Coptotermes priscus Emerson

Krishna, Kumar & Grimaldi, David, 2009, Diverse Rhinotermitidae and Termitidae (Isoptera) in Dominican Amber, American Museum Novitates 2009 (3640), pp. 1-48 : 6-20

Coptotermes priscus Emerson

Figures 1 , 4 , 5 ; tables 2, 3

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

TABLE 2 Measurements (mm) of imagoes of three species of Coptotermes 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, 30 X 1.50, 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

TABLE 3 Measurements (mm) of three soldiers of Coptotermes priscus Emerson 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

Figures 6 , 8 ; table 4

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 round- ed; posterior margin faintly emarginate.

SPECIMEN: Holotype (imago) AMNH PB- 258 .

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

Figures 6 , 8 ; table 4

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.

SPECIMEN: Holotype (imago) AMNH DR10- 1512 .

ETYMOLOGY: This new species is given the Taino name for chief.

Anoplotermes carib , new species

Figures 6 , 9 , 12 ; table 4

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.

SPECIMEN: Holotype (imago) AMNH DR10- 1541 .

ETYMOLOGY: This new species is given the name of the fierce, indigenous peoples of South America who invaded the Tainos.

Anoplotermes maboya , new species

Figures 6 , 9 , 12 ; table 4

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.

SPECIMEN: Holotype (imago) AMNH DR10- 1567 .

ETYMOLOGY: This new species is given the Taino name of the nocturnal god and destroy- er of crops.

Anoplotermes naboria , new species

Figures 7 , 10 , 12 ; table 4

DIAGNOSIS: Anoplotermes naboria is closest to A. nitaino and A. quisqueya , n. spp., in its

(head slightly oblique), AMNH DR-PB258. B. A. cacique , n. sp., AMNH DR10-1512.

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 light- er 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 narrow- er than head; anterior margin deeply angular; posterolateral corners broadly rounded; posterior margin slightly indented medially. Forewing with median unbranched; Cu with 10 branches.

SPECIMEN: Holotype (imago) AMNH DR10- 1537 .

ETYMOLOGY: This new species is given the name for worker in the Taino language.

TABLE 4 Measurements (mm) of imagoes of eight new species of Anoplotermes

Anoplotermes nitaino , new species

Figures 7 , 10 ; table 4

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.

SPECIMEN: Holotype (imago), AMNH PB- 260 .

ETYMOLOGY: This new species is given the Taino name for nobleman or subchief.

Anoplotermes quisqueya , new species

Figures 7 , 11 ; table 4

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; situat- ed 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.

SPECIMENS: Holotype (imago) AMNH DR10-1528 . Paratypes (imagoes) AMNH DR10-1255 , AMNH PB-255 , PB-261 .

ETYMOLOGY: This new species is given the Taino name for Hispaniola , also meaning ‘‘mother of the earth.’’

Anoplotermes taino , new species

Figures 7 , 11 , 12 ; table 4

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.

SPECIMEN: Holotype (imago) AMNH PB- 257 .

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 visible........ 2

— Fontanelle teardrop shaped, oval, or slitlike .......................... 3

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 setae........................ 4

4. Fontanelle small, width 0.03 mm ........ 5

— Fontanelle large, width 0.05 mm ........ 7

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 mm ........................ 6

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 ; table 5

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 pigment- ed 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.

SPECIMENS: Holotype (imago) AMNH DR- 10-1255 . Paratype (imago), AMNH DR10- 1855 .

ETYMOLOGY: This new species is named after the Latin lucidus , ‘‘light’’, referring to the coloration of the head.

Figures

Fig. 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.

Fig. 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.

Fig. 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.

Fig. 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.

Fig. 8. Heads and pronota of Anoplotermes spp. in Dominican amber, dorsal views. A. A. bohio, n. sp.

Fig. 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.

Fig. 12. Forewings of Anoplotermes spp. in Dominican amber. A. A. carib, n. sp., AMNH DR10-1541. B. A. maboya, n. sp., AMNH DR10-1567. C. A. naboria, n. sp., AMNH DR10-1537. D. A. taino, n. sp., AMNH PB257.

Fig. 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).

Fig. 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.

Fig. 11. Heads and pronota of Anoplotermes spp. in Dominican amber, dorsal views. A. A. quisqueya, n. sp., AMNH DR10-1528 (paratype). B. A. taino, n. sp. (dorsofrontal view of head), AMNH DR-PB257 (holotype).

Fig. 13. Head, pronotum (dorsal view) and detail of manidibles (as preserved in Dominican amber) of Amitermes lucidus, n. sp., AMNH DR10-1563.