median, with 9–11 subbranches to posterior border of wing. Hind wing with costal and radius as in forewing; median arising from radius outside of scale; median and cubitus as in forewing. Tibial spurs 3∶2∶2.
Soldier (fig. 4) (first description of a Coptotermes soldier in the fossil record): Head brown to yellowish brown; pronotum, antennae and legs lighter than head; mandibles brown. Head and pronotum with a few short and medium-sized bristles; area around fontanelle with several short bristles; tergites covered with several medium-sized bristles; sternites with short bristles. Head pear shaped; fontanelle large, typical of Coptotermes, with a large, round opening near base of postclypeus. Labrum subtriangular, longer than broad, with a pointed hyaline tip. Mandibles saber shaped, gently incurved at tips. Postmentum club shaped, approximately twice as long as wide; sides narrowing posteriorly to form a “waist” slightly below middle. Antennae broken; with as least 13 articles; second longer than third. Pronotum flat, narrower than head; anterior margin with a notch in the middle; sides rounded; posterior margin emarginate.
Material Examined
AMNH DR15-280 (soldier and imago); DR10-1248 (soldier); DR14-1179 (soldier); DR10-1200 (imago); PB 280 and PB 281 (three imagoes in each). AMNH No. 11339 (a large, pear-shaped piece, 3″ × 1.5″, containing several imagoes and wings, along with a partial wing of Mastotermes electrodominicus, from La Toca mines).
FAMILY TERMITIDAE Latreille, 1802
subfamily apicotermitinae Grassé and Noirot, 1955
Genus Anoplotermes Müller, 1873
The genus Anoplotermes was first described by F. Müller in 1873, with A. pacificus Müller from Brazil as the type species. After Müller, a large number of species from the Ethiopian, Neotropical, and Nearctic regions were added by various authors (see Snyder, 1949). In a major revision, Sands (1972) removed all the species from the Ethiopian Region previously included in this genus, assigning them to several new genera and leaving only the Neotropical and Nearctic species in Anoplotermes. As soil and humus feeders, the genus Anoplotermes and the related African genera created by Sands are important constituents of the soil fauna in tropical forests (Sands, 1972). Anoplotermes and the African genera of this group are distinguished by having no soldier caste. The workers defend the colony by committing suicide: when encountering an enemy, they rupture their abdomens by muscular contractions, spilling the abdominal contents over the attacker. At present there are 34 living species, along with several undescribed species in the AMNH collection. Of the 34 species, six are reported from the West Indies, of which only one is from Hispaniola (table 1). The Neotropical and Nearctic species presently placed in Anoplotermes need a revision along the lines of that done by Sands (1972), to include studies of the internal characters, such as gut anatomy and Malphigian tubules. Until that is done, we are including the fossil species in the genus Anoplotermes.
Anoplotermes bohio, new species
Figure 6
Photomicrographs of Anoplotermes species in Dominican amber; all are holotypes. A. A. bohio, n. sp., AMNH DR-PB258. B. A. cacique, n. sp., AMNH DR10-1512. C. A. carib, n. sp., AMNH DR10-1541. D. A. maboya, n. sp., AMNH DR10-1567.
Figure 7
Photomicrographs of Anoplotermes species in Dominican amber. A. A. naboria, n. sp., AMNH DR10-1537 (holotype). B. A. nitaino, n. sp., AMNH DR-PB260 (holotype). C. A. quisqueya, n. sp., AMNH DR-PB255 (paratype). D. A. taino, n. sp., AMNH DR-PB257 (holotype).
Figure 8
Heads and pronota of Anoplotermes spp. in Dominican amber, dorsal views. A. A. bohio, n. sp. (head slightly oblique), AMNH DR-PB258. B. A. cacique, n. sp., AMNH DR10-1512.
Table 4
Measurements (mm) of imagoes of eight new species of Anoplotermes
Diagnosis
Anoplotermes bohio differs from all of the other fossil species described here in having its head and pronotum densely covered with long setae. It is closest to A. carib and A. maboya, n. spp., in head size, but differs from A. maboya in having a wider head, slightly larger eyes, and a larger pronotum. It differs from A. carib in its narrower head, smaller eyes, and larger pronotum.
Description
Dealate imago: Head, pronotum, and wing scale dark brown; antennae yellowish. Head, pronotum, and wing scale densely covered with long, interspersed with short, setae. Head longer than wide. Eyes small, nearly oval. Ocelli oval, not touching eyes (0.05 mm from eye). Fontanelle not clearly visible, but apparently oval. Postclypeus arched; length equal to half its width (length to width index 0.50). Antennae with 15 articles; third slightly shorter than fourth; second subequal to fourth. Pronotum as wide as head; anterior margin raised in middle, forming two mild concavities on either side of median; posterolateral corners broadly rounded; posterior margin faintly emarginate.
Etymology
This new species name is directly from the Taino word for Hispaniola. The Taino were a Greater Antillean tribe of the Arawaka people.
Anoplotermes cacique, new species
Diagnosis
This species is closest to Anoplotermes naboria, and A. nitaino, n. spp., in its head length. It differs from A. naboria in its narrower head, smaller eyes, larger pronotum, and longer and wider forewing. It differs from A. nitaino in its slightly wider head, larger eyes, and larger pronotum.
Description
Imago: Head and pronotum metallic brown appearing (coloration obscure due to preservation). Head and pronotum pilosity also obscure; anterior margin of forewing with a row of short setae; wing membrane with dotlike punctations. Head longer than wide. Eyes roundish, moderately sized, bulging. Ocelli not touching eye (about 0.03 mm from eye). Fontanelle not visible. Postclypeus arched, divided medially by a fine line; length slightly less than half its width (length to width index 0.47). Antennae with 15 articles; third very short; second subequal to fourth. Pronotum narrower than head; anterior margin very faintly emarginate; posterolateral corners widely rounded; posterior margin with a faint median indentation.
Etymology
This new species is given the Taino name for chief.
Anoplotermes carib, new species
Figure 9
Heads and pronotum of Anoplotermes spp. in Dominican amber; both are holotypes. A. A. carib, n. sp., dorsal view. AMNH DR10-1541. B. A. maboya, n. sp., dorsolateral view of head (pronotum not fully visible), with detail of mandibles and lacinia. AMNH DR10-1567.
Figure 10
Heads and pronota of Anoplotermes spp. in Dominican amber, dorsal views. A. A. naboria, n. sp., AMNH DR10-1537. B. A. nitaino, n. sp., AMNH DR-PB260.
Diagnosis
This new species is closest to A. bohio and A. maboya, n. spp., in its head length, but differs from the latter in having a wider head, shorter and narrower forewings, and larger eyes. It differs from A. bohio in having shorter setae, slightly larger eyes, and a narrower pronotum.
