New fossil paussids from Dominican amber with notes on the phylogenetic systematics of the paussine complex (Coleoptera: Carabidae)

Systematic Entomology(1997)22, 345–362

New fossil paussids from Dominican amber withnotes on the phylogenetic systematics of thepaussine complex (Coleoptera: Carabidae)

P E T E R N A G E L Institut fur Natur-, Landschafts-, Umweltschutz, Biogeographie, Universita¨t, Basel,

Switzerland

Abstract. The first fossil representative of the Protopaussinae and the secondfossil Eohomopterus(Paussinae) are described from amber inclusions of Tertiaryage collected in the Dominican Republic on the West Indian island of Hispaniola(Protopaussus pristinussp.n., Eohomopterus poinarisp.n.). These finds increasethe total number of paussids present in Dominican amber to four species in threegenera (Protopaussus, Eohomopterus, Homopterus). A key is provided for theidentification of the New World genera of Protopaussinae and Paussinae and forthe species ofEohomopterus. Based on synapomorphic character states, the paussidbeetles are split into two subfamilies, the monobasic Protopaussinae and the moreinclusive Paussinae (the twenty-three genera are placed into two supertribes: theCarabidomemnitae comprising two tribes and the Paussitae comprising eight tribes.The extant congeners ofEohomopterusand Homopteruslive in the NeotropicalRegion, while the extant members ofProtopaussusare known only from theOriental Region. This record of New World TertiaryProtopaussusadds anotherexample of the Old World relationships of taxa from Dominican amber. Thetemporal and geographical distribution ofProtopaussusindicates that its presentzoogeographical range may represent the relict of a wide Laurasian distribution inthe past. It does not support the previously suspected Oriental centre of origin forthe ancestral stock of {Protopaussinae1 Paussinae}.

Introduction

Paussid (or ant nest) beetles are highly specialized,myrmecophilous carabids. Recent comprehensive treatmentsinclude Darlington (1950), Nagel (1987a) and Luna de Carvalho(1989). The number of extant species has been estimated at460 (Darlington, 1950), 500 (Nagel, 1987a) or 600 (Luna deCarvalho, 1983) and several fossil species have been recordedfrom Tertiary Baltic and Dominican ambers (twenty-six andtwo, respectively) (Wasmann, 1929; Nagel, 1987b).

This paper reports on two ant nest beetles in Dominicanamber which appear to be extremely interesting in structureand with regard to their zoogeographical implications. Thespecimen described below as a new species of the genusEohomopterusis conspicuous because of its unusual tibial

Correspondence: Prof. Dr Peter Nagel, Institut fu¨r Natur-,Landschafts- und Umweltschutz, Biogeographie, Universita¨t Basel,St.Johanns-Vorstadt 10, CH-4056 Basel, Switzerland.

© 1997 Blackwell Science Ltd 345

shape and unlike the other specimen is aProtopaussusspecies,for which no fossil records had been available and whichwould not initially have been expected in New World amberdeposits given its extant distribution restricted to the OrientalRegion.

A proper treatment of the newly discovered taxa requiredthe updating of records of fossil paussids since their lastcomprehensive treatment (Nagel, 1987b) and a considerationof the classification of the possibly relevant genera. As somephylogenetic conclusions drawn by Darlington (1950) had tobe revised, because of new discoveries and interpretations ofcharacter states (Nagel, 1987a) and as different opinions onthe genus and tribe level classifications of the paussine lineageexist (Darlington, 1950; Nagel, 1987a; Luna de Carvalho,1989), it was necessary to consider this phylogeny in moredetail. Similarly, the formal ranking of the taxa belongingto the ozaenine–protopaussine–paussine lineage had to bereconsidered.

The surprising discovery of the extant strictly OrientalgenusProtopaussusin New World amber deposits required a

346 Peter Nagel

reconsideration of previous biogeographical hypotheses on thedistribution history of paussids. There are a few similar, yetnot identical, distribution patterns among Dominican amberinsects (Poinar, 1992; Baroni Urbani, 1995) and such OldWorld affinities require particular attention.

The main purposes of the present paper are, thus, to contributeto the knowledge of fossil paussid beetles, to substantiatethe applied classification of paussid beetles and to discussbiogeographical hypotheses concerning the distribution historyof Protopaussus.

Fossil paussids

The first comprehensive description of Paussinae in Balticamber was given by Wasmann (1929) and the first inclusionsin Dominican amber were mentioned by Wunderlich (1986)and described by Nagel (1987b). The last reference contains alist of all fossil paussids. A phylogenetic classification ofthe fossils was proposed by Nagel (1987a) and Luna deCarvalho (1989).

Further references to fossil Paussinae not listed by Spahr(1981a,b), Keilbach (1982) or Nagel (1987b) are givenimmediately below: Heyne & Taschenberg (1908), Wheeler(1910) and Wasmann (1934) mentioned fossil paussids. Hieke& Pietrzeniuk (1984) provided information on the collectionof amber paussids at the Museum fu¨r Naturkunde, Berlin.

Schumann & Wendt (1989) reported ‘Paussinae’ in Saxonianamber from the Bitterfeld area (Lower Miocene, Germany),based on a single specimen which I have studied. The state ofpreservation of the fossil is poor and it is extensively maskedby a whitish, opaque layer. The antennae are short but notflattened. The compact flagellum consists of about sevensimilar, cylindric articles. It somewhat resembles the antennaeof the genusDryops, although an enlarged pedicel is notdiscernible. The relatively long, terminal tarsomere with itsclearly visible claws is also reminiscent of certain Dryopoidea.The specimen is definitely not a fossil paussid which aretherefore still only known from Baltic and Dominican amber.

Poinar (1992: 134, Fig. 73) illustrated an ant nest beetlewhich is described here together with a second specimen fromDominican amber inclusions in the Poinar collection.

Classification and ranking of Protopaussinae andPaussinae

During the past few decades a number of authors have proposeddifferent ranking systems for the adephagans and particularlyfor the carabidoids. Ranks such as Paussidae, Paussinae,Paussitae, Paussini and Paussina have been used for the sameor different monophyla (see Nagel, 1987a: 28; Ball & McCleve,1990: 37f). No standardized ranking system prescribing theuse of a particular term for one monophylum such as thePaussinae (sensuBasilewsky, 1953, 1962; Crowson, 1955,1981; Luna de Carvalho, 1983, 1989; Nagel, 1987a; Zaballos& Jeanne, 1994) is accepted generally. The same taxon wascalled Paussini by Darlington (1950), Kryzhanovskij (1976,

© 1997 Blackwell Science Ltd,Systematic Entomology, 22, 345–362

1983), Casaleet al. (1982), Erwin (1984, 1985, 1991), Erwin& Sims (1984), Stork (1986) and Lawrence & Britton (1994),{Cerapteritae1 Paussitae} by Jeannel (1941, 1946) andPaussidae by most authors until about 1950. On the other hand,the monophylum comprising ozaenines, protopaussines andpaussines has been referred to as Paussidae (Crowson, 1955;Deuve, 1993), Paussinae (Ball, 1973; Erwinet al., 1977;Reichardt, 1977; Vigna Taglianti in Casaleet al., 1982;Kryzhanovskij, 1983; Stork, 1986) and Paussina (Bousquet &Larochelle, 1993). The use of the rank Paussitae (Arnett, 1985;Arndt, 1993) implies that the classification of Erwin (1984,1985, 1991) and Erwin & Sims (1984) had been followed,who includedNototylusas a member of this monophylum (seealso Lawrence & Newton, 1995). Deuve (1993, 1994) presentsdata which are ambiguous with regard to the classification ofNototylus. He suggested a sister group relationship between‘Nototylidae’ and ‘Paussidae’, but with some uncertainty.

In contrast to the recognition of sister group relationships(adelphotaxa) and, consequently, their relative ranking in theclassification system, the question of formal ranking inphylogenetic systematics is not a scientific problem but amatter of convention. Table 1 shows the classification and thenomenclatural use of the taxa as applied in this paper. InTable 1, some inconsistencies regarding the requirement ‘samehierarchic level–same suffix’ are accepted against thebackground that it is necessary to change the rank when newsister group relationships are discovered.

