Stuttgarter Beiträge zur Naturkunde

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1 Stuttgarter Beiträge zur Naturkunde Serie B (Geologie und Paläontologie) Herausgeber: Staatliches Museum für Naturkunde, Rosenstein 1, D Stuttgart Stuttgarter Beitr. Naturk. Ser. B Nr pp., 8 figs. Stuttgart, Osoriinae (Coleoptera: Staphylinidae) from Dominican Amber By Ulrich Irmler, Kiel With 8 Figures Abstract In total, 13 fossil specimens of Osoriinae in Dominican amber were investigated which are kept in the collections of the Staatliches Museum für Naturkunde Stuttgart. Two new extinct species, Thoracophorus palaeobrevicristatus n. sp. and Nacaeus dominicanensis n. sp., a new extinct genus and species Lispinomimus atavus n.g. n.sp., and further 5 indeterminate species of the genera Liberiana, Osoriellus, and Neosorius are described. Some extinct species are compared with extant species of the same region. Zusammenfassung Insgesamt 13 Exemplare von Osoriinae aus Dominikanischem Bernstein wurden untersucht, die sich im Staatlichen Museum für Naturkunde Stuttgart befinden. Zwei neue, ausgestorbene Arten, Thoracophorus palaeobrevicristatus n. sp. und Nacaeus dominicanensis n. sp., eine neue ausgestorbene Gattung und Art Lispinomimus atavus n.g. n.sp. und weitere 5 unbestimmte Arten aus den Gattungen Liberiana, Osoriellus und Neosorius werden beschrieben. Einige der ausgestorbenen Arten werden mit rezenten neotropischen Arten verglichen. Introduction Within the extensive collections of fossiliferous amber in the Staatliches Museum für Naturkunde Stuttgart, 13 pieces of Dominican amber exist with the same number of fossil Osoriinae specimens. At present, many fossil rove beetles of different genera are described from the Baltic amber, e.g. of Lathrobium (PASNIK & KUBISZ 2002), Adenopsis (ZERCHE 1999) and others. From the Mexican amber, too, originating from the Late Oligocene or Early Miocene, different species of the subfamilies Trichopseninae, Aleocharinae, and Paederinae are described by SEEVERS (1971). Although many species of Osoriinae are adaptated to trees or woody debris, only one species, Osorius brunnicornis HOPE, 1837, has been described thus far according to the catalogue of amber and copal beetles (SPAHR 1981). POINAR (1992) comprised the present knowledge on Dominican amber and its

2 2 stuttgarter beiträge zur naturkunde Ser. B, Nr. 342 fossils. According to POINAR s publication, Dominican amber originated from Late Eocene to Early or Mid Miocene, thus, being some 20 to 40 million years old. The amber producing tree was the fossil leguminous tree Hymenaea protera related to the extant Hymenaea verrucosa, at present occurring in East Africa and adjacent islands. Later, POINAR (1999) described an illustration of the Dominican amber forest which resembles much the present rain or cloud forests in the Neotropical region, while the Baltic amber forest is supposed to have been a coniferous forest with wetland conditions (HIEKE & PIETRZENIUK 1984). There are no more detailed informations on the mines where the investigated 13 pieces of amber were collected. Thus, it is not possible to give more exact information on the age of the Osoriinae species. Methods. Pictures were taken by a Wild (Heersbrugg) Makroskop M420 with Apozoom 1.6 and a digital camera. It was renounced to determine length or width of the insects because a shrinking process occurred during fossilisation in amber. Only total length was measured to give a rough idea of the beetle s size. The museum s identification numbers are indicated under Material of the descriptions. Acknowledgements Thanks are due Dr. G. Bechly of the Staatliches Museum für Naturkunde Stuttgart who put the 13 pieces of amber to my disposal. In particular, I thank Dr. Karin Wolf-Schwenninger, who excellently prepared the fossiliferous pieces so that it was easy to investigate the specimens. Descriptions of genera and species Order Coleoptera LINNAEUS, 1758 Family Staphylinidae LINNAEUS, 1758 Subfamily Osoriinae ERICHSON, 1839 Genus Thoracophorus MOTSCHULSKY, 1837 The genus Thoracophorus is most related to the genus Aneucamptus SHARP, It is differentiated from that genus by the absence of the prosternal process that divides the front coxae in Aneucamptus. Other related genera live in the southern part of South America, e.g. Rhopalopherus BERNHAUER, 1909 and Mesotrochus WAS- MANN, 1890, associated with ants and characterised by a subocular furrow and clavate antennae. Thoracophorus palaeobrevicristatus n. sp. Fig. 1A, 9 Material: 3 specimens; holotype with number DO-189-K, paratypes with numbers DO M, DO-2405-K. Etymology: The specific epithet is a combination from the Greek word palaios meaning old and brevicristatus referring to the high similarity to the extant species in the region. Diagnosis. The species can be hardly differentiated from the extant T. brevicristatus HORN, 1871 (see description in IRMLER 1985) which is distributed in Central America, West Indies and the southern USA (Fig. 9). The antennae seem to be slightly more slender in the extinct species and the elytra seem to be longer. In T. brevicristatus the antennomeres 7 to 10 are 2 to 3 times wider than long (Fig. 1B), while