Description
Imago: Head brown; pronotum, forewing scale brown, lighter than head; antennae brownish yellow. Head and pronotum densely covered with short setae, interspersed with a few long setae; forewing scale with several long setae; anterior margin of forewing with a row of short setae; wing membrane with dotlike punctations and a few short, scattered setae. Head slightly longer than wide. Eyes small, nearly round, faintly protruding. Ocelli oval, not touching eyes (about 0.03 mm from eyes). Fontanelle oval; width 0.03 mm; situated about 0.31 mm from posterior margin of head. Postclypeus arched; length equal to half its width (length to width index 0.50); medial line not clearly visible. Antennae with 15 articles; third shortest; fourth subequal to second. Pronotum narrower than head; anterior margin moderately concave; posterolateral corners broadly rounded; posterior margin faintly indented medially. Median vein branched into two apically.
Etymology
This new species is given the name of the fierce, indigenous peoples of South America who invaded the Tainos.
Anoplotermes maboya, new species
Diagnosis
Anoplotermes maboya is closest to A. bohio and A. carib, n. spp., in its head length and size of the eyes. It differs from A. bohio in having smaller eyes and sparse setae on its head and pronotum. It differs from A. carib in its longer and wider forewing and smaller eyes.
Description
Imago: Head and pronotum brown; antennae yellowish brown; wings brownish. Head and pronotum with a few short setae; forewing with a row of short setae; wing membrane with dotlike punctations. Head poorly preserved, compressed laterally; longer than wide. Eyes small, round, protruding slightly. Ocelli oval, small, not touching eyes (0.02 mm from eye). Fontanelle small, oval; width 0.03 mm; situated about 0.31 mm from posterior margin of head; medial line faintly visible. Postclypeus arched; length equal to half its width (length to width index 0.50). Mandibles visible, characteristic of Anoplotermes. Antennae with 15 articles; third shortest; fourth subequal to second. Pronotum not clearly visible, apparently narrower than head; anterior margin apparently angular.
Etymology
This new species is given the Taino name of the nocturnal god and destroyer of crops.
Anoplotermes naboria, new species
Diagnosis
Anoplotermes naboria is closest to A. nitaino and A. quisqueya, n. spp., in its pilosity and its head length. It differs from A. quisqueya in having a narrower head and fontanelle, larger eyes, a smaller pronotum, and a shorter and narrower forewing. It differs from A. nitaino in having a wider head, larger eyes, and narrower forewing.
Description
Imago: Head and pronotum dark brown; postclypeus brown, slightly lighter than head; antennae brown. Head densely covered with very short setae; pronotum densely covered with short setae, like head, with a few long setae along margins; anterior margin of forewing with a row of short setae; wing membrane with dotlike punctations. Head longer than wide. Eyes of moderate size, roundish, bulging. Ocelli oval, not touching eye (0.03 mm from eye). Fontanelle oval; width 0.03 mm; situated about 0.26 mm from posterior margin. Postclypeus flat; length slightly less than half its width (length to width index 0.47); medial line very faint. Antennae with 15 articles; third shortest; fourth subequal to second. Pronotum narrower than head; anterior margin deeply angular; posterolateral corners broadly rounded; posterior margin slightly indented medially. Forewing with median unbranched; Cu with 10 branches.
Etymology
This new species is given the name for worker in the Taino language.
Anoplotermes nitaino, new species
Diagnosis
Anoplotermes nitaino is distinguished from all the other fossil species by its small, round, dotlike fontanelle (vs. oval or teardrop shaped in all the other species).
Description
Imago: Head and pronotum chestnut brown; antennae brownish. Head and pronotum densely covered with short setae, bases of setae appearing as small dots; anterior margin of forewing with a row of setae; wing membrane with dotlike punctations. Head longer than wide. Eyes nearly round, bulging. Ocelli small, oval, not touching eyes (0.03 mm from eye). Fontanelle small, round, dotlike; situated about 0.33 mm from posterior margin of head. Postclypeus flat; length slightly less than half its width (length to width index 0.47). Mandibles partially visible, suggestive of Anoplotermes (see Ahmad, 1950: fig. 12). Antennae with 15 articles; third shortest; second subequal to fourth. Pronotum narrower than head; anterior margin concave; posterior margin broadly emarginate.
Etymology
This new species is given the Taino name for nobleman or subchief.
Anoplotermes quisqueya, new species
Diagnosis
This species differs from all the other fossil species of this genus in having a large, teardrop-shaped fontanelle (width 0.05 mm vs. 0.03 mm in all other species).
Description
Imago: Head, pronotum, and wing scale brown; antennae brown, lighter than head. Head and pronotum densely covered with short setae, interspersed with a few long setae; anterior margin of forewing with a row of setae; wing membrane with dotlike punctations. Head longer than wide. Eyes medium sized, nearly oval, bulging. Ocelli oval, not touching eyes (0.06 mm from eye). Fontanelle large, teardrop shaped; width 0.05 mm; situated about 0.36 mm from posterior margin of head. Postclypeus very slightly arched; length slightly less than half its width (length to width index 0.48); divided by a medial line. Antennae with 15 articles; third shortest; second subequal to fourth. Pronotum narrower than head; anterior margin widely concave; posterolateral corners widely rounded; posterior margin almost straight. Partial forewing showing median with four branches.
Etymology
This new species is given the Taino name for Hispaniola, also meaning “mother of the earth.”
Anoplotermes taino, new species
Diagnosis
Anoplotermes taino is close to A. quisqueya, n. sp., in its fontanelle size, but differs from it in having small eyes, a narrower head, and a narrower pronotum.
Description
Imago: Head and pronotum dark brown; antennae brown; wings brownish. Head covered with numerous short setae interspersed with moderate-sized setae; tergites, sternites, and legs with short setae; anterior margin of forewing with a row of short setae; wing membrane with dotlike punctations. Head longer than wide. Eyes small, round, bulging. Ocelli oval, not touching eyes (about 0.03 mm from eye). Fontanelle large, pear shaped; situated posteriorly below level of eyes, close (0.10 mm) to posterior margin of head. Postclypeus arched; length slightly less than half its width (length to width index 0.45). Antennae with 16 articles (apparently): third very short; second subequal to fourth. Pronotum narrower than head; anterior margin roundish (somewhat unclear); posterolateral corners widely rounded; posterior margin with a faint median indentation. Forewing with median vein branched apically; cubitus with 11 branches.
Etymology
This new species is given the name of the Taino, the indigenous people of the Greater Antilles.