Ball & McCleve (1990) considered the possible paraphylyof the Ozaeninae & Beutel (1992, 1995) presented furtherevidence based on larval characters (see also Arndt, 1993) (seefurther discussion in ‘The taxonomic position ofProtopaussus’ section).

The classification of Protopaussinae and Paussinae given inTable 1 is mainly based on shared derived characters asdiscussed by Nagel (1987a, 1994) (see Table 2). The reasonsfor the present position of the Protopaussinae are given below(see ‘The taxonomic position ofProtopaussus’ section). Itdiffers from Basilewsky’s (1953, 1962) interpretation but agreeswith Darlington (1950), Luna de Carvalho (1989) and Ball &McCleve (1990). The main differences from Darlington’s(1950) classification of Paussinae are the splitting of his‘Cerapterina’ and the establishment of the two monophylaCarabidomemnitae and Paussitae. The ‘Cerapterina’ ofDarlington (1950) were founded on symplesiomorphies suchas nine free articles in the flagellum (the presumed ancestralstate) and the lack of trichome-bearing clefts or their vestigeson the medium part of the pronotum. Autapomorphic characterstates of the main taxa are given on Table 2. The Arthropterinihave recently been confirmed to be monophyletic (Nagel, 1994).

The fossil taxaEopaussusWasmann,SuccinarthropterusKolbe, PleurarthropterusWasmann,BalticarthropterusNagel,Cerapterites Wasmann, Protocerapterus Wasmann andArthropterites Wasmann belong either to the supertribeCarabidomemnitae or they are sister groups of{Carabidomemnitae1 Paussitae}. Several specimens ofBalticarthropterusandCerapteriteswere proven to be membersof the Carabidomemnitae as defined in Table 2 below (Nagel,1987a,b).

New fossil paussids 347

Table 1. Classification of the Protopaussinae and Paussinae (revised from Nagel, 1987a: 33, Fig. 4, p. 35ff; see also Ball & McCleve, 1990; fordiscussion on the higher taxa). The listed taxa are monophyletic unless stated otherwise. The position of the taxa indicates phylogenetic relationship(see Hennig, 1981). Unknown relationships are present where more than two taxa are listed within one rank.

{Metriinae 1 Ozaeninae1 Protopaussinae1 Paussinae}Metriinae{Ozaeninae1 Protopaussinae1 Paussinae}

Ozaeninae (paraphyletic!){Protopaussinae1 Paussinae}

ProtopaussinaeProtopaussusGestro

PaussinaeCarabidomemnitae

EohomopteriniEohomopterusWasmann

CarabidomemniniCarabidomemnusKolbe s.l.

PaussitaeArthropterini

MegalopaussusLeaArthropterusMacLeay

CerapteriniMesarthropterusWasmannCerapterusSvederus

PentaplatarthriniHexaplatarthrina

HexaplatarthrusJeannelPentaplatarthrina

HyperpentarthrusKolbePentaplatarthrusWestwood

HomopteriniHomopterusWestwood

HeteropaussiniHeteropaussusThomson

PaussiniPlatyrhopalina

PlatyrhopalopsisDesneuxEuplatyrhopalusDesneuxLebioderusWestwoodStenorhopalusWasmannPlatyrhopalusWestwood

{Ceratoderina1 Paussina}Ceratoderina

MerismoderusWestwoodCeratoderusWestwoodPaussomorphusRaffray

PaussinaPaussusL. s.l.LeleupaussusLuna de CarvalhoGranulopaussusKolbeHylotorusDalman

New paussid taxa

Protopaussus pristinussp.n.

Type material

Holotype, DOMINICAN AMBER: Sex indet., No. C 7–59 B(Poinar collection, Department of Entomology, Oregon StateUniversity, Corvallis).


Amber matrix

Size: 353 15 3 8 mm. The piece of amber contains severalsmall air bubbles and a minute darkish part which may be aremnant of rotten wood.

Etymology

The epithet, a Latin adjective, means former, earlier, previous,referring to theProtopaussusspecies of former times.

348 Peter Nagel

Table 2. Main arguments for the monophyletic status of the major taxa of Protopaussinae and Paussinae (synapomorphic character states of allsubordinate taxa) (see Nagel, 1987a, for discussion of additional autapomorphic character states of tribes and higher taxa). Larval characters arenot considered here because sufficiently detailed descriptions are available only for representatives of Paussini (Luna de Carvalho, 1992; Arndt &Beutel, 1995).

{Protopaussinae1 Paussinae}—Labial lacinia small, without dense brush-like pilosity typical of other Carabidae (see discussion in ‘The taxonomic position of

Protopaussus’ section).Protopaussinae

—Unique pattern of the prothoracic trichome system (Darlington, 1950).Paussinae

—Median cell (oblongum) of inner wings (alae) triangular, narrowed in front (Darlington, 1950).—Antennomere 2 (pedicel) very small, deeply sunken into the apex of antennomere 1 (scape) (Darlington, 1950; Basilewsky, 1962), connected

to antennomere 3 (flagellomere 1) by rigid wedging (Nagel, 1980, 1987a).Carabidomemnitae

—Dovetailed part of elytral suture considerably shortened, ending at or short distance behind middle of elytra (Nagel, 1987a)Paussitae

—Front coxae very prominent and contiguous or almost contiguous. Pleural lobes (proepimeres) enlarged, prolonged and contiguous behindcoxae, leaning on prosternal process (Nagel, 1987a; cf. Darlington, 1950; Baehr, 1979).

Fig. 1. Photograph ofProtopaussus pristinussp.n.

Diagnosis

Generic membership is mainly established by the vaultedforehead with vertical clypeus, large latero-basal trichomes ofthe pronotum, the presence of full-size pedicel (antennomere2) and the missing raised lateral margin of the elytra. It isdistinguished from all congeneric species by the presence of apronounced rim around the frontal part of the vertex borderinga deep depression and by its uniformity in colour (althoughthe latter might be an artefact due to poor preservation).

Description(see Figs 1, 2)

Body length4.0 mm. The real proportions of the specimen’sbody parts are difficult to reconstruct because the specimen


was obliquely compressed laterally and from behind duringthe embedding process. The wrinkled basic structure of almostthe entire surface seems to be an artefact due to (mostly lateral)pressure and/or shrinking.

Uniformly blackish, no particular colour pattern discernible;head (except for temporae and dorsal part of rim) and prothorax(except for latero-basal trichomes) glabrous, antennae denselycovered with short setae, especially from antennomere 3onwards; legs with adjacent scattered hair; elytra with loosepilosity of mostly erect, slightly backward slanting hair, eachinserted in a large puncture.

Head marked by a semicircular rim around frontal part ofvertex; frontal part of rim very high and interior edge abruptlydropping away towards vertex; rim steeply falling, level withtemporae further backward, at level of hind parts of eyes;lateral parts of epistome approximately level with dorsal surface


Fig. 2. Habitus ofProtopaussus pristinussp.n. with the right legs indorsal view (reconstruction). Adjacent, basic pilosity of antennae andlegs not shown. Scale bar5 1 mm.

of eyes, i.e. markedly lower than adjacent rim; few scattered,very short hairs are evident at dorsal part of rim; a central stripof the epistome rises evenly from fore margin to top of rim;real clypeus including antennal insertions vertically positionedon forehead beneath epistome (as generally found inProtopaussus); temporae not lying flat against eyes but slightlyyet distinctly projecting in an angle; no pubescence discernibleon head with exception of very short and almost recumbentpilosity on temporae; although surface structure of head isindiscernible, no coarse punctation likely to be present.

Antennaewith 11 antennomeres, reaching back almost toend of first third of elytra; scape™ 2.5 times longer thanwide; pedicel globular, inserted into scape by globular socket;antennomeres 3–10 of decreasing length, all with straight,parallel margins from above (i.e. more or less cylindrical), 7–10 almost globular with convex margins when seen frombehind; antennomeres 3–5 distinctly longer than wide, 10 atrace wider than long; antennomere 3™ 1.5 times longer thanpedicel; apical article (antennomere 11) a little more than twiceas long as wide with marked, rounded tip (from all sides); allflagellomeres (antennomeres 3–11) with dense punctation and


pubescence, on scape and pedicel slightly more widely spaced;right antenna cut off at the 5th and 6th article at the surfaceof the piece of amber.

