updated manuscript to PETALURA vol. 2,
      1996
    
    
    
    
    
      FOSSIL ODONATES IN TERTIARY AMBER
    
    
    
    
    
      Günter Bechly
    
    
    
    
    Dragonflies and damselflies are one of the most spectacular,
    but also one of the rarest, insect inclusions in Tertiary
    amber. Up to now there are no odonates known from any Mesozoic
    amber. Because of this rarity dragonflies are not even
    mentioned in a recent book on insects in amber (KRZEMINSKA
    & KRZEMINSKI, 1992). This paper is a short review and
    preliminary revision of all currently known specimens, based on
    a revised and amended version of BECHLY (1993). LARSSON (1978)
    explained the presence of odonate larvae or exuviae in Baltic
    amber. The fact that nearly all amber preserved odonates are
    damselflies, can be explained by the conditions of
    fossilisation: Damselflies (Zygoptera) get easier trapped and
    enclosed in resin than dragonflies (Anisoptera). Thus it is
    evident that Anisoptera preserved in amber will always be
    extremely rare and consequently of course also very expensive
    fossils (see ORR, 1993).
    The following damselfly-fossils from the Dominican amber are
    located in the paleontological "Museum am Löwentor
    des Staatlichen Museums für Naturkunde" in Stuttgart
    / Germany (SCHLEE, 1990 and pers. comm.1993):
    
      - 1.) A complete damselfly, which is not
      very well visible, because the insect is surrounded by
      dirt.
- 2.-4.) A relatively large piece of
      amber, containing three (!) damselflies.
- 5.) A fragmentary damselfly-wing.
- 6.) A fine preserved distal half of a
      damselfly-wing in very clear amber.
- 7.) A damselfly (head, thorax, legs,
      proximal abdomen and wing bases) in excellent condition,
      preserved in clear, polished amber!
Colour-photos of the last two mentioned specimens, probably
    Coenagrionidae, have been published in SCHLEE (1990: 83) (see
    fig. 1 and fig. 2). Unfortunately the magnificent
    amber collection of the Stuttgart museum is presently (fall
    1996, probably till end of 1997) not open for scientific
    studies, because of the unexpected retirement of Dr. Schlee.
    There are only three other records of odonate specimens from
    Dominican amber:
    
      - 8.) ORR (1993) reported about a
      libelluloid-like dragonfly completely preserved in Miocene
      amber, certainly from the Dominican Republic. Its present
      location unfortunately is unknown and it has apparently never
      been illustrated or described (ORR, pers. comm. 11/96). It
      would be very important to find out its present location (if
      any of the readers should known something about it, I would
      be most grateful for a short information by email to: bechly@mac.com),
      and to check if this remarkable specimen indeed represents a
      genuine Tertiary amber fossil or maybe just a more recent
      copal inclusion.
- 9.) DONNELLY (1993; and pers. comm.
      11/96) has a damselfly wing (similar to Telebasis)
      in amber from the Dominican Republic too. It is still in his
      private collection, but has never been illustrated or
      described yet.
- 
        10.) POINAR (1996) described a new species
        of the recent coenagrionid genus Diceratobasis
        (see fig. 16) from a piece of Dominican
        amber that is located in the private collection of Jim Work
        (Ashland, Oregon, USA). The larvae of this species probably
        lived in phytotelmata of tank bromeliads.
        
          - POINAR (1996: 382-383): "Diceratobasis
          worki sp. nov."
 
    The Dominican amber originated in the Middle Eocene to Upper
    Oligocene (45-25 Mio. years b.p.), maybe even to the Lower and
    Middle Miocene (15-20 Mio. years b.p.). The Dominican
    "amber-tree" most probably has been a member of the
    Recent genus Hymenaea (H. protera Poinar), a
    neotropical leguminous that is known for its high production of
    resin (SCHLEE, 1986). Amber from the Dominican Republic is
    extraordinary for the following reasons:
    