Key to the Species of Anoplotermes in Dominican Amber
1. Fontanelle dotlike or not visible2
— Fontanelle teardrop shaped, oval, or slitlike3
2. Eye diameter 0.31 mm; pronotal width 0.81 mm; pronotum with posterolateral corners widely rounded (fig. 8B)A. cacique
— Eye diameter 0.26 mm; pronotal width 0.77 mm; pronotum with posterolateral corners narrowly rounded (fig. 10B)A. nitaino
3. Head densely covered primarily with long setae (fig. 8A)A. bohio
— Head densely covered with short setae or very few setae4
4. Fontanelle small, width 0.03 mm5
— Fontanelle large, width 0.05 mm7
5. Eye diameter 0.21 mm; head with few setae; forewing length 7.00 mm (fig. 9B)A. maboya
— Eye diameter 0.26 to 0.33 mm; head densely covered with setae; forewing length 5.60 to 6.30 mm6
6. Eye diameter 0.26 mm; pronotal width 0.69 mm; forewing length 5.60 mm forewing width 1.58 mm (fig. 9A)A. carib
— Eye diameter 0.33 mm; pronotal width 0.77 mm forewing length 6.30 mm; forewing width 1.50 mm (fig. 10A)A. naboria
7. Eye diameter 0.20 mm; head width 0.79 mm; pronotal width 0.69 mm (fig. 11B)A. taino
— Eye diameter 0.27 mm; head width 1.02 mm; pronotal width 0.87 mm (fig. 11A)A. quisqueya
subfamily termitinae Latreille, 1802
Genus Amitermes Silvestri, 1901
The genus Amitermes is a mostly circumtropical genus, with 16 living species reported from the Neotropical Region, of which only one is from the West Indies (Cuba) (table 1). Amitermes species generally feed on wood and are found in a variety of habitats, from desert to savannah to rainforest. Some nest underground and others are mound builders, most notably Amitermes excellens (Silvestri 1923) from Brazil, reported to build mounds up to 15 feet (5 m) high. No living species has been reported from Hispaniola. Amitermes lucidus, n. sp., is the first record of this genus, living or fossil, from Hispaniola and the first fossil record of this genus from any deposit.
Amitermes lucidus, new species
Figure 13
Head, pronotum (dorsal view) and detail of manidibles (as preserved in Dominican amber) of Amitermes lucidus, n. sp., AMNH DR10-1563.
Table 5
Measurements (mm) of imago of Amitermes lucidus, new species
Diagnosis
The imago of this new fossil species has a narrower head and pronotum, smaller eyes, and a larger fontanelle than the living species A. beaumonti Banks from Cuba, the only species of Amitermes heretofore known from the West Indies.
Description
Imago: Head light yellow; pronotum yellowish, darker than head; antennae yellowish; wings brownish. Head, postclypeus, and pronotum densely covered with short bristles interspersed with several long ones; wing scale with several short bristles; wing membrane with a few hairs and pigmented punctations. Head longer than wide. Eyes small, nearly oval, protruding. Ocelli small, oval, not touching eyes (0.05 mm from eye). Fontanelle large, teardrop shaped; 0.39 mm from posterior margin of head. Postclypeus arched; width slightly less than half its length (length to width index 0.53), with a faint median line. Left mandible faintly visible; dentition characteristic of the genus Amitermes (see Ahmad, 1950: fig. 12). Antennae with 15 articles; third very short; second subequal to fourth. Pronotum narrower than head; anterior margin slightly raised medially; posterolateral corners broadly rounded; posterior margin distinctly emarginate. Forewing with costa thickly sclerotized; radius equally sclerotized, running parallel to costa; median weak, emerging separately from scale, running close to cubitus, with six branches, joining apical margin; cubitus weak, approximately 8 branches to lower margin.
Etymology
This new species is named after the Latin lucidus, “light”, referring to the coloration of the head.
Genus Microcerotermes Silvestri, 1901
Microcerotermes is a genus most abundant in the tropical regions of the world, with seven living species from the Neotropical Region, two of which are reported from the West Indies, of which one is from Hispaniola (Haiti only) (table 1). Most species are wood feeders and build hard carton nests of fecal matter. The two species described here, Microcerotermes insulans and M. setosus, are, with the exception of a dubious species in gum copal, the first record of this genus, living or fossil, from the Dominican Republic and the first fossil record of this genus.
Microcerotermes insulanus, new species
Figures 14–Figure 1516; table 6
Figure 14
Photomicrographs of Microcerotermes spp. in Dominican amber. A. M. insulanus, n. sp., AMNH DR10-1526. B. M. setosus, n. sp., AMNH DR10-1559.
Figure 15
Heads and pronota of Microcerotermes spp. in Dominican amber (dorsal views). A. M. setosus, n. sp., AMNH DR10-1553. B. M. insulanus, n. sp., AMNH DR10-1526.
Figure 16
Wings of Microcerotermes spp. in Dominican amber. A. M. insulanus, n. sp., forewing, AMNH DR10-1526. B, C. Fore- and hindwing, respectively, of M. setosus, n. sp., AMNH DR10-1559.
Table 6
Measurements (mm) of imagoes of two species of Microcerotermes
Diagnosis
The imago of this species differs from M. setosus, n. sp., in having smaller eyes, shorter and narrower forewings, a narrower pronotum, and its head and pronotum less densely covered with setae.
Description
Imago: Head and pronotum dark brown; antennae lighter brown; wings light brown. Head and pronotum covered with a moderate number of short setae; forewing scale with numerous long setae; wing membrane with a few short setae and dotlike punctations (microtrichia). Head with sides subparallel below eyes. Eyes small, nearly round. Ocelli barely visible (appear oval), approximately 0.05 mm from eyes. Fontanelle not clearly visible due to whitish film over head (appears dotlike). Postclypeus faintly arched; length half its width (length to width index 0.50). Antennae with 15 articles; third shortest; second subequal to fourth. Pronotum distorted in preservation, narrower than head; anterior margin concave; posterolateral corners broadly rounded. Forewing with median vein unbranched.
Etymology
This new species is named after the Latin insulanus, “islander”.
Microcerotermes setosus, new species
Diagnosis
The imago of this species differs from M. insulanus, n. sp., in having larger eyes, longer and wider forewings, and its head and pronotum densely covered with setae.
Description
Imago: Head and pronotum blackish brown; antennae brownish, lighter than head; forewing scale yellowish brown. Head with a dense mat of short setae, with many long setae interspersed; pronotum densely covered with short setae and longer setae along its margin; wing scale, tergites, sternites heavily covered with setae; anterior margin of forewing with a row of short setae; wing membrane with dotlike punctations (microtrichia). Head oblong, with sides subparallel below eyes. Eyes nearly round. Ocelli oval, very close (0.03 mm) to eyes. Fontanelle not visible due to debris on surface of head (in one specimen appears dotlike). Postclypeus apparently flat, perhaps distorted in preservation; length more than half its width (length to width index 0.58). Antennae with 15 articles; third shortest; second subequal to fourth. Pronotum narrower than head; anterior margin concave; anterolateral corners narrow; posterior margin faintly emarginate. Forewing with median vein with five branches.