Mouthpartssimilar to those of extantProtopaussusspecies;mandibles protruding, rather long, slender, sharply pointed;maxillary palpus slender with apical article slightly longer thanthe 2 preceding ones together; labial palpus with apical articleslightly swollen and little longer than the apical joint ofmaxillary palpus; its tip narrowly blunted; labrum notdiscernible.

Distal part of pronotum narrower than head, divided alongmid-line by a deep triangular furrow; surfaces of the 2 wing-like plates of frontal part difficult to discern, but there seemsto be at least one impression each; proximal part of pronotumwith 2 large wing-like projections, each ending in a bigtrichome, typical ofProtopaussus; proximal, central portion ofpronotum with a central, narrow, low, but sharp carina;additional surface structures, such as further depressions orbumps, were not detected; their presence on the whole pronotumcannot be excluded, due to poor transparency of the amber inthis area.

Elytral disc flattened dorsally; subapical folds (flanges ofCoanda) clearly visible; humeri protruding to the front, similarto other Protopaussusspecies; disc with dense and irregularpunctation, resulting in a rugose surface structure similar toextant species of this genus; pilosity almost erect, of ratherlong, scattered hair; 11 straight, long setae of the right seriesumbilicata visible.

Hindwingsof full size (macropterous); right hindwing partlyunfolded, exposing the large, roundish, darkened area(pterostigma) adjacent to the tip of frontal marginal vein(subcosta plus radius anterior).

Legssimilar to those of extantProtopaussusspecies; inneredge of fore tibiae with weak, oblique emargination lined withequally short hair (comb organ); tarsomeres 1–4 of decreasinglength, tarsomere 5 almost as long as 2–4 together; clawssimple; tarsomeres 1–4 underneath with dense, shortpubescence; pubescence of at least tarsomeres 2 and 3 offore tarsi possibly consisting of adhesive pads (not clearlydetectable); similar to extantProtopaussusspecies, terminalspurs missing or indistinguishable from apical pubescence oftibiae; fine longitudinal carinae on tibiae not detectable whichis possibly due to the disadvantageous position of the legs inthe piece of amber.

Proepimeres(pleural lobes) broad, closing the forecoxalcavities from behind; pro-, meso- and metacoxae not clearlyvisible; ratio of length of abdominal sternites similar to extantProtopaussusspecies.

Temporal and geographical distribution

Known only from the Dominican Republic, Hispaniola, WestIndies, from amber of Upper Eocene to Lower Miocene age.

Chorological affinities

All extant species of the genusProtopaussusare knownonly from the Oriental Region (Fig. 5).Protopaussus pristinus

350 Peter Nagel

sp.n. is the first fossil species of this genus and the firstfound outside the Oriental Region and is thus allopatricand allochronic to its congeners. Its Early to Mid-Tertiaryoccurrence on the island of Hispaniola, West Indies and itszoogeographical relationship to the extant Oriental distributionis discussed below (see ‘The distribution of theProtopaussinae’ section).

Phylogenetic affinities

All major apo- and plesiomorphic characters of extantProtopaussusspecies are present inP.pristinus sp.n. (seesections on ‘Diagnosis’ and ‘Discussion’ for details). Thepronounced relief of the vertex represents a more derivedcharacter state compared to all congeneric (described) species.The unicoloured elytra may represent a plesiomorphic characterstate, provided that it is not an artefact due to fossilization andthat it is not a secondary derived character state. The lack offine longitudinal carinae on the tibiae would represent aplesiomorphic character state provided that they are reallymissing (see sections ‘Description’ above and ‘Discussion’below). The shape of the antennomeres seems to be intermediatebetween the two groups regarded as subgenera by Luna deCarvalho (1989). Only specimens ofP.walkeriandP.almorensiswere available to me for comparison and the publisheddescriptions of the other species are not detailed enough toallow conclusions on the phylogenetic relationships within thegenus. A modern revision of the genus is required to allowfurther conclusions.

Discussion

The subapical elytral folds (flanges of Coanda) place thespecimen as a member of the monophyletic group{Ozaeninae1 Protopaussinae1 Paussinae}. The missingraised lateral margin of the elytra, the presence of a full-sizepedicel (antennomere 2), the large latero-basal trichomes ofthe pronotum, the vaulted forehead with a vertical clypeus andthe modified mouthparts are the decisive characters that placethe specimen within the Protopaussinae. Apart from thesecharacters, the overall shape of the body (habitus) immediatelyis clearly the Protopaussinae-like.

The extant Protopaussinae are represented by seven describedspecies within the genusProtopaussus. Recently, more specieshave been collected on Sulawesi and Borneo (Stork, BMNH,in litt . 1992). They are not yet described.Protopaussus pristinussp.n. is very similar to extant species in its general habitusand, particularly, the shape and structure of the legs and elytra.The structural details of the head and the pronotum are unique,yet consistent with the basic pattern of the extant species. Thepronounced rim around the frontal part of the vertex borderinga deep depression is not known from any other species. Noneof them has such a pronounced relief, although elevations anddepressions are present. Only a few characters of the pronotumwere detected in the amber specimen because of the poorquality of the amber in this area. There may well be a stronger


similarity in pronotal structure to extant species than is atpresent indicated by the description and the correspondingfigure.

Protopaussus pristinussp.n. differs from all extant speciesby its uniformity in colour. The extant species have eitherblackish elytra with light brown to orange spots, or light brownto orange elytra with blackish spots. It must be borne in mindthat surface coloration in amber inclusions is not alwaysproperly preserved. The specimen in hand has a somewhat‘carbonized’ appearance and it may originally well have hadsuch a colour pattern.

Unfortunately, it is not possible to confirm with certaintythe presence or absence of fine longitudinal carinae on thetibiae of P.pristinus sp.n. Carinae are clearly visible on alltibiae of a specimen ofP.walkeri Waterhouse (1897) (CHINA:Chinkiang, 32°129N, 119°309E, 29.v.1924, Elgin Suenson leg.,ZMK), available to me for comparison. Such carinae were alsodescribed by Gestro (1892) forP.feae, but they are missing inthe NHMB specimens ofP.almorensiscurrently available tome (see Appendix).

The extant species are 5–6 mm long, except forP.kaszabiLuna de Carvalho from Formosa (Taiwan) which is 4.5 mmlong. The fossil species has a length of only 4 mm. The latterwas obviously compressed and/or shrank during embedding.This shrinking, however, must have affected its width ratherthan its length as is indicated by the longitudinal direction ofthe wrinkles.

Luna de Carvalho (1967, 1989) distinguished two subgenera:ProtopaussusGestro, 1892,s.str. (type species:P.feaeGestro;syn.: AtavopaussusKolbe, 1924) andArchipaussusLuna deCarvalho, 1989 (type species:P.javanus Wasmann). Thenominate subgenus has blackish elytra with orange spots andmore cylindrical antennomeres andArchipaussushas orangeelytra with blackish spots and more globular flagellarantennomeres. Within this classification the new fossil speciesresembles the subgenusProtopaussus s.str., as the flagellarantennomeres are more cylindrical than globular andparticularly because antennomere 3 is clearly longer than thepedicel (antennomere 2) and also much longer than it is broad.On the other hand, with antennomeres 7–10 more globularwhen seen from behind, it resembles more closely the subgenusArchipaussus. The two subgenera have been established on thebasis of structural similarity of their respective members. Nophylogenetic analysis has yet been attempted to test themonophyly of the two subgenera. A modern revision of thegenus is needed to allow conclusions on the phylogeneticrelationships of its members, to recognize monophyla withinthe genus and, thus, to diagnose and denominate possiblesubgenera. Without such background information, there seemsto be no good reason to placeP.pristinussp.n. in either one ofthe two established subgenera or in a new one.

Eohomopterus poinari sp.n.

Type material

Holotype, DOMINICAN AMBER: Sex indet., No. C 7–59 A(Poinar collection, Department of Entomology, Oregon StateUniversity, Corvallis).


Amber matrix

Size: 29 3 17 3 11 mm. The piece of amber containsnumerous, mostly minute air bubbles. Other inclusions are notdiscernible. A large, thin, opaque, densely streaked layer masksthe head including the antennae, the left fore and middle legs,the left prothorax and the left basal part of the abdomen. Thinlayers of air cover large parts of the body thus masking theview of surface structures.