      - numerous places of discovery, with a large output of
      high-quality amber.
- regular findings of large pieces of amber, up to 13
      kg!
- frequent and diverse fossils in excellent condition:
      Plants (blossoms, leaves and bark), arachnids (incl.
      scorpions, amblypigids and pseudoscorpions), insects (incl.
      odonates, mantids, membracids, strepsipteres and fleas) and
      even vertebrates (mammal bones and bird feathers, as well as
      complete small frogs, gekkos and anolis-iguanas)! There are
      also single pieces of amber with mass-catches of insects,
      e.g. containing 2000 ants, or 1000 dolichopodid flies, or 15
      moths etc.
Therefore it is certainly not over-optimistic to expect
    further odonates from Dominican amber in the future, but there
    are also some fossil odonates known from the Baltic amber of
    eastern Europe, which originated in the Upper Eocene (about
    40-50 Mio. years b.p.) of Scandinavia, but is found in
    secondary deposits of glauconitic sands ("blue
    earth") of the Upper Eocene to Lower Oligocene (about
    30-40 Mio. year b.p.) at the Baltic coast. Based on the
    monograph of CONWENTZ (1890) the Baltic "amber-tree"
    has long been assumed to be an extinct conifer, which has been
    named Pinus (or Pinites) succinifera
    Goepp., although this taxon is still undefined and could
    include five different species (SCHLEE, 1986), however since
    the studies of KATINAS (1971) it is regarded as more likely
    that the Baltic amber was produced by a ceder (close to the
    Recent species Cedrus atlanticus) and maybe also an
    araucaria of the genus Agathis (= kauri-pine).
    
    HAGEN (1854) mentioned five odonates and HANDLIRSCH (1906-1980)
    mentioned six odonates from Baltic amber. Handlirsch's list
    was incomplete and contained several errors, which
    unfortunately have been frequently perpetuated, even recently
    by the author himself (BECHLY, 1993), although they had been
    corrected already by ANDER (1942). Unfortunately the Odonata
    chapter in the well known catalogue of amber fossils by
    KEILBACH (1982: 208-209) is likewise incomplete and incorrect,
    and furthermore even contains some additional errors too. The
    following new list will hopefully be rather complete and
    error-free:
    
      - 
        11.-12.) Two fossil damselflies of the
        famous "Koenigsberg amber collection" are now
        located in the "Geologisch-Paläontologisches
        Institut der Universität" Göttingen /
        Germany (species A: no. 3 B 696; and species B: no. K
        8088). These specimens were described and figured by PFAU
        (1975) (see fig. 3, fig. 4 and fig. 5). They are remarkably well
        preserved and seem to represent female specimens of two
        different Coenagrionoidea species of the Upper Eocene,
        which are still unnamed. Pfau's suggestion that these
        fossils might belong to the Recent family Platycnemididae
        is not really supported by the available evidence, so that
        these fossils should rather be regarded as Coenagrionoidea
        incertae sedis. According to Pfau one of the species might
        be conspecific with "Agrion antiquum"
        Hagen.
        
        
          - PFAU (1975: 1): "..., Zygoptera -
          möglicherweise Platycnemididae"
- KEILBACH (1982: 209): "Platycnemididae spec. A.
          u. spec. B."
- NEL & PAPAZIAN (1990: 254):
          "Platycnemididae, genre incertae sedis (Pfau,
          1975)"
- BECHLY (1993: 14): "... seem to represent two
          different platycnemidid species, which are still
          unnamed."
 
- 
        13.-17.) Four pieces of amber (with
        remains of five specimens of damselflies), of which three
        have been located in the "Collection Berendt" of
        the "Paläontologisches Museum des Museums
        für Naturkunde der Humboldt-Universität" in
        Berlin / Germany. PFAU (1975) could only locate the two of
        these pieces in the Berlin museum (see fig. 6 and fig. 7). The location of the
        fourth specimen is unknown. One of the two remaining
        specimens is one piece (no. 16) with a basal wing fragment
        and a male and a female abdomen (fig. 8). It probably represents
        the remains of a trapped pair, and the appendices of the
        male abdomen indicate that they belong to the same species
        as the second specimen (PFAU, 1975). The second piece
        contains a wing fragment without base and apex (fig. 9), head (fig. 10), all 6 legs but without
        coxae and tarsi (fig. 11), and the apex of a male
        abdomen. The labels for both pieces are marked with a red
        spot, which is indicating a status as original and/or type,
        and by the description there is no doubt that both pieces
        represent originals of HAGEN & PICTET (1856), so that
        these two pieces seem to represent syntypes of
        "Agrion antiquum". The female specimen
        described by PICTET (1856: 79) is apparently lost, as
        already supposed by PFAU (1975). Therefore I decided to
        designate the second specimen (original of HAGEN, 1856: 79)
        as lectotype of "Agrion antiquum", since
        even PICTET (1856: 79) already mentioned that the specific
        identity of piece no. 16 can not be decided.
        