Etymology
This new species name is from the Latin setosus, for its being densely covered with setae.
Genus Termes Linnaeus, 1758
The genus Termes is a circumtropical genus with a wide range of nesting habits. Of the 28 living species, nine have been recorded from the Neotropical Region, with four from the West Indies, only one of which is from Hispaniola (table 1). Termes fatalis, a living species from Surinam, was the first named termite, described by Linnaeus in 1758. One impression fossil species is known from the Oligocene of France, diagnosed on the basis of wing venation. The species described here, Termes primitivus, is the first amber fossil of this genus to be described and the first Termes fossil to be reported from the Dominican Republic.
Termes primitivus, new species
Figure 17
Photomicrographs of A. Termes primitivus, n. sp., AMNH DR 10-1521 (paratype). B. Subulitermes insularis, n. sp., in Dominican amber, AMNH DR8-332 (holotype).
Table 7
Measurements (mm) of imago of Termes primitivus, new species
Diagnosis
The imago of this new fossil species has a narrower head and pronotum and larger eyes than the living species T. hispaniolae, the only species of Termes heretofore reported from Hispaniola.
Description
Imago: Head and pronotum dark brown; legs brown; wing scale brown, lighter than head. Head, postclypeus, and pronotum covered with a dense mat of short bristles and a few long ones; wing scale with several short bristles; wing membrane with a few hairs and pigmented punctations; femur and coxa densely covered with short hairs. Head longer than wide; posterior margin broadly concave. Eyes medium sized, circular. Ocelli oval, arched, very close to eyes. Fontanelle ovalish, with a faint slit in front. Postclypeus arched, with a faint median line; length about half its width (length to width index 0.49). Left mandible faintly visible; apical tooth long, about 0.13 mm from anterior margin of first marginal tooth, characteristic of the genus Termes. Antennae with 15 articles; third shortest; second subequal to fourth. Pronotum flat, narrower than head; anterior margin concave, lateral sides broadly rounded; posterior margin angular, emarginate. Forewing suture faintly arched. Forewing with costa thickly sclerotized; radius equally sclerotized, running parallel to costa; median weak, emerging separately from scale, running close to cubitus, unbranched, joining apical margin; cubitus weak; approximately 14–16 branches to lower margin. Hind wing, costa, and radius similar to forewing; median weak, arising from radius, outside of scale. Tibial spurs 3∶2∶2.
Etymology
This new species is named after the Latin primitivus, “first”, earliest of its kind.
subfamily nasutitermitinae Hare, 1937
Genus Atlantitermes Fontes, 1979
The genus Atlantitermes was first described by Fontes in 1979 for three new species and later expanded (Fontes, 1982) to include five species previously included in the genus Subulitermes. As this exclusively Neotropical genus is comprised today, only one living species of the eight is reported from the West Indies, which is actually from the subcontinental island of Trinidad (table 1). Like Subulitermes, the Atlantitermes species are tropical soil dwellers, feeding on soil rich in humus. No fossil species of this genus has been hitherto reported. The three new fossil species described here are the first record of this genus, living or fossil, from Hispaniola.
Atlantitermes antillea, new species
Figure 19
Photomicrographs of Atlantitermes spp. in Dominican amber; all are holotypes. A. A. antillea, n. sp., AMNH DR10-1543. B. A. caribea, n. sp., AMNH DR10-1555. C. A. magnoculus, n. sp., AMNH DR10-1259.
Figure 20
Heads and pronota of Atlantitermes spp. in Dominican amber. A. A. antillea, n. sp. (reconstructed), AMNH DR10-1543. B. Detail of mandibles of A. antillea, n. sp., as observed through labrum, AMNH DR10-1543. C. A. caribea, n. sp., AMNH DR10-1555 (slightly oblique view). D. A. magnoculus, n. sp., AMNH DR10-1259.
Table 8
Measurements (mm) of imagoes of three new species of Atlantitermes
Diagnosis
Atlantitermes antillea is close to A. caribea, n. sp., but differs from it in having a wider head and pronotum, smaller eyes and ocelli, and a shorter fontanelle.
Description
Imago: Head and pronotum chestnut brown; postclypeus brown, lighter than head; antennae brown; wings brownish, membrane clear. Head and pronotum sparsely covered with short setae; anterior margin of forewing with a row of short setae; wing membrane with dotlike punctations. Head longer than wide. Eyes medium-sized, round, bulging. Ocelli large, oval, very close (0.01 mm) to eyes. Fontanelle ovalish, extended as a faint line in front; located between eyes, 0.28 mm from posterior margin of head. Postclypeus arched; length less than half its width (length to width index 0.42); with a median line. Antennae with 15 articles; third shortest; second subequal to fourth. Pronotum narrower than head; anterior margin almost straight; lateral margains angular posteriorly; posterior margin almost straight.
Etymology
This new species name is derived from Antilles, another name for the Caribbean Islands.
Atlantitermes caribea, new species
Diagnosis
Atlantitermes caribea is close to A. antillea, n. sp., but differs from it in having a narrower head and pronotum, larger eyes and ocelli, and a longer fontanelle.
Description
Imago: Head and pronotum brown; postclypeus brown, lighter than head; antennae yellowish brown; wings brownish, membrane clear. Head densely covered with short setae; pronotum densely covered with short setae, longer setae along margins; anterior margin of forewing with a row of short setae; wing membrane with dotlike punctations. Head longer than wide. Eyes medium-sized, roundish, bulging. Ocelli large, oval, long, almost touching (0.03 mm) eyes. Fontanelle long, ovalish, located between eyes, 0.28 mm from posterior margin of head. Postclypeus arched, with a faint median line; length less than half its width (length to width index 0.43). Antennae with 15 articles; third shortest; second subequal to fourth. Pronotum narrower than head; anterior margin concave; posterolateral corners broadly rounded; posterior margin distinctly emarginate.
Etymology
This new species name is from the stem of Caribbean, in reference to the extralimital occurrence of the fossil species in this area.
Atlantitermes magnoculus, new species
Diagnosis
Atlantitermes magnoculus differs from all other fossil species of Atlantitermes in being much larger in all respects, notably the head, pronotum, and eyes. In eye size, fontanelle shape, and overall appearance it is close to the living species A. osborni Emerson, from Guyana.
