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AN OXYTELINE ROVE BEETLE IN DOMINICAN AMBER WITH POSSIBLEAFRICAN AFFINITIES (COLEOPTERA: STAPHYLINIDAE: OXYTELINAE)Author(s): Michael S. Engel and Stylianos ChatzimanolisSource: Annals of Carnegie Museum, 77(4):425-429. 2009.Published By: Carnegie Museum of Natural HistoryDOI: http://dx.doi.org/10.2992/0097-4463-77.4.425URL: http://www.bioone.org/doi/full/10.2992/0097-4463-77.4.425
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AN OXYTELINE ROVE BEETLE IN DOMINICAN AMBER WITH POSSIBLE AFRICAN AFFINITIES
(COLEOPTERA: STAPHYLINIDAE: OXYTELINAE)
Michael S. engelDivision of Entomology (Paleoentomology), Natural History Museum,
and Department of Ecology and Evolutionary Biology,1501 Crestline Drive – Suite 140, University of Kansas, Lawrence, Kansas 66049-2811
StylianoS chatziManoliS[Research Associate, Section of Invertebrate Zoology, Carnegie Museum of Natural History]
Department of Biological and Environmental Sciences, University of Tennessee at Chattanooga, Dept. 2653, 615 McCallie Avenue, Chattanooga, Tennessee 37403
ABSTRACT
An enigmatic new genus and species of rove beetle (Coleoptera: Staphylinidae) is described as Dolichoxenus newtoni, new genus and spe-cies, in early Miocene (Burdigalian) amber from the Dominican Republic. The species is the first fossil oxyteline discovered as an amber inclusion. The genus is apparently a member of the tribe Oxytelini (Staphylinidae: Oxytelinae) and may be allied to the African genus Jerozenia Herman, 2003. The genus exhibits several remarkable apomorphies for the subfamily, and in general possesses features otherwise known in the subfamily for inquilines of ants, suggesting a possible paleobiological association for the genus.
Key WordS: amber, Oxytelinae, paleontology, Staphyliniformia, Tertiary
A N N A L S O F C A R N E G I E M U S E U M
Vol. 77, nuMber 4, PP. 425–429 28 February 2009
The beetle family Staphylinidae is one of the largest polyphagan families with more than 51,000 described species (A.F. Newton, Jr., unpublished database). The subfamily Oxytelinae contains more than 2,000 species (Herman 2001) and has a worldwide distribution. Over the last decades there have been considerable changes in the classification of tribes within the Oxytelinae. Herman (1970) revised the genera in Oxytelinae and recognized only two tribes: Oxytelini and Coprophilini. Following the work of Newton (1982), Newton and Thayer (1992), and Hansen (1996), Newton et al. (2000) recognized four tribes: Deleasterini, Coprophilini, Thinobiini, and Oxytel- ini. Their action essentially broke up Coprophilini (sensu Herman 1970) into the tribes Deleasterini, Coprophilini sensu stricto, and Thinobiini, with the addition of genera described between 1970 and 2001. Gildenkov (2003) pro-posed a five-tribe classification (adding the tribe Mandini) and considerably reshuffled the genera within the tribes. Most recently, Makranczy (2006) also recognized five tribes; Deleasterini, Coprophilini, Planeustomini (= Man-dini of Gildenkov 2003), Blediini, and Oxytelini sensu novo, combining the tribes Oxytelini and parts of the tribe Thinobiini of Newton et al. (2000). The oldest definitive polyphagan beetle is a staphylinid, from the Late Triassic (Carnian) of Virginia (ca. 225–230 Ma, Fraser et al. 1996) and despite the great number of extant species, relatively few staphylinid fossils have been described (vide Herman 2001 for a summary). This is most likely due to the lack of interest from staphylinid system-atists, because there is a plethora of fossil staphylinids in
