The Fauna of Staphylininae in Dominican Amber (Coleoptera: Staphylinidae)

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The Fauna of Staphylininae in Dominican Amber (Coleoptera: Staphylinidae)Author(s): Stylianos Chatzimanolis Michael S. EngelSource: Annals of Carnegie Museum, 81(4):281-294. 2013.Published By: Carnegie Museum of Natural HistoryDOI: http://dx.doi.org/10.2992/007.081.0401URL: http://www.bioone.org/doi/full/10.2992/007.081.0401

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THE FAUNA OF STAPHYLININAE IN DOMINICAN AMBER (COLEOPTERA: STAPHYLINIDAE)

STYLIANOS CHATZIMANOLIS[Research Associate, Section of Invertebrate Zoology, Carnegie Museum of Natural History]

Department of Biological and Environmental Sciences, Dept. 2653, University of Tennessee at Chattanooga, 615 McCallie Avenue, Chattanooga, Tennessee 37403

[email protected]

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 66045

[email protected]

A N N A L S O F C A R N E G I E M U S E U MVOL. 81, NUMBER 4, PP. 281–294 30 SEPTEMBER 2013

ABSTRACT

The fauna of staphylinine rove beetles (Staphylinidae: Staphylininae) in early Miocene (Burdigalian) amber of the Dominican Republic is described and gured. In total, seven species are recognized representing two tribes, four subtribes, and ve genera. The following ta a are described as new species: Heterothops cornelli, Heterothops infernalis, Holisus funeratus, Philonthus hades, Philonthus rhadamanthus, Quedius electrodominicanus, and Neoxantholinus apolithomenus. A brief summary of Dominican and Me ican amber Staphylinoidea is pro-vided and comments included on the biogeography of ancient Hispaniola.

KEY WORDS: amber, Miocene, paleontology, rove beetles Staphylininae, ta onomy

INTRODUCTION

The rove beetles, family Staphylinidae, constitute one of the most diverse and ecologically successful clades of Coleoptera, with over 57,000 described species (Newton unpubl. data) and an innumerable number of yet unde-scribed ta a. The subfamily Staphylininae contains more than 320 genera and 6,900 species (Thayer 2005) and is one of the most speciose subfamilies in Staphylinidae. Yet, the number of fossils described in Staphylininae is remarkably small in contrast to other subfamilies (see for e ample Chatzimanolis and Engel 2011 Chatzimanolis et al. 2012). The fossil species described from mid-Eocene Baltic amber were recently summarized by Chatzimanolis and Engel (2011), while the Mesozoic record of the fam-ily, compressions and amber inclusions, has been given by Chatzimanolis et al. (2012) and updated by Chatzimanolis et al. (2013). Since the late 1960s the amber mines of the Dominican Republic have produced one of the most abundant sources of amber-preserved insects in the world, second only to the massive deposits of Baltic amber, which, in contrast, have been e cavated for millennia. Recent catalogs of arthro-pods, insects in particular, in Dominican amber have been provided by Pérez-Gelabert (1999, 2008) and Arillo and Ortuño (2005), although these are now incomplete (and were missing some ta a and references even when pub-lished), and a new annotated checklist of this fauna and

ora is needed. Nonetheless, these sources are a good en-try point into the literature for Dominican amber insects. Herein, we describe the rst fossils of Staphylininae from Dominican amber and provide an overview of the

described species from Tertiary ambers of the New World as well as those genera recorded in the literature (Table 1). In total, there are nine described species of Staphylini-dae (e cluding the current study) from Dominican amber (contra Pérez-Gelabert 2011, who lists 24), as well as numerous records of undescribed material, most in private collections (e.g., Poinar and Poinar 1999 Irmler 2003). In this paper we describe seven species, all previously unknown.

MATERIALS AND METHODS

As is true for most staphylinid genera, many of the groups discussed below have never been revised and several ge-neric concepts are old and or ambiguous as de ned by the included living species. This naturally hinders interpreta-tion of the fossil fauna, particularly as full revision of the living species for comparison with the fossils is well be-yond the scope of the present study. Despite this challenge, we place the fossils based on current operational concepts for the genera, recognizing that future monographs and cladistic tests may signi cantly alter the circumscription of some of these genera (e.g., Philonthus). Nonetheless, our hope is to provide an initial account of the Domini-can amber staphylinine diversity in the light of current knowledge of the associated living fauna. Morphological terminology generally follows that used elsewhere in the systematics of Staphylininae (e.g., Blacklwelder 1936). Measurements were made using an ocular micrometer on an Olympus SZX-12 stereomicroscope and microphotog-raphy using a Nikon D1 digital camera attached to an In nity K-2 long-distance microscope lens. Metrics are

282 ANNALS OF CARNEGIE MUSEUM VOL. 81

TABLE 1. Hierarchical classi cation and checklist of described or recorded (to at least genus level) Staphylinoidea(sensu Bouchard et al. 2011) in early Miocene Dominican and Me ican amber, including ta a proposed herein.

Some of the generic records must be considered tentative as they have appeared merely as unveri ed names in listsand without any specimen numbers or repositories, and many of these appear dubious (e.g., Poinar and Poinar 1999).