3 irmler, osoriinae from dominican amber 3 A B C D Fig. 1. A: Thoracophorus palaeobrevicristatus n. sp. (DO-189-K) from Dominican amber and B: the extant species T. brevicristatus Horn, 1871 (Coll. American Museum of Natural History, New York). C: Nacaeus dominicanensis n. sp. from the Dominican amber in total view (Do-176-K) and D: the extant species Nacaeus planellus SHARP, 1887 (Coll. of the author).

4 4 stuttgarter beiträge zur naturkunde Ser. B, Nr. 342 they are almost not wider than long in T. palaeobrevicristatus. The extant species is also slightly longer with mm, but this may be also referred to drying effects by amber. Description. Length: 2.0mm. Colour: piceous. Head: broad, with wide lateral carinae that cover the eyes; thus, eyes not visible in dorsal view; with the typical two longitudinal prominences as in T. brevicristatus. Antennae: with long 2 nd antennomere, that is at least two times longer than wide; the following 5 antennomeres more or less quadrate; antennomeres 9 and 10 distinctly wider than long; the last three antennomeres distinctly thicker than the preceding antennomeres. Pronotum: sides parallel, strongly narrowed shortly in front of the posterior angles; margin denticulate; with distinct lateral furrow; on the disc with flat depression, which is divided by an indistinct circular prominence; surface with distinct isodiametrical reticulation, dull. Elytra: with similar micro-reticulation as on the pronotum; surface dull, with the typical 3 longitudinal carinae of the genus; it seems that a punctuation exists consisting of large punctures, but this cannot be definitely identified; the structure may be also referred to small air bubbles. Genus Nacaeus BLACKWELDER, 1942 The genus Nacaeus is characterised by the absence of morphological structures that are developed in the related genera (IRMLER in press). The head is orbicular with eyes not prominent like in the genus Tannea BLACKWELDER, Strigae on abdominal sternites like in Neolosus BLACKWELDER, 1942 and Lispinus ERICHSON, 1840 are also not developed. Nacaeus dominicanensis n. sp. Fig. 1C Material: 3 specimens; holotype with number Do-176-K; paratypes with numbers Do- 250-K, Do-1852-Ü. Etymology: The specific name refers to the state of the Dominican Republic, where the species was found in amber. Diagnosis. The species is distinctly dorsoventrally depressed, similar as the present N. planellus (SHARP, 1887) (Fig. 2B). The colour of the three specimens varies between brown and dark yellow. The amber species is darker than the extant N. planellus and the microsculpture is more distinct. But it seems that the surface is also polished and shiny. Description. Length: 1.8 2mm. Colour: unicoloured, brown to dark yellow. Head: more or less circular; eyes not prominent; punctuation seems to be sparse and fine and surface more or less shiny; setation seems also to be like in the present Nacaeus species, with two supraocular setae, an additional seta near the anterior supraocular seta and two setae at the anterior margin. Antennae: with relatively long first two antennomeres that are two times longer than wide; antennomeres 4 to 6 more or less quadrate and the following ones wider than long. Pronotum: sides arcuate in the anterior part and distinctly emarginate in the posterior part; at the posterior angles with distinct depressions; with coarser punctuation than on the head, in particular near the posterior margin; on the disc with finer and sparser punctuation; ground sculpture sparse and fine, longitudinally reticulate; surface more or less shiny; with distinct midline and an indistinct longitudinal de-