Description
Imago: Head brown; pronotum brownish; postclypeus brownish, lighter than head; wings brownish, membrane clear. Head densely covered with short setae; pronotum densely covered with short setae, longer setae along margins; anterior margin of forewing with a row of short setae; wing membrane with dotlike punctations. Head (partially damaged in preservation) longer than wide. Eyes very large, oval, bulging. Ocelli large, oval, touching eyes. Fontanelle damaged, apparently narrow, slitlike, forked at tip; located between eyes, about 0.38 mm from posterior margin of head. Postclypeus arched; length less than half its width (length to width index 0.35); median line not visible. Left mandible dentition visible, Atlantitermes-type (Fontes, 1979). Antennae with 15 articles; second article equal to third; fourth subequal to second and third. Pronotum narrower than head; anterior margin faintly concave; lateral margins angular posteriorly; posterior margin faintly emarginate.
Etymology
This new species name is a combination of the root of the Latin magnus, “large”, and oculus, “eye”.
Genus Nasutitermes Dudley, 1890
Nasutitermes is the largest genus of the order Isoptera, with a circumtropical distribution. Of the total of 262 living species and subspecies, 71 are Neotropical, 10 of which are found in the West Indies, with three species from Hispaniola (table 1). The species of this genus are found in a variety of ecological habitats, with a great variation in feeding habits and nest construction. Many construct large terrestrial mounds, others build underground or arboreal nests. Their preferred diet is wood, and many species build runways of soil and carton to get to the wood source. Until now, one fossil species from Dominican amber, based on the soldier caste, and one fossil species from Mexican amber, based on the imago caste, have been known. Seven new fossil species, six based on the imago caste and one on the soldier, are described here.
Nasutitermes amplioculatus, new species
Figure 21
Photomicrographs of imagoes of Nasutitermes spp. in Dominican amber; all are holotypes. A. N. amplioculatus, n. sp., AMNH DR10-1680. B. N. incisus, n. sp., AMNH DR-PB263. C. N. medioculatus, n. sp., AMNH DR14-643.
Figure 22
Photomicrographs of imagoes of Nasutitermes spp. in Dominican amber; both are holotypes. A. N. pilosus, n. sp., AMNH DR10-1627. B. N. seminudus, n. sp., AMNH DR10-1574.
Figure 23
Nasutitermes imagoes in Dominican amber, heads and pronota, dorsal views. A. N. amplioculatus, n. sp., AMNH DR10-1680. B. N. incisus, n. sp., PB263 (head slightly distorted).
Table 9
Measurements (mm) of imagoes of six new species of Nasutitermes
Diagnosis
Nasutitermes amplioculatus resembles N. magnocellus and N. incisus, n. spp., in having large eyes. It differs from both in having slightly smaller eyes and a small, dotlike fontanelle (vs. oblong in N. magnocellus and large and round in N. incisus).
Description
Imago: Head and pronotum dark brown; postclypeus lighter than head; antennae brownish. Head densely covered with short setae; pronotum with several short setae, longer setae along margins; tergites and sternites densely covered with long setae; forewing scale with several long setae; anterior margin of forewing with a row of short setae; wing membrane with dotlike punctations. Head longer than wide. Eyes large, oval, bulging laterally. Ocelli oval viewed dorsally, almost touching eyes. Fontanelle very small; located on a level with the dorsal margin of eyes, 0.28 mm from posterior margin of head. Postclypeus arched; length less than half its width (length to width index 0.41). Antennae broken, with 15 articles visible; third shorter than second and fourth; second subequal to fourth. Pronotum narrower than head; anterior margin slightly concave; posterolateral margins widely rounded; posterior margin with a faint medial indentation.
Etymology
This new species name is a combination of the Latin prefix ampli- “large”, and oculatus, “of eyes”.
Nasutitermes incisus, new species
Diagnosis
Nasutitermes incisus differs from all of the other Nasutitermes fossil species described here in having very large eyes and the anterior margin of the pronotum deeply incised.
Description
Imago: Head and pronotum dark brown; wings light brown, membrane clear. Head and pronotum densely covered with long setae; legs, tergites, and sternites densely covered with setae; wing scales with long setae; anterior margin of forewing with a row of short setae; wing membrane with dotlike punctations. Head oblong. Eyes large, ovalish, bulging. Ocelli from side view round, touching eyes. Fontanelle round; diameter 0.08 mm; located between eyes, about 0.56 mm from posterior margin of head. Postclypeus arched; length less than half its width (length to width index 0.37). Antennae with 15 articles visible; second longer than third or fourth; third subequal to fourth. Pronotum narrower than head; anterior margin deeply incised; posterior margin with a deep medial indentation.
Etymology
This new species name is from the Latin incisus, “incised”, referring to the deeply incised pronotum.
Nasutitermes magnocellus, new species
Diagnosis
Nasutitermes magnocellus, n. sp., differs from all the other fossil species of this genus in having a very large ocellus that is also forked at the anterior end.
Description
Imago: Head brown; pronotum and postclypeus brown, lighter than head; antennae brownish; wings yellowish. Head densely covered with short setae, interspersed with a few longer setae; pronotum with several short setae, longer setae along margins; sternites, tergites, and legs densely covered with setae; forewing scale with several long setae; anterior margin of forewing with a row of short setae; wing membrane with dotlike punctations. Head longer than wide; right side not visible due to poor preservation. Eyes large, roundish, bulging laterally. Ocelli large, oval, almost touching eyes (about 0.02 mm). Fontanelle large, oblong, faintly forked at tip; located between eyes, about 0.56 mm from posterior margin of head. Postclypeus faintly arched; length less than half its width (length to width index 0.33). Antennae with 15 articles; articles appear elongated, due to poor preservation; second, third, and fourth appear equal in size. Pronotum narrower than head; anterior margin widely concave; posterolateral corners broadly rounded; posterior margin with a deep medial indentation. Hind tibia long. Forewing (partial) 10.00 mm long.
Etymology
This new species name is a combination of the root of Latin magnus, “large”, and ocellus.
Nasutitermes medioculatus, new species
Diagnosis
Nasutitermes medioculatus is close to N. pilosus and N. seminudus, n. spp., in the size of its eyes. It differs from N. pilosus in having a larger fontanelle, larger eyes, a wider head, and longer ocelli. It differs from N. seminudus in having a larger fontanelle, larger eyes, and its head and pronotum more densely covered with setae.
Description
Imago: Head and pronotum chestnut brown; postclypeus lighter than head; antennae brownish; legs yellowish. Head moderately covered with short setae, along with a few longer setae; pronotum with several short setae, longer setae along margins; forewing scale with several long setae; anterior margin of forewing with a row of short setae; wing membrane with dotlike punctations. Head oval. Eyes roundish, bulging laterally. Ocelli oval, close to eyes (about 0.03 mm). Fontanelle long, ovalish, forked at tip; located between eyes, about 0.36 mm from posterior margin of head. Postclypeus faintly arched; length less than half its width (length to width index 0.39). Antennae broken, with only five articles visible; third very short; fourth subequal to second. Pronotum narrower than head; anterior margin widely concave; posterolateral corners broadly rounded; posterior margin with a deep medial indentation.