Family LEIODIDAE Fleming Subfamily Leiodinae Fleming Tribe Scotocryptini Reitter Aglyptinus poinari Perkovsky, 2000 DominicanFamily STAPHYLINIDAE Latreille Subfamily Pselaphinae Latreille Tribe Batrisini Reitter Subtribe Batrisina Reitter Arthmius sp. [mentioned (Poinar and Poinar 1999)] Dominican Tribe Dimerini Raffray Barroeuplectoides sp. [mentioned (Poinar and Poinar 1999)] Dominican Tribe Jubini Raffray Jubus sp. [mentioned (Poinar and Poinar 1999)] Dominican Tribe Trichonychini Reitter Subtribe Panaphantina Jeannel Thesium sp. [mentioned (Poinar and Poinar 1999)] Dominican Subtribe Trimiina Bowman Trimium sp.1 [mentioned (Poinar and Poinar 1999)] Dominican Tribe Brachyglutini Raffray Subtribe Brachyglutina Raffray Brachygluta sp.2 [mentioned (Poinar and Poinar 1999)] Dominican Scalenarthrus sp. [mentioned (Poinar and Poinar 1999)] Dominican Tribe Arhytodini Raffray Caccoplectus sucineus Chandler and Wolda, 1986 Me ican Tribe Tyrini Reitter Subtribe Somatipionina Jeannel Apharus sp. [mentioned (Poinar and Poinar 1999)] Dominican Subfamily Tachyporinae MacLeay Tribe Tachyporini MacLeay Tachyporus sp. [mentioned (Poinar and Poinar 1999)] Dominican Subfamily Aleocharinae Fleming Tribe O ypodini Thomson Oxypoda bindosa Seevers, 1971 Me ican Tribe Hypocyphtini Laporte Ambracyptus minutissima (Seevers, 1971) Me ican Tribe Trichopseniini LeConte and Horn Prorhinopsenius alzadae Kistner, 1998 Dominican Prorhinopsenius mexicanus (Seevers, 1971) Me ican Prorhinopsenius poinari Kistner, 1998 Dominican Prorhinopsenius seeversi Kistner, 1998 Dominican Subfamily Osoriinae Erichson Tribe incertae sedis Lispinomimus atavus Irmler, 2003 Dominican

2013 CHATZIMANOLIS AND ENGEL—STAPHYLININAE IN DOMINICAN AMBER 283

TABLE 1. Hierarchical classification and checklist of described or recorded (to at least genus level) Staphylinoidea(sensu Bouchard et al. 2011) in Early Miocene Dominican and Me ican amber, including ta a proposed herein

(continued from previous page).

Tribe Osoriini Erichson Neosorius spp. [mentioned (Irmler 2003)] Dominican Osoriellus sp. [mentioned (Irmler 2003)] Dominican Tribe Thoracophorini Reitter Subtribe Thoracophorina Reitter Thoracophorus palaeobrevicristatus Irmler, 2003 Dominican Subtribe Lispinina Bernhauer and Schubert Liberiana sp.3 [mentioned (Irmler 2003)] Dominican Nacaeus dominicanensis Irmler, 2003 Dominican Subfamily O ytelinae Fleming Tribe O ytelini Fleming Dolichoxenus newtoni Engel and Chatzimanolis, 2009 Dominican Subfamily Scydmaeninae Leach Tribe Mastigini Fleming Mastigus sp.4 [mentioned (Poinar and Poinar 1999)] Dominican Tribe Cyrtoscydmini Schaufuss Euconnus sp. [mentioned (Franz 1983)] Dominican Homoconnus conservatus Franz, 1983 Dominican Microscydmus sp. [mentioned (Poinar and Poinar 1999)] Dominican Neuraphes fossilis Franz, 1983 Dominican Subfamily Paederinae Fleming Tribe Pinophilini Nordmann Subtribe Procirrina Bernhauer and Schubert Palaminus spp. [mentioned (Seevers 1971 Herman 2010)] Me ican Dominican Subfamily Staphylininae Latreille Tribe Staphylinini Latreille Subtribe Amblyopinina Seevers Heterothops cornelli, new species Dominican Heterothops infernalis, new species Dominican Subtribe Hyptiomina Casey Holisus funeratus, new species Dominican Subtribe Philonthina Kirby Philonthus hades, new species Dominican Philonthus rhadamanthus, new species Dominican Subtribe Quediina Kraatz Quedius electrodominicanus, new species Dominican Tribe Xantholinini Erichson Neoxantholinus apolithomenus, new species Dominican Xantholinini undet. [Poinar and Brodzinsky 1985: g. 1] Dominican 1 Listed as “Trimiena, but such a genus does not e ist. Presumably they were referring to subtribe Trimiina and likely genus Trimium Aubé.2 Listed as “Brachyglutina,” i.e., with the subtribe erroneously treated as a genus (Poinar and Poinar 1999). The authors presumably meant Brachygluta Thomson.3 Liberiana Blackwelder is considered a synonym of Nacaeus Blackwelder by Herman (2001).4 Misspelled as “Mastigonius” by Poinar and Poinar (1999).


provided for the holotype. The ideal angle for measuring was not always possible given the state of preservation or orientation of specimens relative to the amber surface, the latter of which could not always be polished given the

pro imity of inclusions. The age and origin of Dominican am-ber has been summarized by Grimaldi (1995), Itturalde-Vinent and MacPhee (1996), and Grimaldi and Engel (2005). Fos-sils were studied from the following institutions: American

Fig. 1.—Photomicrographs of Heterothops cornelli (A–F) and H. infernalis (G–H). A, ventral view of paratype AMNH DR-10-764, scale bar = 0.35 mm B, dorsal view of holotype AMNH DR 10-1212, scale bar = 0.44 mm C, holotype AMNH DR-10-1212 showing details of head and pronotum, scale bar = 0.31 mm D, dorsal view of paratype AMNH DR-10-1214, scale bar = 0.64 mm E, paratype AMNH DR-10-1213, scale bar = 0.35 mm F, lateral view of specimen 11202 (originally in J. Cornell collection, now in FMNH), scale bar = 0.64 mm G, ventral view of holotype of H. infernalis (FMNH), scale bar = 0.5 mm H, dorsal view of holotype of H. infernalis (FMNH), scale bar = 0.5 mm.


Museum of Natural History, New York, New York (AMNH, David A. Grimaldi), James Cornell personal collection (to be deposited in FMNH), and the Field Museum of Natural History (FMNH, Margaret K. Thayer, Alfred F. Newton).