5 irmler, osoriinae from dominican amber 5 pression on both sides of the midline. Elytra: one third longer than the pronotum; punctuation weak and sparse; with distinct ground sculpture, that is longitudinally or netlike reticulate; surface shiny; in the centre of the disc with a long seta inserted in a large puncture. Abdomen: with the typical structure of present Nacaeus species; very finely punctuate and with the rhomboid ground sculpture. Genus Liberiana BLACKWELDER, 1942 Within the related genera, Lispinus, Neolosus, Nacaeus and Tannea, the genus Liberiana is characterised by a coarsely punctuate abdomen, the absence of diagonal strigae at abdominal sternites and of the raised line on the hypomeron (see key of the Neotropical genera of the subtribe Lispinina in IRMLER, in press). Liberiana sp. Fig. 2A, B Material: Do-1410-K, (Do-206-K?). Main characters. The species shows the characteristics of the genus concerning abdominal punctuation, absence of abdominal strigae and prosternal suture. The extinct species is still more similar to the genus Lispinus than the extant Liberiana auratus (Fig. 2C), that is the only known species of the genus Liberiana from Central and South America. The sides of the pronotum of the amber species are also more or less parallel and punctuation is dense and relatively coarse, while in the extant L. auratus pronotal sides are shortly narrowed near the posterior angles. Both species have distinct longitudinal depressions at posterior angles. Description. Length: 3.0mm. Colour: black, antennae and legs reddish. Head: with parallel sides, eyes as long as temples; with distinct, but much weaker and sparser punctuation than on pronotum and elytra; without ground sculpture, surface polished and shiny. Antennae: short, with 2 nd antennomere globular, 3 rd conical and at least twice as long as 2 nd ; the following three antennomeres quadrate and small; antennomeres 7 to 10 larger than the preceding ones and wider than long. Pronotum: with nearly parallel sides; very weakly and straightly narrowed to posterior angles; shortly arcuate at anterior angles; with dense and coarse punctuation; distance between punctures shorter or as long as diameter of punctures; without ground sculpture; surface more or less polished and shiny; at posterior angles with deep longitudinal depressions. Elytra: as wide and long as the pronotum; also punctuation and ground sculpture similar as on the pronotum. Abdomen: tergites with still coarser and denser punctuation than on pronotum and elytra and with distinct more or less longitudinally reticulate ground sculpture, which is similar on sternites; last abdominal sternites with still denser microsculpture than tergites and hairy. Remarks. Within the collection exists another Liberiana species labelled Do- 206-K. This species cannot be described as thoroughly as the preceding, because of inclusions that prevent a distinct view on the punctuation and ground sculpture of pronotum and elytra. However, a coriaceous ground sculpture and finer punctuation seem to exist on the elytra. Also the abdomen appears to have a much weaker and sparser punctuation, but a distinctly denser microsculpture. No diagonal strigae on abdominal sternites could be detected.

6 6 stuttgarter beiträge zur naturkunde Ser. B, Nr. 342 A B C Fig. 2. Liberiana sp. (Do-1410-K) from Dominican amber in dorsal (A) and lateral (B) views compared with the extant species Liberiana auratus (IRMLER, 1994) (C; Coll. of the author).