          - 
            HAGEN (1848: 7): "Agrion antiquum
            Pictet" (nomen nudum; knows 2 specimens)
            
          
- HAGEN in SELYS (1850: 357-358): "Agrion
          ? antiquum Pictet"
- SELYS (1850: 366): "Platycnemis ?
          antiquum"
- HAGEN (1854: 227): "Agrion antiquum P.
          3."
- GIEBEL (1856: 273): "Agrion
          antiquum"
- PICTET in BERENDT (1856: 78-79, fig. 4a-d & 5 on
          tab. vi): "Agrion antiquum" (first
          description; knows 3 specimens)
- HAGEN in BERENDT (1856: 79-80, fig. 11a-c on tab.
          viii): "Agrion antiquum"
          (supplementary description; knows 4 specimens)
- KIRBY (1890: 175): "Coenagrion antiquum
          Hag."
- SCUDDER (1890: 127): "Platycnemis
          antiquum"
- HANDLIRSCH (1908: 899): "Platycnemis
          antiqua Hagen"
- ANDER (1942: 76): "Agrion antiquum
          Pictet et Hagen 1856 ... Die systematische Stellung
          dieser Art ist noch nicht entschieden"
- WEIDNER (1958: 52-53): "Platycnemis ?
          antiqua ?" (see no. 23.))
- PFAU (1975: 4): "Agrion antiquum Pictet
          (= Platycnemis antiqua Hagen)"
- LARSSON (1978: 83): "Platycnemis
          antiqua"
- KEILBACH (1982: 209): "Platycnemis
          antiqua (Pictet & Hagen, 1856)"
- NEL & PAPAZIAN (1990: 254): "Platycnemis
          antiqua (Pictet & Hagen, 1856)"
- BRIDGES (1993: VII.15): "Platycnemis
          antiquum (Pictet & Hagen), 1856"
- BECHLY (1993: 14): "Three
          Platycnemis-like damselflies, described by Hagen
          (1848, 1856) as Platycnemis antiqua."
- BRIDGES (1994: VII.16): "Platycnemis
          antiquum (Pictet & Hagen), 1856"
 
- 
        18.) A piece of amber with one basal and
        two apical fragments of the wings of a relatively large
        dragonfly (wing span about 3 inches according to Hagen) in
        the "Collection Menge". The present location is
        unknown.
        
        
          - HAGEN (1854: 227): "Aeschna.
          Flügelspitzen. - ... wahrscheinlich zu
          Gomphoides"
- HAGEN in BERENDT (1856: 81): "Gomphoides
          occulta Hagen" (nomen nudum; no valid
          description)
- KIRBY (1890: 168): "Gomphoides occulta
          Hag."
- HANDLIRSCH (1908: 900): "Gomphoides
          occulta Hagen"
- HANDLIRSCH (1921: 217): "Gomphoides
          Selys"
- ANDER (1942: 77): "Aeschnidae s.ll."
- FRASER (1957: 94): "... Gomphoides ...
          have also been reported from Bavarian amber and the
          Miocene."
- LARSSON (1978: 83): "Gomphoides
          occultus"
- KEILBACH (1982: 209): "Gomphoides
          occultus Hagen, 1856"
- CARPENTER (1992: 81): "Gomphoides
          SELYS-LONGCHAMPS in SELYS-LONGCHAMPS & HAGEN, 1850,
          p. 360 [Generic assignment of fossil doubtful.] PICTET
          & HAGEN, 1856."
- BRIDGES (1993: VII.167): "Gomphoides
          occulta Hagen, 1856"
- BECHLY (1993: 14): "An adult dragonfly,
          described by Hagen (1854, 1856) as Gomphus
          resinatus."
- NEL & PAICHELER (1994: 60): "Gomphoides
          occultus Hagen, 1856 (in Berendt :
          81)"
- BRIDGES (1994: VII.170): "Gomphoides
          occulta Hagen, 1856"
 
- 
        19.) A damselfly exuvia (Zygoptera) in the
        "Collection Berendt". My recent (November 1996)
        examination of the fossil, which is still preserved in the
        Coll. Berendt at the Natural History Museum
        (Paleontological Museum) in Berlin / Germany, basically
        confirmed the redescription and conclusion of HAGEN (1856)
        (see fig. 12 and fig. 13). It is most probably an
        exuvia of a Coenagrionoidea incertae sedis. If this fossil
        is conspecific with Agrion antiquum can neither be
        confirmed by positive evidence nor discarded by conflicting
        evidence. It seems to be very unlikely that this exuvia was
        embedded at the original site of emergence, because of the
        following reasons: The first evidence is the reasonable
        assumption that the conifer trees that produced the resin
        for the Baltic amber most probably were adapted for dry
        soils just like Recent conifers too, while damselfly larvae
        almost exclusively emerge on small plants that are very
        close to their breeding waters. The second evidence is the
        circumstance that all tarsi as well as the end of the
        abdomen with the caudal gills are missing. The most likely
        explanation seems to be, that an old exuvia was blown
        during a storm on a blotch of resin and became embedded. On
        the other hand it should be noted that two specimens of
        Gammaridae (Crustacea) are known from Baltic amber
        (BACHOFEN-ECHT, 1949 (reprinted 1996): 42-44), of which at
        least one was embedded when it was still alive. Some
        aquatic habitats thus must have been close enough to the
        amber trees that such purely aquatic animals could become
        embedded.
        