museums around the world and new material is continually coming to light (Engel and Chatzimanolis personal obser-vations). Several fossils have been described belonging to the subfamily Oxytelinae, mostly from formations in Rus-sia or from the prolific shales of Florissant, Colorado, in the United States. Herman (2001) listed 22 fossil species of Oxytelinae, 16 of them belonging to extant genera (eight in Bledius Leach, 1819, one in Deleaster Erichson, 1839, five in Oxytelus Gravenhorst, 1802, and two in Platystethus Mannerheim, 1830), and four belonging to extinct genera (two in Mesoxytelus Tikhomirova, 1968, one in Morda Ryvkin, 1990, and one in Turgaphloeus Ryvkin, 1990). The extinct genus Megalymma Tikhomirova, 1980, with two species, was also listed by Herman (2001) as it was explicitly placed in Oxytelinae by Tikhomirova (1980). However, Tikhomirova (1980) included the Omaliinae as a tribe of Oxytelinae and placed her genus near Micra-lymma Westwood, 1838. Accordingly, Megalymma should perhaps be tentatively considered an omaliine of uncertain identity. A further likely fossil oxyteline species, Pliosyn-tomium schmidti Korge, 1967, has been described from the Pliocene of Willershausen, Germany (Korge 1967). All of these are preserved as compressions with little or no re-lief, and in some cases cannot be conclusively assigned to tribe or even subfamily. A thorough revision of the fossil staphylinids is certainly needed, and fossil species incor-porated into cladistic analyses with their modern counter-parts. Herein we describe the first fossil oxyteline discovered as an amber inclusion, and the first from the Tertiary of
INTRODUCTION
426 annalS oF carnegie MuSeuM Vol. 77
Hispaniola. We discuss the possible affinities of this spe-cies with the African genus Jerozenia Herman, 2003. Some oxytelines have been reported in middle Eocene Baltic am-ber (e.g., Menge 1856; Scudder 1885, 1886, 1891; Helm 1896; Handlirsch 1907; Klebs 1910; Bachofen-Echt 1949; Larsson 1978; Spahr 1981). These authors report the genera Bledius, Oxytelus, Carpelimus Leach, 1819, and Planeu- stomus Jacquelin du Val, 1857, but none has been described nor have specimens been located to confirm generic iden-tifications.
MATERIALS AND METHODS
Morphological terminology generally follows that used elsewhere in the systematics of Oxytelinae (e.g., Herman 1970, 2003; Newton 1982). Measurements were made using an ocular micrometer on an Olympus SZX-12 ste-reomicroscope and microphotography using a Nikon D1x digital camera attached to an Infinity K-2 long-distance
microscope lens. The age and origin of Dominican amber has been summarized by Grimaldi and Engel (2005).
SYSTEMATIC PALEONTOLOGY
Order Coleoptera Linnaeus, 1758Suborder Polyphaga Emery, 1886
Family Staphylinidae Latreille, 1802Subfamily Oxytelinae Fleming, 1821
Tribe Oxytelini Fleming, 1821
Dolichoxenus, new genus
Type species.—Dolichoxenus newtoni, new species.
Diagnosis.—The new genus can be immediately recog-nized from other genera of Oxytelinae by the combination of a largely vestigial compound eye, subtrapezoidal head, apparently broad gula (refer to description), long antenna with tightly adjoined scape and pedicel (appearing in many
Fig. 1.—Photomicrographs of holotype of Dolichoxenus newtoni, new genus and species (MACT-1810). A, Dorsal oblique aspect of entire specimen (specimen ca. 2.8 mm in length, excluding antennae); B, Dorsal oblique aspect of head and thoracic dorsum (antennal length 1.1 mm).
2009 engel and chatziManoliS—oxyteline roVe beetle in doMinican aMber 427
views as a single article), dense punctation, and elongate pronotum are diagnostic for the genus.