7 Genus Osoriellus FAGEL, 1959 The complex of osorian species of the Neotropical region is still badly known and a detailed revision is urgently necessary. Actually, no Osoriellus species is described from the Neotropical region according to HERMAN 2001, but obviously the genus is represented by a lot of species in the region, because more than 10 species are in my collections from Central and South Brazil. The genus is differentiated from other osorian genera by the structure of the front tibia. Front tibiae in Osoriellus have a smooth outer margin with spines directly inserted on the outer margin. Usually the front tibia is wide forming a more or less large plate, which contains no hairs at the inner side, while the outer side is densely covered by thick and long hairs. In my collections there are species with extraordinary large front tibiae that form circular plates covering nearly totally the usual emargination of the ventral margin. In contrast, another species has very slender front tibiae similar as in Holotrochus. Size of species in my collections varies between 2.4 mm and 6 mm and eyes can be as large as temples or much smaller. Material: Do-326-K, Do-381-K. irmler, osoriinae from dominican amber 7 Osoriellus sp. Fig. 3, 7E Main characters. This species was found in the amber collections by two specimens. It certainly represents the genus Osoriellus FAGEL, 1959 described from the African region. The amber species represents a common type with front tibiae forming a slender oval plate, with a distinct emargination at the ventral margin, distinct hairy outer side, and well developed eyes. Description. Length: 2.6mm. Colour: seems to be black. Head: oval, with well developed eyes, as long as temples; length in front of eyes slightly longer than diameter of eyes; front margin without teeth or prominence; labrum transverse, twice as long as wide, with several long setae at front margin; surface seems to be polished without ground sculpture; punctuation distinct and moderately dense; distance between punctures as wide as diameter of puncture; each puncture with a short yellow seta. Antennae: 1 st antennomere long; the followings inserted in a rectangular angle to the first; antennomeres 2 6 short, more or less quadrate; 7 10 wider than long forming an indistinct club. Pronotum: trapezoid; widest at anterior angles; straightly narrowed to posterior angles; punctuation similar as on the head and each puncture with a short yellow seta; surface seems also to be polished. Elytra: slightly longer than pronotum; more or less quadrate; surface seems to be coriaceously reticulate and punctures coarser than on the pronotum; punctuation denser than on the pronotum and setae seem to be shorter. Abdomen: hardly to investigate, because it is covered by the hind wings; punctuation seems to be coarse and dense and also hairy. Legs (Fig. 7E): front tibia seems to be relatively slender and much longer than wide; emargination at ventral margin distinct; the number of spines at outer margin cannot be definitely seen, it may be 7 or 8; inner side of the tibia smooth without spines or hairs, outer side distinctly and densely hairy.

8 8 stuttgarter beiträge zur naturkunde Ser. B, Nr. 342 Fig. 3. Osoriellus sp. in dorsal view on front body of Do-326-K and view on outer side of front tibia of Do-381-K (right). Genus Neosorius FAGEL, 1959 The genus Neosorius FAGEL, 1959 is presently not recorded from the Neotropical region. The reasons are already discussed for the preceding species. The genus, however, is also certainly represented in Central and South America. I have several species of the genus in my collections. The genus was described from the African region with many species. According to FAGEL (1959) the species of the genus are characterised by a small to medium size and the anterior tibia with spines inserting on more or less long digits. In his key to the African genera of the Osorius group, FAGEL (1959) did not mention the hairy abdomen as differentiation to the genus Osorius. In my collections the species of the genus Neosorius have a more or less densely haired abdomen, while in the genus Osorius the abdomen is not hairy. The present two species in the Dominican amber are conspicuous by very long spines of the front tibia and relatively long mandibles. The abdomen is also hairy as in the other Neotropical Neosorius species of my collections. In contrast to the present Neotropical Neosorius species of my collections, which have a transverse rectangular labrum, the two amber species have a large prominently arcuate labrum. In the present species of the genus Neosorius or of the genera Osorius, Osoriocanthus, and Osoriellus a similarly structured labrum is not found. Nevertheless, I hesitated to create a new genus for two amber species, because the group is badly investigated and representatives of this type might be found also in the present fauna. Material: Do-631-K. Neosorius sp. 1 Fig. 4, 7A, C Main characters. This species is differentiated from the following by the slightly shorter size and, in particular, by the shorter elytra and eyes. The colour is darker and the antennae seem to be slightly longer than in the following species. It cannot be definitely decided, if the spines of the front tibia are longer because the front tibia is hardly to investigate in the following species.

9 irmler, osoriinae from dominican amber 9 Fig. 4. Neosorius sp. 1 (Do-631-K) in dorsal view (left) and enlarged view of front tibia (right). Description. Length: 2.2mm. Colour: seem to be between black and dark reddish. Head: more or less quadrate and wider than the pronotum, sides parallel, the base of antennae covered by a prominent plate; labrum arcuately prominent and with several long setae at the anterior margin (Fig. 7C); mandibles long, with a distal and two central acute teeth; maxillae very long, thin, and acute at the top; eyes cannot be exactly investigated, but they seem to be relatively small, it is also not clear whether the surface is sculptured or punctuate. Antennae: with long basal antennomere, 2 nd antennomere longer than the followings; 3 rd still longer than wide, while the following antennomeres are quadrate, antennomeres 5 and 6 smaller than the preceding ones and the following ones. Pronotum: widest near the anterior edge, distinctly and straightly narrowed to the posterior edge; with several setae at the lateral margin; punctuation and ground sculpture of surface cannot be exactly investigated. Elytra: As long as pronotum; as on head and pronotum the elytral surface cannot be exactly studied. Abdomen: anterior abdominal segments scarcely smaller than elytra; last abdominal segments much wider and thicker than the anterior segments; surface covered by hairs. Legs (Fig. 7A): front tibia with long spines, the longest one, nearly as long as width of the tibia; at the anterior edge with two long spines followed by a distinctly shorter one, the following marginal spines separated from these three anterior spines by a distinctly larger gap; while on the dorsal side of the tibia seem to exist no hairs, the ventral side is densely covered by long hairs. Material: Do-488-K. Neosorius sp. 2 Fig. 5, 7B, D Main characters. This species is mainly differing from the preceding one by the longer elytra and the reddish colour. While in the preceding species elytra