SYSTEMATIC PALEONTOLOGYFamily Staphylinidae Latreille, 1802

Subfamily Staphylininae Latreille, 1802Tribe Staphylinini Latreille, 1802

Subtribe Amblyopinina Seevers, 1944Genus Heterothops Stephens, 1829

Heterothops cornelli, new species(Figs. 1A–F)

Diagnosis.—Heterothops cornelli has similar head chaeota y to H. confertus Smetana, but it can be easily distinguished from that species based on the overall size: H. confertus is rather large (2.8–3.8 mm), while H. cornelli is much smaller (1.81–1.92 mm). Heterothops cornelli can be distinguished from the two Recent West Indies species (H. ocularis Blackwelder and H. rambouseki Blackweld-er) based on the punctation patterns of the head: in H. cor-nelli there are two punctures between the eyes, while in the Recent West Indies species there are none. Comparisons with H. conticens Scudder from Florissant, Colorado are hard due to the incomplete preservation of that species, but H. conticens Scudder is clearly much larger than H. cornelli. Additionally, H. cornelli can be distinguished from H. infernalis (described below) because the general habitus of H. cornelli is larger and wider, and the head is less elongate than in H. infernalis.

Description.— small size, length 1.81–1.92 mm. Coloration.—(Fig. 1B): color of head brown pronotum reddish-brown elytra, legs, antennae, and abdomen yellowish-brown antenno-mere 11 lighter yellow. Head.—(Fig. 1C) length 0.29–0.31 mm, width 0.25 mm head longer than wide, much narrower than pronotum chaeota y of dorsal side of head (see Smetana 1971, 1978) with one anterior and posterior frontal setiferous puncture on each eye and two other punctures between them but closer to the posterior one. Eyes medium-sized, as long as 1/3 of head, positioned anteriorly. Antennomeres (Fig. 1A) 1–2 longer than wide, club-like, much wider than antennomeres 3–5 antennomeres 3–4 longer than wide, antennomere 4 slightly longer than 3 antennomeres 5–10 sub uadrate antennomere 11 longer than wide, as wide as anten-nomere 10 but 2.5 times as long. Mandibles (Fig. 1C) short but thin and sharp apically. Ma illary palpi with palpomere 1 short palpomere 2–3 club-like, longer than wide, palpomere 3 more dilated apically than 2 palpomere 4 conical, sharp, shorter than 3, less than ½ as wide as pal-pomere 3. Labial palpi with palpomere 1 not visible palpomere 2 slightly shorter and much wider than palpomere 3 palpomere 3 straight, narrow and thin. Neck absent. Thorax.—Pronotum (Fig. 1D) length 0.33 mm, width at posterior margin 0.38–0.39 mm, wider posteriorly, narrower and shorter than elytra, with rounded anterolateral corners. Lateral margins of pronotum conve , e cept posterior 1/5 concave. Pronotum with one row of two setiferous punctures on each side of the central disk, and few setifer-ous punctures along posterior and lateral borders. Mesoscutellum promi-nent, medial length 0.13 mm. Elytra (Fig. 1E) length 0.44 mm, width 0.58 mm, slightly wider posteriorly, with small uniform punctures. Wings fully developed. Legs with tarsal formula 5-5-5 long, densely punctated.

Protibia with pair of spines distally, meso- and metatibia with several spines. Tarsi setiferous. Abdomen.—(Fig. 1E–F) with segments gradually narrowing posteri-orly ma imum abdominal width 0.46 mm. Abdominal terga and sterna with dense uniform setiferous punctures. : Unknown.

Holotype.— : AMNH DR-10-1212, labeled “AMBER: Oligo-Miocene, Dominican Republic, AMNH no DR-10-1212” / “Dominican Republic, Palo Alto, Santiago Prov.” / “Staphylinidae, Staphylininae, Heterothops det. L. Her-man” / “HOLOTYPE Heterothops cornelli Chatzimanolis and Engel” deposited in AMNH.

Paratypes.—6 : Specimens DR-10-1213, DR-10-1214 with labels: “AMBER: Oligo-Miocene, Dominican Re-public, AMNH no DR-10-1213 [or 1214 for other para-type]” / “Staphylinidae, Staphylininae, Heterothops, det. L. Herman” deposited in AMNH. Specimen DR-10-764 with locality label as above but with det. label, “Staphy-linidae, Staphylininae, Quediini det. L. Herman” deposited in AMNH. Specimen DR-8-367 with labels, “Amber: Do-minican Republic, Oligo-Miocene, speci c provenance un-known. Purchased in Santo Domingo by D. Grimaldi. AMNH DR-8-367” / “Staphylinidae, Staphylininae, Heterothops, det. L. Herman” deposited in AMNH. FMNH (Jim Cor-nell 11202 and an unnumbered Jim Cornell specimen) deposited in FMNH. All paratypes with label “PARATYPE Heterothops cornelli Chatzimanolis and Engel”.

Etymology.—The speci c epithet is a patronym honoring the late James F. Cornell, Jr. (1940–2012), proli c collec-tor of living and fossil Staphylinoidea and who permitted us to study his personal material as part of this study (spec-imens now in the FMNH).

Heterothops infernalis, new species(Figs. 1G–H)

Diagnosis.—This is a rather poorly preserved specimen, but distinctive enough from H. cornelli the general hab-itus is much smaller and slimmer and the head is more elongate than in H. cornelli. Heterothops infernalis can be distinguished from the two Recent West Indies species (H. ocularis and H. rambouseki Blackwelder) based on the punctation of the head: the head of H. infernalis is impunc-tate, while the head the head of the two Recent species is punctate. Comparisons with the fossil species known from Florissant, Colorado are hard due to the incomplete preser-vation of that species, but H. conticens Scudder is clearly much larger than H. infernalis.