Etymology

Named for George O. Poinar, Jr., Professor of EntomologicalSciences at the University of California, Berkeley (at present:Department of Entomology, Oregon State University,Corvallis), who discovered the specimen, identified it as apaussid beetle and illustrated it in his bookLife in Amber(Poinar, 1992: 134, Fig. 73).

Diagnosis

Generic membership established by shortened dovetailedpart of elytral suture, dilated apical article of labial palpus,presence of protibial emargination and small tarsomere 4 whichis largely concealed at base of slightly lobed tarsomere 3.Distinguished from all fossil and extant congeneric speciesby its bigger size and its apically lobed, rather short andbroad, tibiae.

Description(see Figs 3, 4)

Length of bent body 8.0 mm, length of naturally orientatedbody presumably roughly 9 mm.

Numerous, some carinate, wrinkles of mostly irregularlongitudinal direction discernible on the elytra, the partiallyexposed propygidium and the pygidium. Pronotum also slightlycompressed laterally, indicating a possible lateral shrinking ofentire specimen.

Colour dorsally and ventrally light brown includingpygidium, but legs and antennae markedly darker; posteriormarginal parts of epipleura in colour very light, almost whitish.This is obviously equivalent to the reduced sclerotization andtransparent appearance of the corresponding parts of extantEohomopterusandCarabidomemnus s.l.species. Glabrous (or,at least, no distinct pilosity discernible) except for ventral tarsalpads, few short hairs close to the hind coxae and row ofequally short hair on tibial lobe; punctures of series umbilicatapresent, indicating presence of corresponding setae.

Head including very large eyes approximately as wide aspronotum; temporae not visible, not projecting; head betweeneyes slightly depressed, with 2 small, shallow tentorial pitsbetween posterior parts of eyes; frons, vertex and neck finelypunctured; view of frontal part including clypeus, labrum,entire right and most of left antenna masked by opaque layer.

Antennaelonger than combined head and pronotum, reaching


Fig. 3. Ventral view of the left antenna ofEohomopterus poinarisp.n.

tip of scutellum if bent backward; scape (antennomere 1)compressed, slightly longer than wide; pedicel (antennomere2) not visible; flagellum of 9 articles (antennomeres 3–11),rather slender, flat, parallel (Fig. 3); first article of the flagellum(3rd antennomere) half as long and almost as wide asantennomere 4, broadly triangular with acute inner (frontal)lateral angle; antennomere 4 rectangular; followingantennomeres with frontal margin obliquely rounded towardsbase; frontal margin of antennae thus slightly serrate;antennomere 11 almost as long as broad, with apex evenlyrounded.

Terminal article of labial palpus very large, triangularlyrounded; ligula not clearly discernible; terminal (apical) articleof maxillary palpus 2.25 times longer than broad, bluntlypointed; mandibles not visible.

Pronotumflat, slightly wider than long, markedly narrowedtowards base; lateral margins slightly convex in distal half,straight towards base; anterior and posterior angles broadlyrounded; lateral margins sharply marked and slightly raisedwith adjacent, shallow furrow on disc; this furrow slightlywidened and deepened towards base; a central, longitudinal,narrow furrow present on disc, ending shortly before anteriorand posterior margins; pronotal disc finely punctate; notopleural(tergopleural) suture complete.

Elytral discwith fine and scattered punctation, slightly rugosetowards lateral margins; distinct pilosity missing, obviouslyglabrous; some punctures of series umbilicata discernible; the

352 Peter Nagel

Fig. 4. Details of the legs ofEohomopterus poinarisp.n. A, Ventral view of right foreleg; B, dorsal view of left middle leg; C, right middle tarsus;D, ventral view of right hind leg; E, ventral view of left hind leg.

shrinking led to longitudinal wrinkling, most distinct on theposterior third of elytra; true suture (dovetailed engagementsof both elytra) ending shortly behind middle; subapical folds(flanges of Coanda) long and narrow; shoulders not projecting,yet distinctly marked by triangular depressions on both sidesof scutellum; the downward flexed epipleura broad alongthe metepisternum, but narrowing and less inclined towardssubapical fold (flange of Coanda); dorsal disc separated frombroad anterior part of epipleura by very fine linear edge (thisstructure may be an artefact caused by shrinking); scutellumcompletely exposed, shaped like a narrow, sharply pointedtriangle.

Macropterous: tips of folded hind wings slightly projecting.Legs (Fig. 4): tibiae compressed with outer apical angle


distinctly lobed; 2 strong terminal spurs present; protibiaeslightly emarginate along apical half of inner edge (antennacleaner); comb (longitudinal row of dense, equally shortbristles) not visible but probably exists as in all other membersof the group; tarsomeres 1–3 large, 2 and 3 slightly lobed;tarsomere 4 very small, largely concealed at base of emarginatedtarsomere 3; tarsomeres 1–3 of forelegs underneath clothedwith tarsal pads; this pubescence only visible on tarsomeres 2and 3 in middle and hind legs; presence or absence of padbeneath tarsomere 1 cannot be confirmed; if present, it mustbe less conspicuous than those of the following tarsomeres.

Front coxaeand total central and proximal part of ventralprothorax masked by middle leg; middle and hind coxae onlylaterally visible.


Fig. 5. Distribution of Protopaussinae. F,Protopaussus pristinussp.n.HISPANIOLA: Dominican amber; (d) Localities of extantProtopanssus.

Pygidium simple, surface in part strongly wrinkled (mostprobably due to shrinkage); propygidium partially exposed,its surface with similar longitudinal wrinkling; additionaltransversal rugosity present, resulting in a net-like structure.

Temporal and geographical distribution

Known only from the Dominican Republic, Hispaniola, WestIndies, from amber of Upper Eocene to Lower Miocene age.

Chorological affinities

The genusEohomopterushas a strict Neotropical distributionwith species known from the South American mainland andthe West Indies.

Eohomopterus poinarisp.n. is sympatric withE.paulmuelleriNagel, 1987. Both species are known from amber collected inthe Dominican Republic on the island of Hispaniola. This kindof amber originates from deposits of different age, rangingfrom the Lower Miocene through the Oligocene to the UpperEocene periods. The precise age of the two pieces of ambercontaining the specimens has not been established. Althoughboth species occurred at the Early to Mid-Tertiary periods,their synchronic existence cannot be confirmed and they maywell be separated by several millions of years.

Two extant species ofEohomopterusare known from SouthAmerica, namelyE.aequatoriensis(Wasmann, 1899) from


Ecuador (Semiradski) andE.centenariusLuna de Carvalho,1960, from Brazil (Rio de Janeiro).

A new extant Eohomopterusspecies has recently beendiscovered on Guadeloupe, Lesser Antilles, West Indies (R. L.Davidson, The Carnegie Museum of Natural History,Pittsburgh,in litt. 1991). Despite the fact that the West Indianground beetle fauna is relatively well known (Erwin, 1979;Erwin & Sims, 1984; Nichols, 1988; Liebherr, 1988a; Ball,1992), this finding indicates that the extant geographicaldistribution of the genusEohomopterusmay well include otherislands of the Lesser Antillean arc.

Phylogenetic relationships

The less pronounced reduction of the 4th tarsomere and theless enlarged tarsomere 3 inE.poinari sp.n. compared to allother congeneric species represent a more ancestral characterstate. The plesiomorphic shape of antennomere 3 (basal articleof the flagellum) is shared with the two described extantspecies, while the fossilE.paulmuelleri displays the morederived rhombic antennomere 3. The relatively short and broadtibiae with their lobed apical angles ofE.poinari sp.n. representa specific, more derived character state compared to the otherspecies of the genus.

No detailed revision of the genus and its sister group(Carabidomemnus s.l.) is available. All species of

354 Peter Nagel

Eohomopterusare known from their holotype specimens only,thus not allowing conclusions on their variability. Therefore,any statement on phylogenetic relationship must be preliminary.The distinct lateral dilatation of tarsomere 3 and concomitantstrong reduction of tarsomere 4 might be considered as asynapomorphy of E.paulmuelleri, E.aequatoriensis andE.centenarius. Consequently,E.poinari might form the sistertaxon of this possibly monophyletic group.