10 10 stuttgarter beiträge zur naturkunde Ser. B, Nr. 342 Fig. 5. Neosorius sp. 2 (Do-488-K) in dorsal view. are not longer than the pronotum, elytra are 1/3 longer than the pronotum in species 2. Description. Length: 2.7mm. Colour: reddish, pronotum slightly lighter red than the head and the elytra. Legs yellow. Head: punctuation seem to be sparse and fine and surface without distinct microsculpture, shiny; sides parallel from base of antennae to the neck, with a distinct prominence at the antennal base, sides of clypeus emarginate to the front edge, which is straightly obtuse; labrum arcuately prominent, with 6 long setae at each side of the middle (Fig. 7D); eyes large and distinctly prominent. Antennae: 2 nd antennomere oblong, much longer than wide and thicker than antennomeres 3 to 5; 3 rd antennomere conical, the following three more or less quadrate; antennomeres 7 to 11 distinctly thicker than the preceding ones, slightly wider than long. Pronotum: widest at the anterior edge, straightly narrowed to the posterior edge; punctuation and microsculpture seem to be similar as on the head, surface shiny. Elytra: conspicuously long, nearly 1/3 longer than pronotum and 1/4 longer than wide; punctuation seem to be coarser but as sparse than on the pronotum; ground sculpture indistinctly coriaceous, surface less shiny than on pronotum; with deep sutural furrow. Abdomen: last abdominal segments wider than basal ones; thus, abdomen posteriorly thicker than anteriorly; surface is covered by the hind wings and, therefore, scarcely visible; it seems to be sparsely hairy. Legs: front tibia is hardly to investigate; a ventral emargination with a comb of setae certainly exists; the outer margin carries several spines inserting on short teeth; a larger distance between the two distal spines and the following row of spines as in the preceding species seems not to exist; the spines seem also to be

11 Diagnosis. The species is differentiated by the characters of the genus from similar species of the genera Liberiana, Nacaeus and Tannea. Description. Length: 2.2mm. Colour: black, legs piceous; colour of pronotum cannot be identified as it is covered by an airy film, which makes it shining white. Head: densely and coarsely punctuate; distance between punctures much shorter than diameter of punctures, surface seem to be more or less shiny without microsculpture; eyes large and laterally prominent, scarcely shorter than head; 3 rd segment of maxillary palpus thick, while 4 th is very thin, needle like. Antennae: as long as head and pronotum; antennomeres 2 to 3 longer than wide, thin, 4 to 6 quadrate, 7 th antennomere seem to be thicker than the small 8 th, which is as thick as 6 th, antennomeres 9 to 10 slightly wider than long. Pronotum: surface of pronotum is covered by an airy film, which prevents to identify punctuation and microsculpture; widest at anterior edge and straightly narrowed to posterior angles; posterior angles and, in particular, anterior angles widely arcuate; sides with several dark setae. Elytra: distinctly longer than pronotum with coarse and moderately dense punctuation; microsculpture seem to be weak and surface, therefore, moderately shiny; laterally with a row of dark setae. Abdomen: with similar punctuation as on the elytra, coarse and moderately dense; a weak, isodiametrically reticuirmler, osoriinae from dominican amber 11 shorter; dorsal side of tibia seems to be without hairs or only with few hairs, while the ventral side is densely hairy. Genus Lispinomimus n. g. Etymology: The genus name refers to the high similarity with the genus Lispinus and derived from the combination of Lispinus and mimeo, the Greek word for imitating. Diagnosis. The new genus differs from the genera related to Lispinus by the absence of strigae on the abdominal sternites. Concerning size and the globular head with relatively large eyes it is similar to the genus Tannea. Species of this genus are characterised by a specific microsculpture and a very fine nearly invisible punctuation of the abdominal segments, while abdominal sternites of the new genus are coarsely punctuate. In this respect, it resembles the genus Liberiana. However, species of the genus Liberiana are larger and have a more parallel aspect. Head and abdomen are more or less parallel and only narrowed at the extreme distal end like in the genus Lispinus. In contrast, Lispinomimus has a more or less globular head with a distinct neck and a straightly narrowed abdomen from the base to the last segment. With respect to the globular head and the neck, it resembles the Hawaiian genus Lispinodes SHARP, Lispinodes, however, is characterised by obscure styles at the terminal abdominal segment, while in Lispinomimus no processes exist. I saw similar species from the South American continent deposited in the Canadian National Collections, Ottawa. But these species were only found in female specimens, which makes a description not appropriate. Moreover, they were all characterised by different abdominal processes like in Lispinodes. Lispinomimus atavus n. g. n. sp. Fig. 6 Material: holotype with number Do-449-K. Etymology: The specific name is derived from the Latin word atavus [substantive] meaning ancestor.