Description.— small size, total length as preserved 1.64 mm. Coloration.— (Fig. 1G) integument throughout dark reddish brown, darker on head and abdomen than remainder of body (perhaps preserva-tional) integument dull. Head.—(Fig. 1H) narrow, narrower than pronotum, longer than wide width across compound eyes 0.19 mm medial length 0.23 mm frontal part (before compound eyes) prolonged apically. Eyes weakly conve . Chaeota y of dorsal side of head (see Smetana 1971, 1978) with two


setiferous punctures at upper and lower angles of posterior border of each eye, without setiferous puncture at anterior border, otherwise head ap-parently impunctate and faintly imbricate temple about as long as (per-haps slightly longer than) eye in dorsal view. Antenna (Figs. 1G–H) thin throughout length, antennomeres 1–3 longer than wide antennomeres 1 and 2 club-like, much wider than remaining antennomeres antennomere 3 as long as 4 but slightly narrower, distinctly shorter and narrower than 2 remaining antennomeres longer than wide antennomere 11 at least three times longer than wide. Mouthparts not clearly visible e cept ma -illary palpus typical of Heterothops. Thorax.—Pronotum (Fig. 1H) wider than long, ma imum width at posterior, width 0.30 mm, medial length 0.26 mm, lateral margins con-verging anteriorly, faintly conve , posterior margin relatively straight, anterior margin relatively straight integument apparently faintly imbri-cate and impunctate e cept where setae are present setae along lateral margin, not evident on disc. Mesoscutellum large, triangular, medial length 0.12 mm. Elytra longer than pronotum, length (as measured along lateral margin) 0.41 mm, width of elytron 0.17 mm, lateral margins par-allel, posterior border concave integument apparently coarsely imbri-

cate to granulose, with numerous, suberect to seemingly appressed setae across surface. Hind wings (Figs. 1G–H) fully developed. Legs with tar-sal formula 5-5-5 long, densely punctated. Abdomen.—(Figs. 1G–H) widest basally, width 0.30 mm, tapering in width apically densely setiferous, apical and lateral setae particularly elongate and stiff integument apparently as on elytra. : Unknown.

Holotype.— , FMNH, labeled, “DOMINICAN REPUB-LIC: La Toca mine in the Northern, coastal range, elev. 1000, 75% probability from this area, Dominican amber, Field Museum Nat. Hist.” / “HOLOTYPE Heterothops in-fernalis Chatzimanolis and Engel” deposited in FMNH.

Etymology.—The speci c epithet is the Latin term infer-nalis, referring to the underground as an allusion to its be-ing a fossil species.

Fig. 2.—Photomicrographs of Holisus funeratus (AMNH DR-10-1199). A, dorsal view, scale bar = 0.5 mm B, ventral view, scale bar = 0.5 mm C. details of the head, scale bar = 0.25 mm.


Subtribe Hyptiomina Casey, 1906Genus Holisus Erichson, 1839

Holisus funeratus, new species(Figs. 2A–C)

Diagnosis.—Among Holisus species distributed in West Indies, H. funeratus can be distinguished from H. laevis

Blackwelder and H. guidingii Erichson by the lack of a carinate labium and the black coloration of the head of these two species (head dark brown in H. funeratus). Holisus funeratus looks rather similar to H. debilis Erich-son but can be distinguished from the later species based on the punctation of the head: in H. debilis punctures are more closely spaced together than in H. funeratus.

Fig. 3.—Photomicrographs of Philonthus hades. A, lateral view of holotype AMNH DR-10-1216, scale bar = 1.13 mm B, lateral view of holotype AMNH DR-10-1216 showing details of head, thora , and legs, scale bar = 0.77 mm C, lateral view of holotype AMNH DR-10-1216 showing details of abdomen, scale bar = 0.77 mm D, lateral view of paratype AMNH DR-10-1215, scale bar = 1 mm E, dorsal view of paratype AMNH DR-10-1215, scale bar = 1 mm F, lateral view of paratype AMNH DR-10-1215 showing details of elytra and pronotum, scale bar = 0.63 mm G, lateral view of paratype AMNH DR-10-1215 showing details of head, mouthparts, and antenna, scale bar = 0.63 mm.


Description.— small size, length 3.42 mm. Coloration.— (Fig. 2): color of head and antennae dark brown, ap-pearing metallic pronotum, elytra, legs, and abdomen reddish-brown posterior margin of abdominal segments III–VI orange. Head.—(Fig. 2C) sub uadrate, length 0.42 mm, width 0.48 mm with large, deep, setiferous punctures dorsally punctures almost contiguous on lateral sides and posteriorly, but distance between punctures increases towards middle of head middle of head without punctures. Postgena (Fig. 2B) with smaller punctures punctures not uniformly distributed. Head covered throughout with equidistant micropunctures. Eyes (Fig. 2C) small, as long as 1/4 of head, positioned anteriorly. Antennomeres with large punctures antennomere 1–3 longer than wide, clavate an-tennomere 2 slightly smaller than antennomere 3 antennomeres 4 sub-quadrate, slightly larger than antennomere 5, antennomere 5 quadrate antennomere 6 quadrate, subequal to antennomere 5 antennomeres 7–10 transverse, gradually increasing in width antennomere 11 longer than wide, as wide as antennomere 10 but twice as long. Mandibles long, sym-metrical, with small medial tooth and shallow longitudinal groove each from base to tooth. Ma illae and labium punctuated, aciculate. Apical ma illary and labial palpomeres shorter than penultimate. Gular sutures fused, e cept anteriorly. Neck with deep contiguous punctures. Thorax.—Pronotum (Fig. 2A) quadrate, length 0.46 mm, width 0.44 mm, subequal in legth to head but shorter than elytra, with rounded an-terolateral corners. Lateral margins of pronotum conve , e cept poste-rior 1/5 concave. Pronotum with deep contiguous punctures. Pronotal hypomeron not visible. Metasternum with shallow carina medially and 2–3 punctures on each side. Mesoscutellum prominent, medial length 0.15 mm, with deep punctures. Elytra (Fig. 2A) longer than wide, length 0.6 mm, width of elytron 0.23 mm with deep setiferous punctures least distance between punctures equal to 1 puncture. Legs (Figs. 2A–B) with tarsal formula 5-5-5 densely punctuated. Tarsomeres setiferous, 1–4 transverse, 5 elongate empodium appearing glabrous. Protibia with three rows of stout setae. Mesotibia with several rows of stout setae metatibia having stout setae apically only. Abdomen.—(Figs. 2A–B) with posterior margins of abdominal terga wider than anterior margins terga gradually becoming wider, ma imum abdominal width 0.46 mm. Abdominal terga and sterna with deep setifer-ous punctures punctures contiguous or separated by ma imum distance of 0.5 puncture diameter. Abdominal terga III–V with faint basal carina subbasal (arch-like) carina not present sterna VII and VIII with no obvi-ous secondary se ual modi cations sternum IX with a shallow U-shaped emargination aedeagus not visible. : Unknown.