Discussion

The following are the most distinctive characters that classifythis species within the Carabidomemnitae: (1) the shorteneddovetailed part of the elytral suture (synapomorphy of{Carabidomemnitae1 [all?] Balticarthropterus1 Cerapter-ites [ 1 other fossil genera?]}) and (2) the strikingly dilatedapical article of the labial palpi (present in Carabidomemnitae,Arthropterini, Homopterini and Protocerapterus). Baltic-arthropterus and Cerapterites have thickened (rather thandilated) terminal articles of the labial palpi. Arthropterini andHomopterini have an elytral suture which almost reaches theapex. In Homopterini the subapical article of the labial palpiis also very wide (an autapomorphy).Protocerapteruswithsimilarly wide apical articles of the labial palpi has a verystout body and the tibiae are very short and wide, only a littlelonger than wide at the apex. The protibial emargination is aplesiomorphic character in the Carabidomemnitae and someArthropterus. The Eohomopterini (Eohomopterus) andCarabidomemnini (Carabidomemnus s.l.) are separated by theresting position of the aedeagus and the position of its apicalorifice as well as by the structure of the tarsi. The copulatoryorgan is not visible in the fossil specimen. The tarsi are slenderand the tarsomeres 1–4 are of decreasing length with tarsomere4 fully visible in Carabidomemnus s.l.By contrast, inEohomopterustarsomere 2 and, in particular, 3 are enlargedand laterally dilated whereas the 4th article is minute andlargely concealed at the distal emargination of the 3rd article.An enlarged 2nd and 3rd tarsomere and/or a minute penultimatearticle are present in several other genera of the Paussinaewhich, however, do not display the other characters listedabove. It is evident from this analysis that the present fossilspecimen must be classified as Eohomopterini within its onlygenusEohomopterus.

Three species ofEohomopterushave been described: theextant E.aequatoriensis (Wasmann, 1899) from Ecuador(Semiradski) andE.centenariusLuna de Carvalho, 1960, fromBrazil (Rio de Janeiro), and the fossilE.paulmuelleriNagel,1987, from Dominican amber. All are known from theirrespective holotype specimen only. A new extantEohomopterusspecies has recently been discovered on Guadeloupe, LesserAntilles, West Indies (R. H. Davidson, The Carnegie Museum,Pittsburgh,in litt. 1991).

Eohomopterus poinarisp.n. is very similar toE.paulmuelleriin the general shape of the antennae and the pronotum. Thevisible mouthparts are similar to those ofE.paulmuellerialthough the apical labial palpomere is more triangular and the


apical maxillary palpomere shorter and blunter inE.poinarisp.n. The most striking differences betweenE.poinari sp.n.and the other three described species are the bigger size ofE.poinari (9 mm as opposed to 5.5–6 mm) and, especially, itsshorter and broader tibiae with lobed apical angles. The dilationof the 2nd and 3rd tarsomeres is less pronounced than in anyof the other three species. The first article of the flagellum(antennomere 3) is not rhombic with acute angles at both sideslike in E.paulmuelleri (Nagel, 1987b: 142, Fig. 3) but it issimilar to that of E.centenarius(as illustrated by Luna deCarvalho, 1960: 150, Fig. 6A).

Key to genera of New World Protopaussinae andPaussinae and species of Eohomopterus

The key is intended to identify the adults of extant and fossilhigher taxa of the ozaenine–paussine lineages known from theNew World and to differentiate theEohomopterusspecies. Itis based on diagnostic (i.e. not necessarily synapomorphic)characters and applies exclusively to the New World taxa. Thekey starts from couplet 11 of Ball’s key to Carabidae of theNeotropical region (in Reichardt, 1977: 357, see Erwin, 1991:4). The taxonomic ranks generally used in the present paperare also applied to the key.

11(10’) Elytron without subapical fold at outer edge; anteriortibia simple, without longitudinal sulcus or completeantenna cleaner (see redescription by Deuve, 1994)(extant species, known only from one female fromEsperitu Santu, Brazil)............................Nototylinae,

Nototylus Gemminger & Harold,Nototylus fryi(Schaum)

11’ Elytron with subapical fold (flange of Coanda) atouter edge; anterior tibia with or without antennacleaner ......................................................................A

A(11’) Antenna of 10 clearly visible articles, article 2inconspicuous and largely concealed at apex ofantennomere 1; antennomeres 3–11 free but flat andbroad, forming a distinct, compact flagellum; terminalarticle of labial palpus markedly enlarged.................................................................................Paussinae, C

A’ Antenna of 11 clearly visible articles, article 2distinct, slightly shorter than article 3, articles 3–11moniliform, short-cylindrical, or slightly flattened;terminal article of labial palpus not markedlyenlarged, parallel-sided to slightly broader towardsapex ..........................................................................B

B(A’) Surface of head and pronotum with pronounced relief;forehead vaulted with clypeus vertical; pronotum withlarge latero-basal trichomes; anterior coxae moreprominent, separated by narrower process (from theNew World known only from a fossil specimenpreserved in amber of Tertiary age from theDominican Republic) ........................ Protopaussinae,

ProtopaussusGestro,Protopaussus pristinussp.n.


B’ Surface of head and pronotum of ordinary carabidshape, not markedly sculptured; forehead and clypeusnormal; pronotum without trichomes; anterior coxaenot much projected, separated from each other bynormal process ...........................................Ozaeninae

C(A) Tibia wide, with apical excavation for completereception of tarsus; fore coxae long, stronglyprojecting, contiguous; prosternal process invisiblebetween coxae; antenna cleaner at anterior tibiaabsent; antennal flagellum wider, with base of articlessinuose, or very wide with bases and apices ofsegments straight; penultimate article of labial palpusenlarged, transverse, apically as wide as base ofapical article; dovetailed part of elytral suture endingshort distance before apex of elytra (12 extant speciesdistributed from northern Mexico to northernArgentina, one fossil species preserved in amber ofTertiary age from the Dominican Republic).............................................................HomopterusWestwood

C’ Tibia narrower with tarsus not retractable and longerthan apical margin of tibia; fore coxae hardlyprominent, only slightly projecting beyond level ofcentral part of prosternum in front of coxae; prosternalprocess clearly visible between coxae, at least in theform of a narrow septum; antenna cleaner present:anterior tibia with longitudinal sulcus, equipped withrow of dense, equally short bristles; antennalflagellum narrower, with apex and base ofintermediate articles straight; penultimate article oflabial palpus small, of normal size; dovetailed partof elytral suture markedly shortened, ending shortdistance behind middle of elytra .......................................................................EohomopterusWasmann, D

D(C’) Tibiae relatively short and broad with outer apicalangles lobed and broadly rounded; apical margin oftibiae as wide as tarsomeres 1–4 together; tarsomeres2 and 3 only slightly dilated; antennomere 3 (1starticle of flagellum) transverse, triangular, with acuteangle at inner side and short, straight margin at outerside; body length 9 mm (known only from a fossilspecimen of unknown sex preserved in amber ofTertiary age from the Dominican Republic).....................................................Eohomopterus poinarisp.n.

D’ Tibiae relatively long and slender with outer apicalangles simple; apical margin of tibiae narrower, atmost as wide as combined tarsomeres 1, 2 and basalhalf of 3; body length 5.5–6 mm ............................ E

E(D’) Pronotum slightly cordiform, with lateral marginsconcave at basal third, forming well-marked basalangles; tarsomeres 2 and 3 strongly bilobed; apicalmaxillary palpomere three times longer thanpenultimate article (extant species, known only fromone male from Semiradski, Ecuador)..........................................Eohomopterus aequatoriensis(Wasmann)

E’ Pronotum with lateral margins convex or straight atbasal third, basal angles rounded; tarsomeres 2 and3 only slightly dilated.............................................. F

F(E’) Antennomere 3 (1st article of flagellum) transverse,


triangular, with acute angle at inner side and short,straight margin at outer side; underside of tarsomereswith glabrous longitudinal mid-line; apical maxillarypalpomere 3 times longer than penultimate article(extant species, known only from one male from Riode Janeiro, Brazil ............Eohomopterus centenarius

Luna de CarvalhoF’ Antennomere 3 (1st article of flagellum) transverse,

rhombic, acute at inner and outer side; tarsomeres 2and 3 completely pubescent underneath; apical andpenultimate maxillary palpomeres of equal length(known only from a fossil specimen of unknownsex preserved in amber of Tertiary age from theDominican Republic) ...............................................................................Eohomopterus paulmuelleriNagel

Subfamily Protopaussinae

Habit

Almost nothing is known about the habits of theProtopaussinae. All we know aboutProtopaussusis what weare able to infer from collection data, from structural featuresof the adults (larvae are unknown) and from comparison withrelated taxa.