Etymology
This new species name is a combination of the Latin prefix medi-, “medium”, and oculatus, “of eyes”.
Nasutitermes pilosus, new species
Figures 22, 25, 26; table 9
Diagnosis
Nasutitermes pilosus resembles N. seminudus, n. sp., but differs from it in having smaller eyes, a differently shaped fontanelle (round vs. oval in N. seminudus), a pronotum with its posterior margin broadly emarginate (vs. deeply notched medially in N. seminudus), and its head and pronotum densely covered with setae.
Description
Imago: Head brown; postclypeus lighter than head; pronotum, antennae, and legs yellowish. Head and pronotum densely covered with a fine mat of short setae, along with several longer setae; forewing scale with a few long setae; anterior margin of forewing with a row of short setae; wing membrane with dotlike punctations. Head oblong. Eyes small, nearly round. Ocelli small, oval, close (about 0.03 mm) to eyes. Fontanelle small, circular, located between eyes, about 0.36 mm from posterior margin of head. Postclypeus nearly flat; length less than half its width (length to width index 0.40). Antennae with 15 articles; third very short; fourth subequal to second. Pronotum narrower than head; anterior margin widely concave; posterior margin broadly emarginate. Forewing with median vein branched (four branches); cubitus with 6–7 branches.
Etymology
This new species is named from the Latin pilosus, “hairy”.
Nasutitermes seminudus, new species
Figures 22, 25, 26; table 9
Diagnosis
Nasutitermes seminudus resembles N. pilosus, n. sp., but differs from it in having larger eyes, a differently shaped fontanelle (oval, vs. round in N. pilosus), a pronotum deeply notched medially, and its head and pronotum with few setae.
Description
Imago: Head and pronotum brown; postclypeus lighter than head; antennae yellowish; forewing scale brownish. Head with very few setae; pronotum with a few short setae on surface, a few long setae along margins; forewing scale with a few moderately sized setae; anterior margin of forewing with a row of short setae; wing membrane with punctations, no setae visible. Head nearly oval. Eyes small, roundish. Ocelli ovalish, not touching eyes (0.03 mm from eyes). Fontanelle slitlike; located about 0.38 mm from posterior margin of head. Postclypeus arched; length less than half its width (length to width index 0.34). Antennae with 15 articles; third shortest; fourth subequal to second. Pronotum narrower than head; anterior margin broadly concave; posterior margin with a deep medial indentation. Forewing with median branched apically. Cubitus with 10 branches.
Etymology
This new species is a combination of the Latin prefix semi-, “partially”, and nudus, “naked, bare”.
Key to the Species of Nasutitermes Imagoes in Dominican Amber
1. Eyes large; diameter 0.41–0.56 mm2
— Eyes medium-sized or small; diameter 0.26–0.37 mm4
2. Ocelli small; length 0.13–0.15 mm3
— Ocelli large; length 0.23 mm; eye diameter 0.51 mm; fontanelle oblong, forked on front (fig. 24A)N. magnocellus
3. Eye diameter 0.56 mm; fontanelle large, round; pronotal width 1.10 mm; anterior margin of pronotum deeply incised (fig. 23B)N. incisus
— Eye diameter 0.41 mm; fontanelle small, dotlike; pronotal width 0.79 mm; anterior margin of pronotum not deeply incised (fig. 23A)N. amplioculatus
4. Head and pronotum densely covered with setae5
— Head and pronotum sparsely covered with setae (fig. 25B)N. seminudus
5. Eye diameter 0.26 mm; fontanelle small; head width 0.87 mm; ocellus length 0.09 mm (fig. 25A)N. pilosus
— Eye diameter 0.31 mm; fontanelle large; head width 0.94 mm; ocellus length 0.13 mm (fig. 24B)N. medioculatus
Genus Subulitermes Holmgren, 1910
The genus Subulitermes, treated today in a more restricted sense than that of Snyder (1949) and Emerson (1955), presently consists of six living species from the Neotropical Region. The distribution is very similar to that of Atlantitermes, with only one species reported from the West Indies, also from the subcontinental island of Trinidad (table 1). Species of Subulitermes are tropical soil dwellers, feeding on soil rich in humus. No fossil species of this genus has been hitherto reported. The two new fossil species described here are the first record of this genus, living or fossil, from Hispaniola.
Subulitermes insularis, new species
Figure 27
Heads and pronota (dorsal views) with details of visible portions of mandibles of Subulitermes spp. in Dominican amber; both are holotypes. A. S. hispaniola, n. sp., AMNH DR8-341. B. S. insularis, n. sp., AMNH DR8-332.
Table 10
Measurements (mm) of imagoes of two new species of Subulitermes
Diagnosis
Subulitermes insularis differs from S. hispaniola, n. sp., in having smaller eyes, a wider head, a wider pronotum with angular sides and posterior margin almost straight.
Description
Imago: Head and pronotum brown; postclypeus lighter than head; wings light brown, membrane clear. Head densely covered with short setae; pronotum densely covered with short setae, longer setae along margins; anterior margin of forewing with a row of short setae; wing membrane with dotlike punctations. Head longer than wide. Eyes small, round, bulging. Ocelli ovalish, very close (0.03 mm) to eyes. Fontanelle not clearly visible, apparently narrow and oblong; located between eyes, about 0.23 mm from posterior margin of head. Postclypeus arched; length less than half its width (length to width index 0.37). Mandible dentition visible, Subulitermes-type (see Ahmad, 1950: fig. 11). Antennae with 15 articles; second article longer than third or fourth; third subequal to fourth. Pronotum narrower than head; anterior margin straight; lateral margins narrowing posteriorly at an angle; posterior margin faintly emarginate.
Etymology
This new species name is based on insular, referring to the island origin.
Subulitermes hispaniola, new species
Diagnosis
Subulitermes hispaniola differs from S. insularis, n. sp., in having larger eyes, a narrower head, narrower forewings, and a narrower pronotum, with its posterolateral corners more broadly rounded and its posterior margin with a medial indentation.
Description
Imago: Head and pronotum reddish brown; postclypeus light brown, lighter than head; antennae brownish; wings light brown, membrane clear. Head densely covered with short setae; pronotum densely covered with short setae, longer setae along margins; anterior margin of forewing with a row of short setae; wing membrane with dotlike punctations. Head longer than wide. Eyes small, ovalish, bulging. Ocelli oval, 0.05 mm from eyes. Fontanelle oval; about 0.31 mm from posterior margin of head. Postclypeus arched; length less than half its width (length to width index 0.39). Mandible dentition visible, Subulitermes type (Ahmad, 1950: fig. 11). Antennae with 15 articles; second longer than third or fourth; third subequal to fourth. Pronotum narrower than head; anterior margin undulating, with a faint median projection; posterolateral corners rounded; posterior margin with a distinct medial indentation.