Holotype.— , AMNH DR-10-1199, labeled “AMBER: Oligo-Miocene, Dominican Republic, AMNH no DR-10-1199” / “Staphylinidae, Staphylininae, Holisus, det. L. Herman” / “HOLOTYPE Holisus funeratus Chatzimano-lis and Engel” deposited in AMNH.

Etymology.—The speci c epithet is the Latin term fu-neratus, meaning “intern” or “bury,” and is a reference to this being a fossil species.

Subtribe Philonthina Kirby, 1837Genus Philonthus Stephens, 1829

Philonthus hades, new species(Figs. 3A–G)

Diagnosis.— The following characters, along with the geologic age of this species, can be used to distinguish this from other Philonthus species: head longer than wide, lat-eral margins of pronotum parallel-sided, disc of pronotum with four pairs of setiferous punctures, and metatarsi with

basitatsus subequal in length to last tarsomere. Compara-tive notes with P. rhadamanthus are given in the diagnosis of P. rhadamanthus below.

Description.— medium size, length 4.69 mm. Coloration.—(Fig. 3A): head, antenna, and pronotum black mouth-parts, mesoscutellum, elytra, and legs dark brown abdomen dark brown, although nearly black. Head.—(Fig. 3B) longer than wide, length 0.88 mm, width 0.63 mm. Head with microsculpture of thin, transverse, weakly wavy lines and im-punctate disc e cept distinct deep setiferous punctures posteriorly and laterally temple behind eye about as long as neck, distinctly shorter than compound eyes. Mandibles (Figs. 3D–G) long and sharp ma illa not visible e cept ma illary palpi. Ma illary palpus with palpomere 1 short palpomere 2 three times as long as palpomere 1 palpomere 3 shorter than 2, palpomere 3 widest apically palpomere 4 conical, narrower and about twice as long as 3. Labial palpomeres 1 and 2 not visible palpomere 3 parallel-sided, long and slender. Antenna (Figs. 3D, G) long, appro i-mately as long as head and pronotum combined antennomeres 1–6 lon-ger than wide antennomeres 7–10 subquadrate to quadrate antennomere 1 long and wide, 1.5 times as long as antennomere 2 antennomere 3 lon-ger than 2 antennomeres 4–5 subequal antennomeres 4–11 with dense setae. Head constricted posteriorly to form neck neck wider than long, with multiple micropunctures and microsculpture similar to head. Thorax.—Pronotum (Fig. 3E) longer than wide, length 0.73 mm, width 0.65 mm. Pronotum with four pairs of deep, setiferous punctures on disc and multiple punctures along the lateral and posterior margin pronotal microsculpturing as on head. Lateral margins of pronotum par-allel anterior border weakly concave posterior border weakly conve . Mesoscutellum large, length 0.29 mm, with numerous ne, sub-erect se-tae e cept basally such setae absent integument apparently imbricate. Elytra (Figs. 3B, D–F) slightly transverse, length 1.00 mm, width of ely-tron 0.57 mm. Elytra with numerous punctures and ne, sub-erect setae as on mesoscutellum integument apparently imbricate lateral margins parallel posterior margin straight. Hind wings (Fig. 3A) fully developed. Legs (Figs. 3A–B) with tarsal formula 5-5-5 long, meso- and metatibia and metatarsi with numerous long spines protarsi slightly dilated meta-tarsi with basitarsus subequal in length to last tarsomere. Abdomen.—Abdominal [no clear dorsal view] terga and sterna (Fig. 3C) appearing covered with dense setae. Abdominal terga III–V with two basal lines elevated area between these two lines appearing shining and with less punctures than rest of tergum. : Unknown.

Holotype.— AMNH DR-10-1216, labeled, “AMBER: Oligo-Miocene, Dominican Republic, AMNH no DR-10-1216” / “Dominican Republic, Santiago” / “Staphylinidae, Staphylininae, Staphylinini, det. L. Herman” / “Philonthina gen. sp. det. A. Newton 2010” / “HOLOTYPE Philonthus hades Chatzimanolis and Engel” deposited in AMNH.

Paratype.— AMNH DR-10-1215, labeled, “AMBER: Oligo-Miocene, Dominican Republic, AMNH no DR-10-1215” / “Dominican Republic, La Toca mine, Santiago” / “Staphylinidae, Staphylininae, Philonthus?, det. L. Her-man” / “PARATYPE Philonthus hades Chatzimanolis and Engel” deposited in AMNH.

Etymology.—The speci c epithet is the name of the Greek god of the underworld, Hades (meaning “the unseen”), and is treated as a noun in apposition.

Remarks.—The genus Philonthus includes more than 1000 species worldwide and it is likely not monophyletic, (Newton et al. 2001). Since the genus is not monophyletic


it makes little sense to try to distinguish this species from others, especially given the diversity of the group. In the absence of a comprehensive monograph and phylogenetic hypothesis for the genus, it is impossible at this time to speculate on the nearest relatives of the fossil species and biogeography, particularly over the span of millions of years. Speculation is of little value, because closely dis-tributed or overlapping species may still be distant rela-tives. It is greatly hoped that e tensive work on the modern species will move forward and one day permit a reconsid-eration of these fossil species.

Philonthus rhadamanthus, new species(Figs. 4A–G)

Diagnosis.—Philonthus rhadamanthus can be distin-guished from P. hades based on the following characters, besides the obvious difference in size: in P. hades the lateral margins of the pronotum are parallel, while in P. rhadaman-thus the lateral sides of the pronotum are parallel anteriorly and concave medially in P. hades the disc of the pronotum has four pairs of setiferous punctures and metatarsi with basitarsus subequal in length to the last tarsomere, while in P. rhadamanthus the disc of the pronotum has two pairs of setiferous punctures and metatarsi with basitarsus equal appro imately to 2/3 the length of the last tarsomere.