Waterhouse (1897: 392) reports that the type specimen ofP.walkeri was collected by ‘sweeping long grass’. In therain forest of the Dumoga-Bone National Park, N. Sulawesi‘Paussinae’ (which include one undescribed species ofProtopaussusand some ozaenines of the generaEustra andPseudozaena; Stork, BMNH, in litt. 1992) were collected in1985 by means of light traps, malaise traps, flight intercepttraps, canopy fogging, unbaited pitfall traps, from leaf litterand soil samples and in or on dead wood (Hammond, 1990:251). A series of several males ofP.feaewas collected in amalaise trap in open, second-growth tropical forest, at 1300 m,Doi Inthanon National Park, Chiang Mai Province, Thailand(Ball, UASM, personal communication, 1995).

These and additional collection data (see appendix) permitonly the inference that the members ofProtopaussuslive inand near tropical forests at 500–2300 m and that, at least, themales fly.

All members of the sister taxon of the Protopaussinae, thePaussinae, are associated with ants, at least during their larvalstage. It was shown that Paussinae are predatory inquilines inants’ nests and that host-specifity is unlikely (Nagel, 1987a;see Kistner, 1979, 1982). Some Ozaeninae are also known tobe myrmecophilous. Termitophily has been suspected to existin some members of the Paussinae, yet no member of the{Ozaeninae1 Protopaussinae1 Paussinae} is known forcertain to be associated with termites (Nagel, 1987a: 118f).The form of body and mouthparts and the presence of trichomeson the pronota of adults indicate thatProtopaussusis associatedwith ants or termites. Termitophily is, however, highly unlikelybecause trichomes are very rare among termitophiles (Kistner,1979, 1982).

356 Peter Nagel

From these data one can infer reasonably that the membersof Protopaussusare myrmecophilous (see Darlington, 1950).Numerous ant genera are known from Dominican amber(Wilson, 1988; Baroni Urbani, 1995).

The taxonomic position ofProtopaussus

The fossilProtopaussusis of little help for the reconstructionof the phylogeny of the related subfamilies but some commentson this problem are presented here. The sister grouprelationships within the monophylum {Ozaeninae1Protopaussinae1 Paussinae} have been the subject of debate.Jeannel (1941, 1946), Darlington (1950), Janssens (1953), Lunade Carvalho (1965, 1989), Hlavac (1975), Nagel (1980) andBall & McCleve (1990) considered the Protopaussinae andPaussinae as more closely related taxa and separated from theOzaeninae. Basilewsky (1953, 1962) regarded theProtopaussinae as members of the Ozaenine complex andseparated from the Paussinae. Nagel (1987a: 27) placed theProtopaussini amongst the Ozaeninae and left it open whetherProtopaussusrepresents the sister group of the Paussinae.

While most early authors followed Gestro’s (1892) originalclassification without further discussion, Luna de Carvalho(1967, 1989) listed the following reasons: the mouthparts,antennal scape, pilosity of basal antennomeres, structure of thehead, pronotum and elytra and structure of the male genitaliaare more similar to the Paussinae than to the Ozaeninae.Darlington (1950) stressed the following similarities betweenProtopaussinae and Paussinae: absence of raised lateral marginsof the elytra, presence of trichomes and modified mouthparts.Basilewsky (1953, 1962) listed the following similaritiesbetween Protopaussinae and Ozaeninae: antennomere2 (pedicel) not reduced, procoxae flat and distinctly separatedby the prosternal process, metacoxae separated (although onlynarrowly), median cell of the hind wing rectangular, tactilesetae present. It must be noted that the flat procoxae are alsopresent in some extant Paussinae (Carabidomemnitae) and allfossil ones and that tactile setae are also occasionally presentin some Paussinae (Carabidomemnus s.l.).

In order to reconstruct the phylogeny one has to findshared-derived character states (synapomorphies) rather thansimilarities. All the characters listed by Basilewsky (1953,1962) are shared ancestral characters (symplesiomorphies) andtherefore cannot be used for phylogenetic reconstruction. Mostof the characters listed by Luna de Carvalho (1967) can beinterpreted as convergences and are thus also not very helpfulfor the phylogenetic reconstruction. The relative enlargementof the antennal scape ofProtopaussusis a character whichis shared with the Paussinae. According to Ball (personalcommunication, 1995), the members of Ozaena (which I havenot seen) also have large antennal scapes. This character mightbe interpreted as a synapomorphy of Ozaena, Protopaussinaeand Paussinae. In addition, the lacinia of the labium ofProtopaussusis basically identical to that of the Paussinae (P.Nagel, unpublished data, see also Nagel, 1994). The shape ofthe lacinia of the Paussinae is unique amongst beetles and,although correlated to the common fluid feeding mechanism


evolved with myrmecophily, cannot be explained byindependent evolution (analogy, convergence) in individualtribes. At present, the unique structure of the lacinia may beregarded as a synapomorphy of the two sister taxaProtopaussinae and Paussinae and, thus, support the existence ofthe monophylum {Protopaussinae1 Paussinae} (see Table 2).

The Ozaeninae are a paraphyletic taxon with theMystropomini most probably forming the sister group of{remaining Ozaeninae1 Protopaussinae1 Paussinae}. Fromanalysis of the larval characters of the three ozaenine generaPachyteles, Afrozaena and Physea, Beutel (1992, 1995)postulated a sister group relationship betweenPhyseaandPaussinae. Ball & McCleve (1990) assumed that either thePhyseaor Ozaenalineage could be the sister group of thepaussines. Recently, Ball (personal communication, 1995)regardsOzaenaas the better candidate for the sister group of{Protopaussinae1 Paussinae}, based on the enlarged antennalscape and the reduced antenna cleaner of the fore tibia.

The distribution of the Protopaussinae

Old World affinities

Protopaussus pristinussp.n. is the first recorded fossilProtopaussus, but its existence back to the Early Tertiary hadindirectly been proven by the discovery of Paussinae fromBaltic amber. A true surprise, however, is the discovery of amember of the Protopaussinae in Dominican amber. The presentdistribution of the Protopaussinae is strictly Oriental (includingSulawesi) (Fig. 5).

The majority of the Dominican amber fauna showsrelationships to extant Neotropical and even Caribbean taxa(Poinar, 1992; for ants see Wilson, 1985a, 1988; Baroni Urbani,1995). I do not know of any other taxa with a distributionpattern identical toProtopaussus. However, some examples oftaxa with essentially similar distribution patterns have beendiscussed by Poinar (1992) and, in detail, by Baroni Urbani(1995).

Given the myrmecophily of Protopaussinae, similaritieswith the distribution of ants is of particular interest. Thedolichoderine ant genusLeptomyrmexhas been detected inDominican amber, with extant species only known from theMoluccas, New Guinea, the Aru Islands, eastern Australia andNew Caledonia (Baroni Urbani, 1980; Baroni Urbani & Wilson,1987). One species of fossil DominicanAnochetus(Ponerinae)is probably related to an extant species from New Guinea,New Britain and the Solomon Islands, whereas other fossilcongeners from Dominican amber have Neotropicalrelationships (de Andrade, 1994). The closest relatives of twoPheidole(Myrmicinae) species known from Dominican amberare extant species restricted to the Malayan, Australian andOceanic regions (Baroni Urbani, 1995). One species of fossilDominicanAphaenogaster(Myrmicinae) is most closely relatedto two Indomalayan species within a species group furtherdistributed in the Indomalayan and Malagasy regions (deAndrade, 1995).

