A new termite bug in Miocene amber from the Dominican Republic ( Hemiptera , Termitaphididae )

A new species of the termite bug genus Termitaradus Myers (Aradoidea: Termitaphididae) is described and fi gured based on a single female preserved in Early Miocene (Burdigalian) amber from the Dominican Republic. Termitaradus mitnicki sp. n. diff ers from the only other termitaphidid in Dominican amber, T. avitinquilinus Grimaldi and Engel, in the integumental ornamentation, number of laminae and lobules, body proportions, and setation. Th is is the third species of fossil Termitaphididae discovered. A revised key to living and fossil termitaphidids is provided.


Introduction
Th e nests of highly eusocial insects are seething with a diversity of associated arthropods, all evolved to take advantage of the rich resources provided by these ubiquitous and ecologically impressive species.Among the more unusual of inquilinous arthropods to evolve in conjunction with their eusocial hosts are those species of the aradoid family Termitaphididae (Hemiptera: Heteroptera: Pentatomomorpha).Termitaphidids, as their name suggests, are obligate inquilines in the nests of termites, particularly those of the families Termitidae and Rhinotermitidae.Termitaphidids are typically small (2-4 mm in length, although some fossils can be as large as 5.5-7 mm), ovoid, and generally dorsoventrally compressed, with fl attened lateral laminae extending from each segment.Th e family currently comprises two genera, Termitaphis Wasmann, a monotypic genus known from Colombia, and Termitaradus Myers, with eight living species in South and Central America, tropical Africa, Asia, and Australia.Although classifi ed as a separate family, it is possible that termitaphidids are highly derived mezirine Aradidae, some of which have structures similar to Termitaphididae and several of which are termitophiles (Grimaldi and Engel, 2008).Th e most detailed accounts of termitaphidid biology are those of Myers (1924Myers ( , 1932)).
Two species of termitaphidids, both of the genus Termitaradus and known only from females, have been discovered in New World Tertiary ambers.Termitaradus avitinquilinus Grimaldi and Engel was described from Early Miocene Dominican amber and is generally similar to modern species in its overall proportions, but differs most notably in the number of laminar lobules as well as the structure of the fl abella (Grimaldi and Engel 2008).Termitaradus protera Poinar and Doyen shares some similarities in lobule number with T. avitinquilinus by comparison to modern species, but is particularly noteworthy for its "gigantic" size (Poinar and Doyen 1992), which at 7.1 mm is nearly twice the length of previously described species.Hosts for these fossil species are unknown, although T. avitinquilinus is likely associated with Mastotermes electrodominicana Krishna and Grimaldi (Mastotermitidae) as two specimens were found in association with a worker of that species (Grimaldi and Engel 2008).Th is is noteworthy in that as far as has been documented modern termitaphidids are only associated with Neoisoptera (sensu Engel et al., 2009) of the families Rhinotermitidae (hosts for species of the genus Termitaradus) and Termitidae (hosts for the sole species of Termitaphis) (Wasmann 1902;Silvestri 1911Silvestri , 1921;;Mjöberg 1914;Morrison 1923;Myers 1924Myers , 1932;;Usinger 1942).Given how little termitaphidids have been studied there are likely a number of new species to be discovered and many elements of their biology remaining to be elucidated, perhaps including yet unsuspected host associations.Since the scant available information indicates an association between termitaphidids and Rhinotermitidae and Termitidae, and that these two termite families are so diverse in Dominican amber (Krishna and Grimaldi 2009), it is perhaps not surprising that a second species of termitaphidid has been discovered in this deposit (Fig. 1).Herein I provide a description of this new species as well as modifi ed keys to the identifi cation of living and fossil Termitaphididae.
Morphological terminology and format for the description generally follows that of Grimaldi and Engel (2008).Measurements were made using an ocular micrometer on an Olympus SZX-12 stereomicrosope.Diagnosis.Th e new species is immediately distinctive for the network of dorsal, segmentally-arranged carinae (Fig. 1) and the absence of the small globular nodule-like setae of other species.Th e species is larger than most other termitaphidids (2-4 mm), approximating in size the anomalously large T. protera in Mexican amber (5.8 mm vs. 7 mm).Like the other two fossil species, T. mitnicki shares a greater number of lobules (4) on the terminal abdominal segment, whereas modern species have only 2-3.Th e number of lobules on most abdominal segments is greater than in any other species, living or fossil, of termitaphidid.

Systematic
Description.Female (adult): Total length 5.8 mm, maximal width 4.0 mm (length/width ratio 1.45).Integument generally reddish brown, although paler on sterna and on dorsal carinae (Fig. 1), lobules largely reddish brown although slightly lighter than body of laminae, apices of lobules more weakly sclerotized, paler than remainder; marginal setae typically pale yellow brown although some cleared, faint (the latter owing to preservation); ventrally faintly imbricate except longitudinally wrinkled on sterna (Fig. 2), with mediolateral areas glabrous, without punctures; dorsally imbricate without punctures, with distinct and complex network of thick carinae arranged segmentally (Fig. 1), such carinae not extending onto laminae (Fig. 1), without setae.Antenna four-segmented, geniculate; fi rst article elongate, length slightly longer than combined lengths of remaining articles combined; remaining articles short, clyindrical, apicalmost apparently slightly swollen (challenging to see in specimen).Labium three-segmented, basalmost segment much shorter than others; second longest, third approximately three-quarters length of second.
Legs with stout femora, greatest widths of femora 2.5-4× times width of tibiae; femora with widely scattered short setae on outer and inner surfaces; mesofemur ventrally with longitudinal row of 3-4 stiff , erect setae at about three-quarters length; metafemur with 4-5 long, fi ne, erect setae at about four-fi fths length; tibiae with scattered short setae along lengths, particularly on outer surfaces, apically with patches of moderately dense, stiff setae; tarsi dimerous, basal tarsomere one-fourth to one-fi fth length of apical tarsomere; pretarsal ungues (= claws) large, simple, without teeth; pulvilli present, slender, straplike, lengths nearly as long as that of pretarsal ungues.
Male: Unknown.Etymology.Th e specifi c epithet is a patronym honoring Mr. Tyler Mitnick, nephew of Keith Luzzi who generously located and donated this fi ne specimen for study.

Key to adult females of Termitaphididae
Th e following key is updated and modifi ed from the one provided by Usinger (1942).1.