Description.— medium size, length 5.78 mm. Coloration.—(Figs. 4A, C) head, pronotum, and elytra dark brown to black legs dark brown. Head.—longer than wide, length 0.92 mm, width 0.75 mm, with mi-crosculpture of thin, transverse, weakly wavy lines with large setiferous punctures posteriorly and laterally, and few scattered punctures anteriorly around central disc temple behind eye about as long as neck, distinctly shorter than eye. Clypeus moderately emarginate. Labrum wide, as wide as frons, with long macrosetae. Mandibles (Figs. 4B, G) long, sharp ma -illa not visible e cept ma illary palpi. Ma illary palpus with palpomere 1 short palpomere 2 three times as long as palpomere 1 palpomere 3 short-er than 2, palpomere 3 widest apically palpomere 4, conical, narrower and about twice as long as 3. Labial palpomere 1 not visible palpomere 2 slender, longer than wide palpomere 3 parallel-sided, about twice as long as palpomere 3. Antenna (Fig. 4B) long, appro imately as long as head and pronotum combined antennomeres 1–5 longer than wide an-tennomeres 6–10 subquadrate to quadrate antennomere 1 long and wide, 1.5 times as long as antennomere 2 antennomere 2 widest distally anten-nomere 3 longer than 2 antennomeres 4–5 subequal antennomeres 5–11 (perhaps even 4) with dense setae. Head constricted posteriorly to form neck, neck wider than long, with multiple micropunctures and microscu-lpture similar to head. Thorax.—Pronotum (Figs. 4D, E) longer than wide, length 0.98 mm, width 0.75 mm. Lateral sides of pronotum parallel to each other anteriorly, concave medially posterior border of pronotum weakly conve . Lateral and anterior borders of pronotum with numerous deep setiferous punc-tures and long macrosetae disc of pronotum with two pairs of setiferous punctures. Pronotal microsculpturing as on head with additional micro-punctures. Mesoscutellum (Fig. 4E) large, length 0.44 mm, with numer-ous ne, sub-erect setae e cept basally such setae absent integument apparently imbricate. Elytra quadrate (Figs. 4D, E), length 1.08 mm, width of elytron 0.54 mm with numerous ne, sub-erect setae as on me-soscutellum integument apparently imbricate lateral margins parallel posterior margin straight. Hind wings fully developed. Legs (Figs. 4A–C) with tarsal formula 5-5-5 long meso- and metatibia with numerous spines protarsi slightly dilated metatarsi with basitarsus appro imately

2/3 length of last tarsomere. Abdomen.—(Fig. 4F) gradually becoming wider posteriorly (wid-est on segment V and VI) ma imum abdominal width 1.02 mm. Terga and sterna with multiple deep setiferous punctures. Abdominal terga III–V with two basal lines elevated area between these two lines appear-ing shining and with less punctures than rest of tergum. Lateral tergal sclerites of IX thin, long (as long as VIII), with large long setae. : Unknown.

Holotype.— AMNH DR-10-756, labeled, “AMBER: Oligo-Miocene, Dominican Republic, AMNH no DR-10-756” / “Staphylinidae, Staphylininae, Staphylinini, det. L. Herman” / “HOLOTYPE Philonthus rhadamanthus Chatzimanolis and Engel” deposited in AMNH.

Paratypes.—2 , FMNH, labeled, “DOMINICAN RE-PUBLIC: La Toca mine in the Northern, coastal range, elev. 1000, 75% probability from this area, Dominican amber, Field Museum Nat. Hist.” / “PARATYPE Philon-thus rhadamanthus Chatzimanolis and Engel” deposited in FMNH.

Etymology.—The speci c epithet is the name of the Greek mythological king, Rhadamanthys, son of Zeus and Europa, who purportedly ruled Crete before Minos and es-tablished its code of conduct. In mythology he eventually became the overseer of the Elysian Fields and sometimes judge of the deceased.

Subtribe Quediina Kraatz, 1857Genus Quedius Stephens, 1829

Quedius (Cyrtoquedius) electrodominicanus, new species(Figs. 5A–E)

Diagnosis.—Quedius electrodominicanus seems to be-long to the subgenus Cyrtoquedius Bernhauer based on the following characteristics [provided by Adam Brunke]: head, pronotum, and elytra with no microsculpture head with one puncture along inner margin of eye between ante-rior and posterior frontal punctures pronotum with single puncture on each side of midline elytra glabrous e cept setiferous punctures in rows and elytral epipleuron gla-brous e cept few macrosetae. Distinguishing this species from all other species of Cyrtoquedius is challenging since the subgenus has not been revised. However, based on the key provided by Scheerpeltz (1955), Q. electrodominica-nus appears to be the smallest species known (at least 2 mm shorter than all other species).

Description.— , medium size, length 5.47 mm. Coloration.—(Fig. 5A) head, pronotum and abdomen dark brown antennae, elytra and legs (e cept metaco a and metafemur dark brown) light brown. Head.—(Fig. 5D) quadrate, length 1.04 mm, width 1.00 mm. Head smooth, with no microsculpture, impunctate on disc e cept one punc-ture on inner margin of eyes between anterior and posterior punctures with few deep setiferous punctures along lateral border eyes large, about as long as distance between eyes. Gular sutures separated anteriorly but joined near middle of head. Clypeus slightly emarginate. Labrum wide,


with long macrosetae. Mandibles (Fig. 5D) long, sharp, each with medial tooth. Ma illa not visible e cept palpus ma illary palpus with palpomere 1 short palpomere 2 appro imately 2.5 times as long as palpomere 1 and wider palpomere 3 shorter and narrower than 2 palpomere 4 long, slender, conical, about twice as long as palpomere 3. Antennomeres 1–5 longer than wide antennomeres 5–6 subquadrate antennomeres 7–10

transverse [appearing distorted on Figs. 5A–D] antennomere 1 1.5 times as long as antennomere 2 antennomere 3 longer than either antenno-meres 2 or 4 antennomere 5 shorter than 4 antennomere 11 twice as long as antennomere 10. Thorax.—Pronotum (Figs. 5B–C) typical of Quediina, with pronotal hypomeron strongly in e ed, not visible in lateral view pronotum with