12 12 stuttgarter beiträge zur naturkunde Ser. B, Nr. 342 Fig. 6. Lispinomimus atavus n. g. n. sp. (Do-449-K) in dorsal view (right) and head in separate view from the other side of the diaphragmatic part (left). late microsculpture may exist, surface only moderately shiny; abdominal segments are relatively long, on average longer than wide; in total, the abdomen gives a relatively long and thin impression becoming successively thinner posteriorly; sides with very long dark setae; sternites without diagonal strigae. Discussion According to our present knowledge, Coleoptera developed in the Permian period (HENNIG 1969) and the oldest Staphyliniformia existed already in the Middle Jurassic (HENNIG 1969, ARNOL DI et al. 1991). The oldest staphylinids in amber are described from the Cretaceous New Jersey amber representing the subfamily Tachyporinae some 90 million years ago (GUSAROV 2000). The subfamily Osoriinae is regarded as the sister group of either the piestine subfamily (NEWTON & THAYER 1992) or the oxyteline subfamily (BEUTEL & MOLDENA 1997). The last subfamily is already reported from Jurassic fossils reflecting that the two subfamilies may exist at least since 150 million years (HERMAN 1986). Dominican amber dates from upper Eocene to lower Miocene meaning that the inclusions are between 40 and 20 million years old. DONNELLY (1988) reported a brief compendium on the development of the Greater Antilles with reference to the biogeographic implications. At the end of Cretaceous, fragments of the Greater Antillean islands developed, while between 38 and 15 million years ago Central America closed against the Pacific Ocean. DONNELLY (1988) emphasised that the Dominican island already existed more or less in a similar position as today and isolations between the Antillean islands began to establish. The extant Thoracophorus brevicristatus seems to represent a very persistent species of the ancient amber forest. As fig. 9 indicates, T. brevicristatus is still distributed in the northern part of the Antilles as may be its very similar ancestor T. palaeobrevicristatus. The most northern collecting location is situated in Arkansas (USA), the most southern on the St. Croix island. Thus, T. brevicristatus may occur

13 irmler, osoriinae from dominican amber 13 Fig. 7. Front legs of A: Neosorius sp. 1 (Do-631-K), B: Neosorius sp. 2 (Do-488-K), E: Osoriellus sp. (Do-381-K); labrum of C: Neosorius sp. 1 (Do-631-K) and D: Neosorius sp. 2 (Do-488-K). Scales: 0.1 mm. in convenient habitats of the whole area from southern USA to northern Panama in Central America and the Greater Antilles to St. Croix island. No collecting locations are known, however, from the South American mainland and the southernmost Lower Antilles, e.g. the islands of the Grenades or Trinidad/Tobago, although a lot of material of the genus was determined from that region. T. brevicristatus is actually not known from the Dominican Republic or Haiti, which can be referred to the bad collecting status or the devastated situation of the rain forest on that island. It is surprising that the actually frequent species of the genus Lispinus were not represented in the Dominican amber, while two species of the rare genus Liberiana were found. These two extinct species are, moreover, very similar to the extant genus Lispinus differing only by the absence of the ventral strigae of the abdominal sternites. This may indicate that Liberiana may be the ancestor of Lispinus. In this case, this old genus is only represented by the relic species Liberiana auratus, while the genera Lispinus, Neolosus, and Allotrochus with abdominal strigae have developed from that ancient genus. Regarding the genus Neosorius, a very decisive change may concern the structure of the labrum. In my collections, the extant species of the genus from the Neotropics have a transverse labrum that is characterised by a straight anterior edge. The two species from the amber forest have a semicircular anterior edge. I never investigated