Description.—f: Body elongate, slender, rather cylindrical (Fig. 1A). Head, pronotum, elytra, and abdomen with numerous setae, those on api-cal abdominal segments longest. Head and pronotum depressed medio-longitudinally. Head elongate, depressed medially, subtrapezoidal with widest section at anterior (Fig. 1B), lateral surfaces with anterior depres-sions bearing antennal insertion, such depressions becoming gradually shallower to just before head midlength, anteriorly making supraäntennal ridge prominent (but ridge not extending posteriorly beyond head mid-length and not forming posterior lobe); clypeal margin slightly reflexed, extending moderately beyond apex of supra-antennal ridge; epistomal sulcus faint but apparently complete; labrum apparently very short, api-cal margin hidden, with medial fringe of long, stout, anteriorly-directed setae; mandibles not visible; labial palpus tetramerous, P3 (= third pal-pomere) largest, swollen, P4 cylindrical, subulate, longer than wide, 0.6x length of P3; P1 and P2 very short; mentum subtrapezoidal, flat, api-cal margin straight; gula relatively wide, apparently fused anteriorly for only a short distance, then broadened rapidly and tapering divergently toward posterior margin of head until slightly converging near posterior head margin (very difficult to discern owing to nature of preservation but apparently somewhat similar to condition seen in Deleaster, although gula appearing broader immediately posterior to short fused section of gular sulci). Antenna long, with 11 antennomeres, each article with whorls of numerous, long, thin setae; scape and pedicel tightly adjoined and appearing as a single article (Fig. 1B). Compound eyes dramatically reduced, eye diameter less than one-half scape width, positioned slightly above antennal insertion (size of ommatidia not clear). Base of head not constricted to form neck. Pronotum much longer than wide. Legs long, slender (Fig. 1A); pro-trochantin hidden; procoxae contiguous; prosternal process short, trian-gular, not carinate; tibiae without spinelike setae, without rows of setae; scattered, moderate-length, curved setae near tibial apices (weakly so on meso- and metatibiae), such setae also present on ventral surface of basitarsi; tarsal formula 3-3-3; mesocoxae nearly contiguous (Fig. 2A), narrowly separated posteriorly by short metasternal carina; mesosternal process short, obtusely rounded, extending just past and slightly over anterior borders of mesocoxae; metasternum depressed, without medi-al carina. Elytra longer than wide; scutellum slightly exposed, surface rounded, not impressed. Abdominal segments taper in width apically, giving abdomen a cy-lindrical appearance (superficially similar to some Piestinae); parater-gites difficult to discern (faint or weak and abdomen largely compressed so distortion makes them challenging to discern), apparently with two paratergites on second through sixth (perhaps seventh) abdominal seg-ments; ninth tergum completely divided by tenth tergum (Fig. 2B); sec-ond sternum fully developed, distinct, apparently mobile, as long as third sternum; second and third sterna without median intercoxal carinae.
Etymology.—The new genus-group name is a combina-tion of the Greek words dolichos (meaning, “long”) and xenos (meaning, “stranger”). The name is masculine.
Dolichoxenus newtoni, new species(Figs. 1–2)
Diagnosis.—As for the genus (see above).
Description.—f: Coloration of integument apparently dark brown, nearly black and not shining; integument throughout body with distinct, contiguous punctures, integument between punctures strongly imbricate. Body elongate and slender, length ca. 2.8 mm as preserved (excluding antennae). Head 0.37 mm in length (as measured along midline), anterior width 0.29 mm, posterior width 0.21 mm; with deep, sparse punctation; with numerous setae. Antennae long, 1.1 mm in length; pedicel compact, wider than long, as wide as scape, and tightly adjoining apex of scape
(most easily discerned ventrally); antennomeres 3–11 each globose, with slender basal neck (shortest on first antennomere), gradually increasing in size from base to apex, each slightly wider than long except anten-nomere 11 longer than wide and with gently conical apex; no distinct club. Compound eyes reduced, compound eye length approximately 12% length of head; neck absent. Pronotum much longer than wide, depressed mediolongitudinally, 1.4 times as long as head and 0.89 times as long as elytra, with numerous setae. Elytra narrow, length 0.57 mm, width of individual elytron 0.19 mm; surface strongly imbricate, without visible grooves or ridges, with punctures more shallow and scattered medially, those along lateral margins more dense, nearly contiguous, with setae largely confined to lateral borders. Legs long, slender; protrochantin hidden; procoxae contiguous; prosternal process short, tri-angular, not carinate; tibiae without spinelike setae, without rows of setae; scattered, moderate-length, curved setae near tibial apices (weakly so on meso- and metatibiae), such setae also present on ventral surface of ba-sitarsi; tarsal formula 3-3-3; mesocoxae narrowly separated posteriorly by short metasternal carina; mesosternal process short, obtusely rounded, extending just past and slightly over anterior borders of mesocoxae; metasternum depressed, without medial carina. Abdomen with punctures progressively becoming more shallow and slightly more widely spaced (0.5 times a puncture width or less) on progressively more apical terga and sterna; terga with scattered, short, posteriorly-directed setae, setae becoming elongate, fuscous, and more dense laterally; sterna with scattered, short, posteriorly-directed setae.
Holotype.—Female, MACT-1810; early Miocene (Burdi-galian) amber; Dominican Republic; Morone Amber Col-lection, Turin.
Etymology.—The specific epithet is a patronymic honor-ing our friend and colleague Dr. Alfred F. Newton, Jr., a leading authority on the systematics of Staphylinoidea, among other lineages of Coleoptera.