Fig. 4.—Photomicrographs of Philonthus rhadamanthus. A, lateral view of FMNH paratype, scale bar = 1.08 mm B, lateral view of FMNH paratype showing details of head, antenna, and legs, scale bar = 0.54 mm C, lateral view of holotype AMNH DR-10-756, scale bar = 1.08 mm D, dorsal view of holotype AMNH DR-10-756, scale bar = 0.92 mm E, dorsal view of holotype AMNH DR-10-756 showing details of head, pronotum, and elytra, scale bar = 0.68 mm F, dorsal view of holotype AMNH DR-10-756 showing details of abdomen, scale bar = 0.68 mm G, dorsal view of holotype AMNH DR-10-756 showing details of head, mouthparts, and legs, scale bar = 0.77 mm.


slight e planate lateral and posterior margins length 1.21 mm, width 1.25 mm. Pronotal central disc with single puncture on each side of mid-line, otherwise impunctate with no microsculpture. Mesoscutellum with setiferous punctures, length 0.42 mm. Elytra (Figs. 5B, C) transverse, length 1.38 mm width of elytron 0.79 mm appearing glabrous e cept for four rows of four setiferous punctures and border of elytron. Elytral

epipleuron glabrous e cept few macrosetae. Hind wings fully developed (Figs. 5C, E). Legs (Fig. 5E) with tarsal formula 5-5-5 long, tibiae with numerous large spines protarsi dilated. Abdomen.—with numerous long setiferous punctures covering sterna and terga pair of long macrosetae e tending beyond the posterolateral margins of terga III–VII ma imum abdominal width 1.13 mm. Distal

Fig. 5.—Photographs of Quedius (Cyrtoquedius) electrodominicanus (A–E) and Neoxantholinus apolithomenus (F–G). A, dorsal view of paratype AMNH DR-15-77, scale bar = 0.82 mm B, lateral view of paratype AMNH DR-15-77, scale bar = 1.3 mm C, lateral view of paratype AMNH DR-15-77, scale bar = 1.3 mm D, ventral view paratype AMNH DR-15-77 showing details of the head and mouthparts, scale bar = 0.82 mm E, lateral view of holotype AMNH DR-10-1211, scale bar = 1.38 mm F, dorsal view of the FMNH holotype, scale bar = 1.26 mm G, dorsal view of the FMNH holotype showing details of the head, pronotum and elytra, scale bar = 0.63 mm.


border of VIII with small emargination medially in males. : Unknown.

Holotype.— AMNH DR-10-1211, labeled, “AMBER: Oligo-Miocene, Dominican Republic, AMNH no DR-10-1211” / “Dominican Republic, Palo Tamboril, Santiago Prov.” / “Staphylinidae, Staphylininae, Staphylinini, det. L. Herman” / “HOLOTYPE Quedius electrodominicanus Chatzimanolis and Engel” deposited in AMNH.

Paratype.— AMNH DR-15-77 [poorly preserved], labeled, “AMBER: Oligo-Miocene, Dominican Repub-lic, AMNH no DR-15-77” / “Dominican Amber: Coleop-tera: Staphylinidae (large, e c.)” / “PARATYPE Quedius electrodominicanus Chatzimanolis and Engel” deposited in AMNH.

Etymology.—The speci c epithet is a combination of electron (Greek, meaning “amber”) and Dominican, refer-ring to the Dominican Republic.

Remarks.—Quedius electrodominicanus looks super -cially similar to some species in the genus Bolitogyrus however, the ne pubescence on the antennae starts on an-tennomere 4 (instead of 6 in Bolitogyrus), and the lateral margins of the pronotum are not as strongly e planate as in Bolitogyrus.

Tribe Xantholinini Erichson, 1839Neoxantholinus Cameron 1945

Neoxantholinus apolithomenus, new species(Figs. 5F–G)

Diagnosis.—Neoxantholinus apolithomenus can be dis-tinguished from all other species in Neoxantholinus based on the slightly asymmetrical mandibles and the punctation pattern of the pronotum: in N. apolithomenus the prono-tum has three dorsal rows of punctures, each with three punctures, while in other species the pronotum has less than three rows of punctures.

Description.— , small size, total length as preserved 3.63 mm. Coloration.—(Fig. 5F) integument throughout dark reddish brown, head appearing metallic, pronotum and elytra translucent, abdominal segments V–VII darker brown. Ventral view of head and pronotum ob-structed by bubble in amber. Head.— (Fig. 5G) longer than wide, length 0.67 mm, width 0.46 mm narrow, parallel-sided with rounded posterior angles. Head with long linear ocular grooves reaching almost middle of head. Head integument with long wavy longitudinal lines (perhaps preservational). Eyes small, positioned anteriorly, appro imately equal to length of head. Antenna (Fig. 5G) subequal in length to head antennomeres 1–4 longer than wide antennomere 1 equal in length to antennomeres 1–3 combined, anten-nomere 1 wider than antennomere 2 antennomere 2 wider than anten-nomere 3 antennomeres 3–4 subequal antennomeres 5–10 subquadrate, and similar in size antennomere 11 longer than wide, appro imately twice as long as antennomere 10. Mouthparts visible labrum with small medial emargination apically, with long setae. Mandibles sharp, with lat-eral furrow, slightly asymmetrical, both with medial tooth left mandible

slightly longer than right, tooth on left mandible slightly more elongate than tooth on right mandible. Ma illary palpus moderately elongate (about as long as mandibles) palpomere 1 not visible, palpomere 2 wider than palpomere 3 palpomere 3 elongate palpomere 4 much shorter and narrower than 3, conical. Labial palpomere 1 shorter and wider than pal-pomere 2 palpomere 3 narrower and slightly shorter than palpomere 2. Neck e tremely narrow, appro imately 1/8 of width of head. Thorax.—Pronotum (Fig. 5G) longer than wide, width 0.42 mm, me-dial length 0.63 mm pronotum widest anteriorly with three dorsal rows of punctures, each with three punctures. Superior marginal line of prono-tal hypomeron turning downward well behind middle of pronotum length and joining inferior line in front of proco ae. Mesoscutellum small, tri-angular, medial length 0.15 mm. Elytra longer than pronotum (Fig. 5G), length (as measured along lateral margin) 0.78 mm, width of elytron 0.31 mm, widest posteriorly with numerous punctures. Hind wings fully developed (Fig. 5F). Legs (Fig. 5G) with tarsal formula 5-5-5 densely punctated mesotarsi with lobed tarsomeres. Abdomen.—(Fig. 5F) densely setiferous, ma imum abdominal width 0.5 mm segments gradually becoming longer and wider. : Unknown.