14 14 stuttgarter beiträge zur naturkunde Ser. B, Nr. 342 Fig. 8. Distribution of the extant Thoracophorus brevicristatus with collecting locations in North and Central America. According to BLACKWELDER (1943), IRMLER (1985), and unpublished material in different collections, mainly in USA. African specimens of the genus and the Neotropical species are badly studied. Nevertheless, the ancient Neosorius from the Dominican amber may also represent a new extinct genus. Species of the subfamily Osoriinae are living in different habitats of the recent rain forest. Few, e.g. also some species of the genus Thoracophorus, the related dirrhocephalid genera, and Mesotrochus, are associated with ants (BURAKOWSKI & NEW- TON 1992). The extant Thoracophorus brevicristatus was found in colonies of the termites Neotermes castaneus (BURM.) and N. jouteli (BANKS & SNYDER) in Cuba (BO- HÁC 1978), which may indicate that its ancestor might also lived together with termites in the amber forest. Most species and genera, however, are living in the litter, under bark or in rotten logs of the rain forest and are, therefore, closely related to the rain forest trees. This also concerns the Osoriinae genera found in the Dominican amber. Most species of the genus Thoracophorus are found in the litter feeding mainly on fungi as well as species of the genus Nacaeus that are living under bark or also in leaf litter. Species of Osoriellus and Neosorius mainly occur under bark or even in the log as predators of wood feeding insects or as feeders on fungi. References ARNOL DI, L.V., ZHERIKHIN, V.V., NIKITIN, L.M., PONOMARENKO, A.G. (1991): Mesozoic Coleoptera. Translated from the Russian for the Smithsonian Institution Libraries. 284 pp.; New Delhi (Amerind Publishing Co).