Other taxa known from Dominican amber also show Old


World affinities. The genusValeseguya(Diptera: Anisopodidae)is only known from one fossil Dominican species and oneextant Australian species (Grimaldi, 1991). The extant membersof the genusMesochria (Diptera: Anisopodidae) are knownfrom Africa, Seychelles, Borneo and Samoa and one fossilspecies has been described from Dominican amber (Grimaldi,1991). The single extant species ofMastotermes(Isoptera:Mastotermitidae) is restricted to Australia while the fossilrecord comprises species of Early to Mid-Tertiary age fromEurope, Mexico and Hispaniola (Dominican amber) (Krishna& Grimaldi, 1991).

Some extant insect taxa of the West Indies are most closelyrelated to Asian and Australian (including Pacific) taxa whichis regarded as an indication of their great age (Liebherr, 1988b;Grimaldi, 1988; see also Schuh & Stonedahl, 1986).

In summary one can state that in a certain number of taxafaunal relationships do exist with the Oriental and Australianregions, despite the generally significant relationship of thefossil Dominican amber fauna with the fauna of the NewWorld tropics. Of particular interest are those which exhibitdisjunct distribution patterns composed of an Early to Mid-Tertiary existence in what is now the West Indies and an extantcongeneric presence in the Oriental and/or Australian regions,for which Protopaussusis another example.

Mesozoic vicariance?

The extant pantropical range of paussid beetles suggests aGondwanian distribution with subsequent vicariance due to thebreak up of this palaeo-continent. Jeannel (1942: 250)postulated that the family Paussidae (corresponding to{Protopaussinae1 Paussinae}) ‘de´rive evidemment de souchesdifferenciees sur l’Inabre´sie avant la se´paration de l’Australie,c’est-a-dire au Trias ou au Jurassique infe´rieur’. Basilewsky(1953: 23) considered the Ozaeninae (includingProtopaussus)and Paussinae already having been separated at the time offragmentation of ‘l’Inabre´sie, au Cre´tacesuperieur’. Luna deCarvalho (1989: 31) regarded the paussid beetles as inhabitants‘deja des anciens continents Gondwana et Le´murie’. AlthoughDarlington (1979: 346) clearly favoured dispersal for theexplanation of the distribution of paussids, he considered apossible ‘Wegenarian explanation’.

In order to contribute to this question one has to considerthe age of the protopaussine and paussine lineages. The ageof Dominican amber from whichProtopaussusas well asPaussinae have been recorded, is estimated at 15–40 millionyears before present, covering the Lower Miocene throughOligocene to the Upper Eocene periods (Poinar, 1992).Paussinae are also known from Baltic amber, the age of whichis estimated at 35–40 (some deposits up to 50) million yearsbefore present (Lower Oligocene–Upper Eocene). FossilOzaeninae have not been reported. The stem species of{Protopaussinae1 Paussinae} was probably already amyrmecophilous member of the ozaenine complex.

The first fossil ants date back to the Lower Cretaceousperiod, between 107 and 114 million years ago (Upper Aptian)(Myrmeciinae, see Brandaõ, 1990; Brandaõ et al, 1990) The


first more advanced subfamilies of ants are known from Mid-Eocene deposits (Wilson, 1985b). Ho¨lldobler & Wilson (1990)suggested that the adaptive radiation of ants, resulting inhigh subfamily and genus diversity, started no later than thebeginning of the Tertiary period, about 65 million years ago. Thefirst myrmecophilous member of the ozaenine protopaussinepaussine lineage could only have evolved after social ants hadarisen, i.e. Mid- to Late Cretaceous or ‘in fact they can hardlybe older than the rise of ants as dominant social insects’(Darlington, 1950; p. 86), i.e. at the beginning of the Tertiaryperiod. The main host ants of paussines belong to moreadvanced subfamilies such as the Myrmicinae and, less oftenrecorded, the Formicinae. Ponerine and dolichoderine ants havebeen recorded only two times each as hosts of paussines.

In summary, the separation of the Protopaussinae andPaussinae happened before the Upper Eocene–Lower Oligocene(because of the presence of Paussinae in Baltic amber deposits)and the stem species of the {Protopaussinae1 Paussinae}arose earlier, i.e. not later than Lower to Middle Eocene andmost probably not earlier than the beginning of the Tertiary(Palaeocene) (parallel to the evolution of ants).

The rifting between the southern parts of Africa and SouthAmerica began during the Lower Cretaceous roughly 135million years ago and the separation of north-western Africaand northern South America was completed and biologicallyeffective about 100 million years ago (Late Cretaceous) (Pitmanet al., 1993; see also Scoteseet al., 1988; Parrish, 1993). Thebreak off from Gondwanaland of Madagascar, the Indiansubcontinent and Australia had also been completed by thisperiod, about 90 million years ago (Late Cretaceous) (Smithet al., 1981; Audley-Charles, 1987).

No data are available to postulate the presence or absenceof representatives of the protopaussine–paussine lineages inparts of the southern land masses during the Late Cretaceousto Early Tertiary periods. The hypothesis of a continuousGondwanian distribution during the Lower Cretaceous andsubsequent vicariance and diversification due to the breakingup of this continent is not supported by any kind of evidence.Mesozoic vicariance can be ruled out with reasonableconfidence, given the presumed younger age of Protopaussinaeand Paussinae.

Range contraction?

Two different explanations for a distribution patternessentially similar to that ofProtopaussushave been reviewedby Baroni Urbani (1995), predominantly based on ants: oneassuming a former cosmopolitanism followed by speciationthrough allopatry and subsequent extinction and rangecontraction in most of the former range and the other assuminga centre of origin followed by speciation events after dispersal.Baroni Urbani (1995) finds evidence only for the formerexplanation but no supporting evidence for the latter. Thehypothesis of Tertiary vicariance (mobilist geologicalhypothesis) fitted best to the area relationships of carabids ofthe extant circum-CaribbeanPlatynusclade (Liebherr, 1988b).

Dispersal within the Greater Antilles and with northern

358 Peter Nagel

Central America was not hampered by wide gaps of openocean for most of the Late Mesozoic and Cenozoic (Donnelly,1985, 1988; see also Ross & Scotese, 1988; Perfit & Williams,1989). Possible dispersal routes during the Late Cretaceous toEarly and Mid-Tertiary include North American-European andNorth American-East Asian land connections which mighthave been passable for the Protopaussinae only for certainperiods, for climatic reasons (see Stehli & Webb, 1985;Behrensmeyeret al., 1992; Prothero & Berggren, 1992; George& Lavocat, 1993; Goldblatt, 1993). Briggs (1994) presentsbiological data which suggest that, despite the most acceptedgeophysical theory, a dispersal of terrestrial and freshwateranimals between North and South America must have beenpossible during the Upper Cretaceous to Palaeocene periods.Darlington’s (1979; p. 346) brief discussion on the distributionhistory of paussids by dispersal has to be partially revised dueto the subsequent discovery of Dominican amber paussids andthe new interpretation of ‘primitive’ and ‘derivative’ paussids.

Darlington (1950: 114) assumed a Late Cretaceous origin ofpaussids (Protopaussinae1 Paussinae) with the place of origin‘probably somewhere in the main regions of the Old Worldtropics. Perhaps (but this is sheer guess-work) it was in whatis now the Oriental Region’. With the new fossil recordavailable, at least the second supposition now seems unlikely.

A Laurasian distribution of protopaussines in Early Tertiarytimes is not contradicted by strong arguments. One argumentcould be the missing evidence of Protopaussinae in Balticamber or fossiliferous beds of other parts of the (northern)world. Despite the comparatively good knowledge of the richBaltic amber fauna, in particular with regard to small specimensup to 5 mm, such a deficiency in the fossil record is no proofof their factual absence.

The limited data base does not allow further tests of thedifferent hypotheses for the protopaussines. Considering theprotopaussines’ myrmecophily one could assume a distributionhistory largely congruent with that of their potential hosts (seeBaroni Urbani, 1995). Although additonal zoogeographicalscenarios may be applicable, parsimony suggests that thepresent Oriental range ofProtopaussusrepresents the relictof a large Tertiary range. The presumed geological age,consideration of geological history and the present and fossillocality records indicate thatProtopaussuswas widespreadthrough tropical Laurasia during the Tertiary.