Etymology
This new species name is derived directly from the island of origin.
nasutitermitinae soldiers
Four new species of nasutitermitines are described below based on the soldiers, which, with two other Dominican amber species described previously (Nasutitermes electronasutus Krishna and Constrictotermes electroconstrictus Krishna), brings to six the total of nasutitermitine species in Dominican amber known from soldiers. The two species of Nasutitermes described on the basis of soldiers may be synonymous with two of the six species of this genus described from imagoes. The other four species of nasute soldiers belong to the genera Caribitermes, Constrictotermes, Parvitermes, and Velocitermes, so it is unlikely that any of these species are synonymous with any described species since imagoes of these genera are as yet unknown in Dominican amber.
Nasutitermes rotundicephalus, new species
Figure 28
Photomicrographs of Nasutitermitinae soldiers in Dominican amber. A. Caribitermes hispaniola, n. sp., AMNH DR15-1256A. B. Parvitermes longinasus, n. sp., AMNH DR15-1255. C. Nasutitermes rotundicephalus, n. sp., AMNH DR10-1789. D. Velocitermes bulbus, n. sp., AMNH DR15-1257.
Figure 29
Caribitermes hispaniola, n. sp. soldiers in Dominican amber, AMNH DR15-1256, pieces A and B, showing locations of soldier a (head shown in figs. C, D) and soldier b (shown in figs. E, F). Figs. C, E. head in dorsal view. D, F. Head in lateral view.
Table 11
Measurements (mm) of soldiers of new species of Nasutitermitinae in Dominican amber
Diagnosis
Nasutitermes rotundicephalus has a much longer, wider, and more rounded head compared to N. electronasutus Krishna, the only soldier of Nasutitermes hitherto known from the fossil record.
Description
Soldier: Head chestnut brown; antennae, pronotum, and legs yellowish. Head with two long setae at base of nasus; four or five shorter setae at tip of nasus; tergites and sternites with very few short setae. Head dorsally viewed very wide and round behind nasus; posterior margin widely concave; nasus at base wide; dorsal margin of head in profile faintly sloping toward tip of nasus. Mandibles with short points. Antennae with 13 articles; third longer than second or fourth; second subequal to fourth. Pronotum saddle shaped; anterior margin with slight indentation in middle; posterior margin almost straight.
Etymology
This new species name is a combination of Latin rotundus, “round”, and Latin cephalus (borrowed from Greek), “head”.
Genus Parvitermes Emerson, 1949
The genus Parvitermes consists of nine living species, all from the West Indies. Of these eight, six are from Hispaniola exclusively (table 1). Parvitermes species are tropical-forest soil dwellers and feed on soil rich in humus. The new species described here, based on the soldier, is the first species of this genus known for the fossil record.
Parvitermes longinasus, new species
Diagnosis
Parvitermes longinasus differs from all the living species of Parvitermes in having a long nasus and the constriction behind the head not as prominent.
Description
Soldier: Head chestnut brown; antennae and tergites brown; legs yellowish brown. Head with four setae at base of nasus, two behind constriction, a few shorter setae at tip of nasus. Head slightly constricted behind antennal fossae; in profile vertex raised in front of constriction; sides of head round behind constriction; in lateral view constriction 0.66 mm from posterior margin of head; nasus long, wide at base. Antennae with 13 articles; third longest, almost double the length of second; second subequal to fourth. Pronotum saddle shaped; slight indentation in middle; posterior margin straight.
Etymology
This new species name is a combination of the root of Latin longus, “long”, and nasus, “nose”.
Genus Caribitermes Roisin, 1996
Caribitermes was described by Roisin et al. (1996) as a monotypic genus for the species C. discolor from the West Indies, previously placed in Parvitermes. The new species described here, based on the soldier, is the first report of this genus in the fossil record.
Caribitermes hispaniola, new species
Diagnosis
Caribitermes hispaniola resembles C. discolor (Banks), a living species from the Caribbean of the hitherto monotypic genus Caribitermes, both species having the head capsule very slightly constricted behind the antennae. The genus was described based on this character, the imago mandibles, and the gut anatomy, the latter of which cannot be studied in this fossil specimen. Caribitermes hispaniola differs from C. discolor in having a longer and wider head and nasus, fewer setae, and a longer third antennal article.
Description
Soldier: Head chestnut brown; antennae and legs yellowish brown; sternites and tergites dark brown. Head capsule with no setae visible; nasus with two or three short bristles at tip. In dorsal view constriction of head below antennal fossae not very prominent; head capsule below constriction round and wide in holotype specimen; in paratype specimen head not as bulbous (head width 0.79 mm vs. 0.82 in holotype); in lateral view constriction 0.51–0.53 mm from posterior margin of head; in profile vertex raised in front of constriction; nasus long and narrow, broad at base. Mandibles with points. Antennae with 13 articles; third slightly longer than fourth; fourth shorter than second or third. Pronotum saddle-shaped; anterior margin arched; sides angular; posterior margin faintly emarginate.
Specimens
Soldier (holotype) AMNH DR15-1256A. Paratype AMNH No. DR15-1256B (in same pieces as holotype).
Etymology
This new species named after the island of Hispaniola (Dominican Republic and Haiti).
Genus Velocitermes Holmgren, 1912
The tropical genus Velocitermes consists of 10 species, all from the Neotropical Region, none of which are reported from the West Indies. Velocitermes species live in savannah and forested areas, feeding on leaf litter (Matthews, 1977). This new species, based on the soldier caste, is the first report of the genus Velocitermes from the West Indies (Hispaniola) and the first report of this genus in the fossil record.
Velocitermes bulbus, new species
Diagnosis
Velocitermes bulbus, differs from all living species of Velocitermes in having a less constricted head and a more prominent posterior lobe of the head.
Description
Soldier: Head brown; antennae brown; abdomen blackish. Head capsule with four or five long setae and four or five short setae at tip of nasus; tergites with very short setae. Head in dorsal view constricted below base of nasus; head below constriction large and bulbous in profile; concavity between base of nasus and head; nasus narrow and slender, in profile upper margin bending downward anteriorly to tip. Mandibles with points. Antennae with 13 articles; third longest, second subequal to fourth. Pronotum saddle shaped; front margin slightly indented medially; posterior margin straight.