15 irmler, osoriinae from dominican amber 15 BEUTEL, R.G. & MOLDENA, R. (1997): Comparative morphology of selected larvae of Staphylinoidea (Coleoptera, Polyphaga) with phylogenetic implications. Zoologischer Anzeiger, 236: 37 67; Jena. BLACKWELDER, R.E. (1942): Notes on the classification of the staphylinid beetles of the groups Lispini and Osoriinae. Proceedings of the United States National Museum, 91: 75 90; Washington. (1943): Monograph of the West Indian beetles of the family Staphylinidae. Smithsonian Institution United States National Museum Bulletin, 182: 1-658; New York. BOHÁC, J. (1978): Description of the larvae and pupa of Thoracophorus brevicristatus (Coleoptera, Staphylinidae). Acta Entomologica Bohemoslovaca, 75: ; Praha. BURAKOWSKI, B. & NEWTON, A.F. (1992): The immature stages and bionomics of the myrmecophile Thoracophorus corticinus Motschulsky, and placement of the genus (Coleoptera, Staphylinidae, Osoriinae). Annali del Museo Civico di Storia Naturale G. Doria, 89: 17 42; Genova. DONELLY, T.W. (1988): Geologic constraints on Caribbean biogeography. In: LIEBHERR, J.K. (ed.): Zoogeography of Caribbean insects. Pp ; London, Ithaca (Cornell University Press). ERICHSON, W.F. (1839): Genera et species Staphylinorum Insectorum Coleopterorum Familiae. 954 pp.; Berlin (Morin). FAGEL, G. (1959): Osoriinae (Coleoptera: Polyphaga, Fam. Staphylinidae). Exploration do Parc National de la Garamba. Mission H. de Saeger, 12: 1 205; Bruxelles. GUSAROV, V.I. (2000): Mesotachyporus puer, a new genus and species of Cretaceous Tachyporinae (Coleoptera, Staphylinidae) from New Jersey amber. In: GRIMALDI, D. (ed.): Studies on fossils in amber, with particular reference to Cretaceous of New Jersey. Pp London (Backhurys Publishers). HENNIG, W. (1969): Die Stammesgeschichte der Insekten. 436 pp.; Frankfurt (Kramer). HERMAN, L.H. (1986): Revision of Bledius. Part IV. Classification of species groups, phylogeny, natural history, and catalogue (Coleoptera, Staphylinidae, Oxytelinae). Bulletin of the American Museum of Natural History, 184: 1 367; New York. (2000): Catalog of the Staphylinidae (Insecta: Coleoptera) to the end of the second millenium. Vol. III. Oxyteline group. Bulletin of the American Museum of Natural History, 265: ; New York. HIEKE, F. & PIETRZENIUK, E. (1984): Die Bernstein-Käfer des Museums für Naturkunde, Berlin (Insecta, Coleoptera). Mitteilungen aus dem zoologischen Museum in Berlin, 60: ; Berlin. HOPE, F.W. (1837): X. Observations on succinic Insects. Part 2 nd Gums and resins. Transactions of the Royal entomological Society of London, 1 st Series, 2: ; London. HORN, G. (1971): Description of new Coleoptera of the United States, with notes on known species. Transactions of the American Entomological Society, 3: ; Philadelphia. IRMLER, U. (1985): Neue Arten der Gattungen Aneucamptus und Thoracophorus (Col., Staphylinidae) aus der Neotropis. Entomologische Blätter, 81: 41 58; Krefeld. (in press): Taxonomy and distribution of the Neotropical species of the genera Tannea BLACKWELDER, 1952 and Nacaeus BLACKWELDER, 1942 with remarks to genus Lispinus (Coleoptera: Staphylinidae). Bulletin de l institut royal des sciences naturelles de Belgique; Bruxelles. LINNAEUS, C. (1758): Systema Naturae. Regnum Animale. 824 pp.; Holmiae (Laurentii Salvii). MOTSCHULSKY, V. (1837): Extrait d une lettre adressée por M.V. Motschulsky à M.B. Zoubkoff. Bulletin de la Société Impériale des Naturalistes de Moscou, 10: ; Moscow. NEWTON, A. & THAYER, M. (1992): Current classification and family-group names in Staphyliniformia (Coleoptera). Fieldiana: Zoology (N.S.), 67: 1 92; Chicago. PASNIK, G. & KUBISZ, D. (2002): A new genus and new species of Staphylinidae (Coleoptera) from Baltic amber. European Journal of Entomology, 99: ; Budejovice. POINAR, G.O. (1992): Life in amber. 350 pp.; Stanford (Stanford University Press). POINAR, G.O. & POINAR, R. (1999): The amber forest. A reconstruction of a vanished world. 196 pp.; New Jersey (Princton University Press).

16 16 stuttgarter beiträge zur naturkunde Ser. B, Nr. 342 SEEVERS, C.H. (1971): Fossil Staphylinidae in Tertiary Mexican amber (Coleoptera). In: PE- TRUNKEVITCH, A. (ed.): Studies of fossiliferous amber arthropods of Chiapas, Mexico. Part II. Pp Berkley, Los Angeles, London (University of California Press). SHARP, D. (1880): On some Coleoptera from the Hawaiian Islands. Transactions of the Royal entomological Society of London, 1880: 37 54, London. (1887): Staphylinidae. In: GODMAN, F.D. (ed.): Biologia Centrali-Americana, Insecta, Coleoptera, 1. Pp ; London (Taylor & Francis). SPAHR, U. (1981): Systematischer Katalog der Bernstein- und Kopal-Käfer (Coleoptera). Stuttgarter Beiträge zur Naturkunde, B, 80: 1 107; Stuttgart. ZERCHE, L. (1999): Eine neue Art der Gattung Adinopsis CAMERON aus dem Baltischen Bernstein. Beiträge zur Entomologie, 49: ; Eberswalde. Author s address Prof. Dr. Ulrich Irmler, Christian-Albrechts Universität, Ökologie-Zentrum, Olshausenstr. 40, D Kiel uirmler@ecology.uni-kiel.de ISSN Schriftleitung: Dr. Gert Bloos, Rosenstein 1, D Stuttgart Gesamtherstellung: Gulde-Druck GmbH, D Tübingen