DISCUSSION
The placement of Dolichoxenus in Oxytelinae can be confirmed by the well-developed second abdominal ster-num (reduced in the other subfamilies) and the complete
Fig. 2.—Thoracic and abdominal structures of Dolichoxenus newtoni, new genus and species (MACT-1810). A, Ventral oblique aspect of meso- and metathoracic segments (drawn as preserved and in view that is most easily seen); B, Dorsal, slightly oblique aspect of abdominal apex depict-ing tenth and divided ninth terga (drawn as preserved – the abdomen is somewhat crumpled and the apical ends of the ninth tergum are slightly pulled away from the extended tenth tergum). Scale bars = 0.25 mm.
428 annalS oF carnegie MuSeuM Vol. 77
division of the ninth tergum by the tenth. Dolichoxenus probably belongs to the tribe Oxytelini (sensu Newton et al. 2000) based on the separation of the mesocoxae by a metasternal process, the 3-3-3 tarsal formula, and the mor-phology of the labial palpomeres. However, the gular sulci appear to be somewhat more broadly separated than in most Oxytelini, reminiscent in some manner to those of Deleaster. Given the difficulty discerning the gular sulci, this trait must await further confirmation based on more finely preserved material. In addition, the presence or ab-sence of basolateral ridges on the abdominal terga and the relative size of the ommatidia are not observable or clearly discernible in the specimen. All members of Oxytelinae have 11 antennal articles (except the African Jerozenia co-mata Herman, 2003, which has nine: Herman 2003), and while Dolichoxenus has this typical complement, it has the general appearance of only ten articles owing to the tight position of the pedicel against the apex of the scape. This structure of the pedicel and scape renders the former difficult to discern and serves to make the genus readily recognizable among other oxytelines. Similar to Jerozenia, the mesocoxae of Dolichoxenus are separated by a nar-row metasternal ridge, while the mesocoxae of most other genera in Oxytelini (sensu Herman 1970) are separated by a broad metasternal process. The peculiar body form is highly reminiscent of other oxytelines that are known or suspected to be specialized inquilines of ants (e.g., Ecitoclimax Borgmeier, 1934, and Jerozenia). It is likely that Dolichoxenus had a similar biology and lived in ant colonies. Even though a formal phylogenetic analysis is lacking, it is reasonable to consider that Dolichoxenus and Jerozen-ia are closely related as evidenced by the aforementioned traits. This affinity is particularly interesting given that Jerozenia is known from central Africa (Democratic Re-public of the Congo), whereas Dolichoxenus was found in Dominican amber. There has been considerable discussion regarding the paleogeographical affinities of Dominican or Mexican amber specimens with extant African or other Old World organisms. Grimaldi (1995) documented flies in amber of the genus Ogcodes Latreille, 1797, that have their closest relatives in Africa. Engel and Grimaldi (2007) discussed examples from Neuroptera, and Poinar and Brown (2002) speculated for species of Hymenaea Lin-naeus, 1753 (Fabaceae), although the last was based on su-perficial similarity and strictly on plesiomorphic features (which are not indicative of relationship) rather than any apomorphic features demonstrating such a relationship. Moreover, regional extinction of Old World lineages other-wise present in the Tertiary of Hispaniola extends to ants of the genus Leptothorax Mayr, 1855 (Baroni Urbani and Wilson 1987), termites of the genus Mastotermes Froggatt, 1897 (Krishna and Grimaldi 1991), protopaussine beetles (Nagel, 1997), and wood gnats of the genus Valeseguya Colless, 1990 (Grimaldi 1991), among others. Dolicho-xenus, like these other examples, was perhaps a relic in its day, representative of an earlier diversification (perhaps in
the mid-Tertiary) that persisted in a few tropical regions of the Northern Hemisphere before eventually becoming broadly extinct and leaving relic taxa in warmer habitats of the Old World.
ACKNOWLEDGMENTS
We are grateful to Dott. Ettore Morone for bringing this interesting mate-rial to our attention, permitting us to study it, and for his gracious hos-pitality during the senior author’s visit to Italy. We also thank Charles Nelson and two anonymous reviewers for studying earlier versions of the manuscript. Support for this work was provided by National Science Foundation grant DEB-0741475 to S. Chatzimanolis and M.S. Engel. This is a research contribution of the Division of Entomology, University of Kansas Natural History Museum.
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