Discussion
In comparing the extant and extinct termite faunas on Hispaniola, it is crucial to first estimate sampling biases. The extant fauna is rather well surveyed for the Caribbean (Roisin et al., 1996; Scheffrahn and Křeček, 1993; Scheffrahn et al., 1994; Scheffrahn and Roisin, 1995; Scheffrahn et al., 1998, 2003, 2006; Snyder, 1956). The paleofauna preserved in Dominican amber, however, is undoubtedly a fraction of the entire paleofauna from Hispaniola. Hispaniola today is 76,480 km2, and the amber deposits are localized within the Cordillera Septentrional of the Dominican Republic to an area of approximately 400 km2 (Grimaldi, 1995). Like most fossil deposits, globules of amber are typically concentrated by tides and waves in shore and near shore sediments prior to burial, so the total area that originally produced the resin was certainly larger than 400 km2. Even if that area was, say, 1000 km2, it is still a small fraction of Hispaniola's present size. Given this fact, plus the bias that resin has in capturing forest arthropods associated with trees, it would not be surprising if 30% or less of the species in the original paleofauna of Hispaniola is actually represented in large samples of Dominican amber.
Despite the biased undersampling, species diversity of termites in Dominican amber is approximately the same as is known for the entire island of Hispaniola (table 1). In order to examine the relationship between termite species number and island size for the Caribbean, we plotted extant termite species diversity on 46 Caribbean islands against a log10-transformed value of island area (in km2) (table 12; fig. 31). The islands of Trinidad and Tobago were plotted in figure 31 but excluded from the regression analysis since they are virtually a peninsula of the Venezuelan mainland and, not surprisingly, have a disproportionately high number of termite species. Trinidad, for example, has 18 more termite species than Cuba, even though its size is 4% that of Cuba. There were several clear outliers in the species-area plot (fig. 31), namely two with disportionately high species numbers (Guana Island and Montserrat), and six with disproportionately few (St. Kitts, Curaçao, and a group of four Bahamas [Andros, Dry Key/Eleuthera, Grand Bahama, and San Salvador]). Guana Island is a small (3.4 km2) island close to Tortola (95 km2), both in the British Virgin Islands; the former is dry and low and has eight species, the latter is higher and even with cloud forest at the highest ridge and has five known species. The discrepancy may be due to the intensive collecting over many years (by Margaret Collins and Barbara Thorne), which may also explain the anomalous diversity reported for Montserrat. Intensive sampling plausibly uncovers rare, newly colonized species that may not become perennial inhabitants of an island. The paucity of termite species in the six islands that are low outliers is probably due to undersampling.
Figure 31
Simple regression of island size (log10 km2) and termite species number for 48 Caribbean islands (y = 1.85 + 4.39x, r = 0.74, p < 0.01). The number for each dot corresponds to the islands listed in table 12.
Table 12
Termite species diversity on Caribbean islands
Despite the outliers and confounding factors (explained below), we found a strong, general relationship between termite species diversity and island, defined by the linear relationship y = −1.85 = 4.39x (r = 0.74, p < 0.01), which follows general patterns of species numbers and island area (MacArthur and Wilson, 1967; Williamson, 1981). Though species-area relationships are explained by hypothetical rates in colonization and extinction, there are confounding factors, such as distance of islands from the mainland, age of islands, and the fact that larger islands usually have more topographic relief (Williamson, 1990), the mountains of which are usually forested and wet—conditions that sustain termites well. The Bahamas, Turks and Caicos, and smaller islands of the Virgin Islands group are low, desert islands, which may explain their paucity of termites. Nonetheless, the species-area relationship applies to islands within a group of low, dry islands as well as to a group that consists of high, forested ones (table 12; fig. 31).
By extrapolation, the large size of Hispaniola's termite paleofauna compared to that of present day suggests the island had a much larger area or a connection to the mainland. The latter possibility is supported by the presence in Dominican amber of taxa that are known to be poor dispersers over water, like stingless and orchid bees, which do not naturally occur in the Caribbean today with the exception of Trinidad and Tobago. (Melipona beecheii, occurring in Cuba, was introduced from Central America and Mexico by islanders for honey production.)
Ecologically, there appear to be significant differences in taxonomic composition between the living and extinct Hispaniolan termite faunas. Specifically, there is an unexpected diversity of soil- and humus-feeding termites in Dominican amber, such as Atlantitermes, Anoplotermes (8 species in amber and 3 living), and Subulitermes. Also, there are significantly more species of nasutitermitines in the amber (17 species) than are presently living in Hispaniola (6 species). Lastly, there are few kalotermitids in the amber (5 species), but 15 species in the Recent fauna. Kalotermitids feed and nest in sound wood, and so would be expected to have been living in close proximity to copious resin, since Hymenaea trees (the genus to which the Dominican amber tree belongs) bleed profusely in response to damage such as wind and insect attacks (Langenheim, 2003). Thus the scarcity of kalotermitids in Dominican amber may reflect a genuine absence of these termites in the amber forest, and not just an absence of data.
Another study surveyed the diversity in Dominican amber of the other major group of eusocial insects in Dominican amber, the ants (Wilson, 1985). When that study was done, 37 genera were known from Dominican amber, and 36 genera presently on Hispaniola (since then, several additional genera have been discovered in Dominican amber). Even though monophyletic genera are not equivalent units like species, this taxic approach may serve as a rough proxy for species diversity, in which case the number of known ant taxa in the amber is approximately equivalent to that in the living Hispaniola fauna, as is the case for the termites. Of the 37 ant genera in Dominican amber, 22 still exist on Hispaniola and 15 genera no longer occur on the island or in the Caribbean. Interestingly, there has been selective extinction on Hispaniola of ant genera that have specialized feeding and nesting habits, such as large colonies, flightless queens, and that are specialized predators or social parasites (Wilson, 1985). Termites are similar, with humus and soil feeding having been replaced by wood feeding, which is the primitive habit. It is not clear whether the faunal turnovers in the ants and termites are due to exclusion by competitively superior colonizers, to populational effects when an area becomes insular, or to both of these.
Biogeographically, the Dominican amber fauna is essentially a Mesoamerican one, with two exceptions. There are three species of Coptotermes in Dominican amber, which is a predominantly Asian and African genus, and in fact C. priscus appears to be the most common termite in Dominican amber. Much more dramatic is the presence of Mastotermes electrodominicus Krishna and Grimaldi in Dominican amber, whose only living congeneric relative is a species in northern Australia. Mastotermes electrodominicus too is not particularly rare in Dominican amber. These examples raise questions about the rates and extents of species turnover in biological communities and whether communities ever truly reach equilibrium.
Acknowledgments
We would like to thank Steve Thurston for his help in preparing the figures; Valerie Krishna for editorial assistance, and Michael Engel for his meticulous reading of the manuscript and his valuable comments.