Acknowledgements

I am very grateful to George O. Poinar, Jr, University ofCalifornia, Berkeley, for allowing me to study the two amberfossils from his collection and for his efforts in re-shaping andre-polishing the amber pieces. The provision of the Saxonianamber fossil by Erika Pietrzeniuk, Museum fu¨r Naturkundeder Humboldt–Universita¨t, Berlin and of Protopaussusspecimens by Ole Martin, Zoologisk Museum, København andMichel Brancucci, Naturhistorisches Museum, Basel, is verymuch appreciated. Many thanks are extended to Nigel E. Stork,The Natural History Museum, London and Robert L.Davidson,The Carnegie Museum of Natural History, Pittsburgh, for


allowing me to use their unpublished records and to ThierryDeuve, Museúm National d’Histoire Naturelle, Paris, andGeorge E. Ball, University of Alberta, Strickland EntomologicalMuseum, Edmonton, for providing locality data of museumspecimens. Thanks also to Erik Arndt, Institut fu¨r Zoologie,Universitat Leipzig, who provided some information onliterature. Cesare Baroni Urbani, Zoologisches Institut,Universitat Basel, was kind enough to read the manuscript andhe provided important information on literature. James K.Liebherr, Cornell University, Ithaca, reviewed a previous draftof the manuscript and provided valuable information, especiallyon new publications and on zoogeographical issues. I amparticularly grateful to George E. Ball, University of Alberta,Edmonton, for his friendly, detailed and constructive reviewof the manuscript. I am indebted to Sibylle Gussmann,University of Pretoria, for the revision of English of a previousdraft of the manuscript. I am also grateful to Eva Weber, Basel,for drawing a fair copy of Fig. 2.

Acronyms

BMNH, The Natural History Museum, London; MCSN, MuseoCivico di Storia Naturale, Genova; MNHN, Museúm Nationald’Histoire Naturelle, Paris; NHMB, Naturhistorisches Museum,NRS; Naturhistoriska Riksmuseet, Stockholm; UASM,University of Alberta, Strickland Entomological Museum,Edmonton; ZMK Zoologisk Museum, København.

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Accepted 23 June 1997

362 Peter Nagel

Appendix

Records of extantProtopaussusused for the compilation of the distribution map (Fig. 5).

1. Protopaussus almorensisChampion,INDIA : United Provinces, Central Almora Division, Sitoli, 21.vii.1921 (Champion) (Champion, 1923)[Almora: 29°369N, 79°409E].

2. Protopaussus almorensisChampion,NEPAL: Khali Ghandaki Valley, Tatopani, 1200 m, iv.1987 (A. Loy) (Vigna de Taglianti collection, Rome,vid./det. Nagel) [Kali Gandak R.: approximately 27°539N, 84°109E].

3. Protopaussus almorensisChampion,NEPAL: O-Nepal, Arun R., Lamobagar Gola, 1400 m, 28. 31.v.1980 (C. Holzschuh) (NHMB, vid./det.Nagel) [Lamobagar Gola: 27°399N, 87°219E].

4. Protopaussus almorensisChampion,SIKKIM : E. Sikkim, Deehiling, 1200 m, 29.iv.1984 (C.J. Rai) (NHMB, vid./det. Nagel) [approximately27°209N 88°409E].

5. Protopaussus almorensisChampion,INDIA : Inde Anglaise, Re´gion de Darjeeling, Pedong, 1935, compared with type G.J.A. (Chasseursindigenes) (Oberthur collection, MNHN & Luna de Carvalho collection) (Luna de Carvalho, 1967, Deuve, MNHN,in litt., 1994) [Darjeeling:27°029N, 88°209E].

6. Protopaussus almorensisChampion,INDIA : Indien, Darjeeling, Kalimpong, Hitli, 900 m, 8.v.1987 (Ch.J. Rai) (NHMB, vid./det. Nagel)[Kalimpong: 27°029N 88°349E].

7. Protopaussus almorensisChampion,INDIA : Indien, Darjeeling, Suruk, 900 m, 18.iv.1985 (Ch.J. Rai) (NHMB, vid./det. Nagel) [Darjeeling:27°029N, 88°209E].

8. Protopaussus feaeGestro,BURMA (MYANMAR ): Birmania, Monti dei Carin Cheba`, 900–1100 m (L. Fea) (MCSN) (Gestro, 1892) [approximately100 km NNE of Toungoo, Leito` / Leiktho / Lutasa village, approximately 19°109N, 96°459E, 1888, leg.Nasuelli, Missione Fea, teste Fea,1896, 1897].

9. Protopaussus feaeGestro,THAILAND : Chiang Mai Province, Doi Inthanon National Park, 1300 m, 2–12.v.1990, open, second growth tropicalforest, malaise trap (E.R.Fuller) [approximately 18°329N, 98°319E] (UASM) (G. E. Ball, UASM, pers. comm., 1995)

10. ?Protopaussus feaeGestro,THAILAND : NW Thailand, Soppong - Pai, 1800 m, 1–6.v.1991 (Pacholatko) (NHMB, vid./det. Nagel) [Soppong:19°309N, 98°159E; Pai: 19°249N, 98°269E].

11. Protopaussus jeanneliLuna de Carvalho,SABAH (Borneo): Ins. Bangeng, Waterstrook, ex Musaeo Van de Poll, 1909 (Oberthu¨r collection,MNHN) (Deuve, MNHN, in litt ., 1994, see Luna de Carvalho, 1960) (interpreted asLAOS: Riv. Bang Hieng, by Luna de Carvalho, 1989;here interpreted as Banggai I. / Banggi I. / Bang I., 07°159N, 117°109E, see Stork, 1986).

12. Protopaussussp.n.,SARAWAK (Borneo): Mts. Mulu (BMNH) (Stork, BMNH, in litt ., 1992) [Mts. Mulu: 04°029N, 114°549E].13. Protopaussus walkeriWaterhouse,CHINA: Fukien, Kuatun, 22.v.1946 (Tschung Sen) (NRS) (Luna de Carvalho, 1967) [CHINA: Fukien, Kuatun,

2300 m, 24.7°N, 117.4°E,testeVoss, 1949].14. Protopaussus walkeriWaterhouse,CHINA: Mont Combe, 30 miles SE of Ning–po, 1600 feet, 2.vi.1892 (J.J. Walker) (BMNH) (Waterhouse,

1897) [Ningpo: 29°549N, 121°339E].15. Protopaussus walkeriWaterhouse,CHINA: Chekiang, Hangtcheóu, A. Pichon 1925 (MNHN) (Deuve, MNHN,in litt ., 1994, see Luna de

Carvalho, 1967) [Hangchow: 30°189N, 120°079E].16. Protopaussus walkeriWaterhouse,CHINA: Chinkiang, 32°129N, 119°309E, 29.v.1924 (Elgin Suenson) (ZMK, vid. Nagel).17. Protopaussus bakeriHeller, PHILIPPINES: Luzon, Laguna Distr., Mons Makiling (C.F. Baker) (Heller, 1914; Schultze, 1916; Darlington, 1950)

[Mount Maquiling: 14°089N, 121°129E].18. Protopaussus bakeriHeller, PHILIPPINES: Mindanao, Butuan (BMNH) (Luna de Carvalho, 1989) [Butuan: 08°569N, 125°319E].19. Protopaussussp.n., INDONESIA: N–Sulawesi, Dumoga–Bone National Park, Project Wallace, 1985 (BMNH) (Stork, BMNH,in litt., 1992)

[Dumoga Bone Nat. Park: approximately 00°359N, 123°509E].20. Protopaussus javanusWasmann,INDONESIA: Java, Nongkodjadjar, 1200 m, i.1911 (E. Jacobson) (Veth collection, Leiden Museum ?)

(Wasmann, 1912) [E Java, near Tengger Mts., approximately 07°509S, 113°009E, see Reichensperger, 1935].21. Protopaussus kaszabiLuna de Carvalho,TAIWAN : Formosa, Takao, 25.vii.1907 (Sauter) (Nat. Hist. Mus. Budapest) (Luna de Carvalho, 1967)

[Takao/Takau/Kao-hsiung: 22°369N, 120°179E]


Documents

New fossil paussids from Dominican amber with notes on the phylogenetic systematics of the paussine complex (Coleoptera: Carabidae)