Holotype.— , FMNH, labeled, “DOMINICAN REPUB-LIC: La Toca mine in the Northern, coastal range, elev. 1000, 75% probability from this area, Dominican amber, Field Museum Nat. Hist.” / “HOLOTYPE Neoxantholinus apolithomenus Chatzimanolis and Engel” deposited in FMNH.

Etymology.—The speci c epithet is derived from the Greek word apolithoma, meaning “fossil,” and is treated as a noun in apposition.

Remarks.—Poinar and Poinar (1999) mentioned the occurrence of Neoxantholinus in Dominican amber with-out formally describing a new species. In an earlier publi-cation, Poinar and Brodzinsky (1985) illustrated a antho-linine beetle in Dominican amber, but that specimen does not belong in Neoxantholinus.

DISCUSSION

It is rather surprising that the fossils described in this paper are the rst Staphylininae to be described from Dominican amber for two reasons: a) Dominican amber is rather com-mon and thousands of fossiliferous pieces are available and b) the recent fauna of the subfamily is very species-rich, and Chatzimanolis et al. (2012) and Solodovnikov et al. (2012) both hypothesized that the subfamily diversi ed during the Early Cretaceous. The species described above are probably just the tip of the iceberg regarding the diver-sity of staphylinines in Dominican amber and many more await description. This is evident in the many ta a men-tioned (but not described) by Seevers (1971), Poinar and Poinar (1999), and Irmler (2003). All species described here were keyed easily to their respective genus by using either the keys to Staphylini-nae in ‘American Beetles’ (Newton et al. 2001) and/or the keys in the ‘Illustrated guide to the genera of Staphylini-dae in Me ico’ (Navarrete-Heredia et al. 2002). Holisus funeratus is the rst fossil for the genus Holisus, a group of appro imately 30 species. Holisus has a primarily Neo-


tropical distribution with one species in Africa and an undescribed species from Arizona (Newton et al. 2001), while two species (H. laevis Blackwelder and H. guidingii Erichson) are known from the West Indies. Heterothops contains more than 150 species with a nearly worldwide distribution (Newton et al. 2001 and discussion about lim-its of Heterothops in Solodovnikov and Schomann 2009). Heterothops cornelli and H. infernalis are the second and third fossils described for the genus, after H. conticens Scudder (Scudder 1900), a compression from Florissant, Colorado. There are two Recent species of Heterothops known from the West Indies, H. ocularis Blackwelder and H. rambouseki Blackwelder. Quedius and Philonthus are both megadiverse gen-era with worldwide distributions and more than 800 and 1000 species, respectively. However, both of them are not monophyletic and some work is in progress to rede ne the ta onomic concepts used (Smetana 1995 Solodovnikov and Schomann 2009 Chatzimanolis et al. 2010). Several fossil species of Philonthus have been described mainly from the Oligocene-Eocene boundary of Florissant, Colo-rado (Scudder 1900), although the concept of Philonthus used by Scudder is probably rather different than the one used today. Fossils described as Quedius are mainly known from Florissant (Scudder 1890 Wickham 1912), the Oligocene of France (Oustalet 1874; Théobald 1937), and the Lower Cretaceous of China (Cai and Huang 2013) although similar to those records for Philonthus, these should be treated as tentative identi cations. E tensive Quedius-like fossils mostly assigned to a composite genus Cretoquedius Ryvnin are known from the Cretaceous pe-riod (Ryvkin 1988 Solodovnikov et al. 2012). There are several species of Philonthus from the West Indies, but to our knowledge this is the rst record of Quedius from that region. It is unclear if this is due to the e tinction of the ge-nus from the islands or under-sampling by modern collec-tors, although it is worth pointing out that the West Indies were sampled by at least two rove beetle systematists (A. Bierig and R. Blackwelder) in the 20th century. Neoxantholinus contains 21 species distributed in the Neotropical, Nearctic, and Australian regions. The ta o-nomic history of the genus was summarized by Smetana (1977), and all species studied by Blackwelder (1943) in the genus Oligolinus auct. (nec. Casey 1906) belong in fact in Neoxantholinus. Interestingly, the description of Steno-linus Bierig (Bierig 1937) from Cuba is remarkably similar to the description of Neoxantholinus and pending further e amination of specimens (no Stenolinus specimens were available), these two genera might require synonymy, with the name Stenolinus having priority. Blackwelder (1943) also stated that Stenolinus macrotrichus Bierig keyed to Oligolinus. The fossil described here is the rst fossil ta -on in Neoxantholinus, although Poinar and Poinar (1999) mentioned the occurrence of Neoxantholinus in Domini-can amber.

ACKNOWLEDGMENTS

We thank Lee Herman for his initial sort of most of the AMNH material, Alfred Newton for identi cation comments on Neoxantholinus, and the late James F. Cornell, Jr. for permitting us to study material from his private collection (now in the FMNH). Adam Brunke was e ceedingly helpful in describing and placing the Quedius specimens in the correct subgenus. Ismael A. Hinojosa-Díaz assisted with photomicrography. Par-tial nancial support was provided by U.S. National Science Founda-tion grants DEB-0741475 (to S.C. and M.S.E.) and DEB-0542909 (to M.S.E.). This is a contribution of the Division of Entomology, University of Kansas Natural History Museum.

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Documents

The Fauna of Staphylininae in Dominican Amber (Coleoptera: Staphylinidae)