          - HAGEN (1848: 8): "Gomphus
          resinatus" (nomen nudum; no description)
- HAGEN in SELYS (1850: 358): "Gomphus
          resinatus Hagen, Nymphe"
- GIEBEL (1852: 639): "Gomphus resinatus
          Hagen"
- HAGEN (1854: 227): "Calopteryx ?,
          Nymphenhaut."
- GIEBEL (1856: 284): "Libellula
          resinata"
- PICTET in BERENDT (1856: 78 and 80, fig. 6 on tab.
          vi): "Gomphus - (larva)" (first
          description)
- HAGEN in BERENDT (1856: 80, fig. 12 on tab. viii):
          "Agrionide" (supplementary description; the
          term "Larva" in the explanation of fig. 12
          refers to Agrion antiquum)
- KIRBY (1890: 168): "Aeschna resinata
          Hag."
- HANDLIRSCH (1908: 896): "Calopteryx ?
          (larva) Hagen"
- HANDLIRSCH (1908: 900): "Gomphus
          resinatus Hagen"
- HANDLIRSCH (1908: 900): "Gomphus -
          (larva) Hagen"
- HANDLIRSCH (1921: 217): "Calopterygidae,
          zweifelhafte Larve"
- HANDLIRSCH (1921: 217): "2 als Gomphus
          bezeichnete Formen"
- ANDER (1942: 76): "Agrioniden-Larve 1"
- FRASER (1957: 94): "... Gomphus ...
          have also been reported from Bavarian amber and the
          Miocene."
- KEILBACH (1982: 209): "Agrion spec.
          Hagen in Berendt 1856, p. 80"
- KEILBACH (1982: 209): "Gomphus Larva
          Hagen in Berendt 1856, p. 80"
- KEILBACH (1982: 209): "Gomphus
          resinatus Pictet, 1856 in Berendt, p. 81"
- CARPENTER (1992: 81): "Gomphus LEACH,
          1815, p. 37. [Generic assignment of fossil (nymph)
          doubtful.] HAGEN, 1848."
- CARPENTER (1992: 87): "Calopteryx
          LEACH, 1815, p. 137. ..... HAGEN, 1848, ... . Oligo.,
          Europe (Baltic), ..."
- BRIDGES (1993: VII.196): "Gomphus
          resinatus Hagen, 1848"
- BECHLY (1993: 14): "A damselfly-larva (!)
          described by Hagen (1854) as
          Calopteryx."
- BECHLY (1993: 14): "Another adult dragonfly,
          described by Hagen (1848, 1856) as Gomphus
          resinatus."
- BECHLY (1993: 14): "A dragonfly larva (!),
          described by Hagen (1856) as Gomphus."
- NEL & PAICHELER (1994: 57): "Gomphus
          resinatus Pictet, 1856 (in Berendt, 1856:
          81) (d'après Keilbach, 1982) ... Elle doit
          être considérée comme un Odonata
          Gomphidae (?) de position incertaine."
- NEL & PAICHELER (1994: 57):
          "Gomphus "larva" Hagen, 1856
          (in Berendt) ... Son attribution est très
          doubteuse."
- BRIDGES (1994: VII.200): "Gomphus
          resinatus Hagen, 1848"
 
- 
        20.) Another Zygoptera-larva in the
        "Collection Hagen". The author could not find
        this specimen in the amber collections of the Museum of
        Comparative Zoology in Cambridge (laboratory of late F.M.
        Carpenter), although the complete Coll. Hagen is presently
        located in this institution.
        
          - HAGEN in SELYS (1850: 357): "Agrion ..... Une
          petite nymphe, ou plutôt l'étui
          vide."
- GIEBEL (1856: 273): "Eine unvollständige
          kleine Larve"
- ANDER (1942: 76): "Agrioniden-Larve 2"
 
- 
        21.) An odonate "larva"
        (certainly an exuvia) of uncertain affinities (location:
        unknown; lost according to ANDER, 1942):
        
      
- 22.) An undescribed abdomen of a female
      damselfly (completely preserved, incl. ovipositor) in the
      collection Bachofen-Echt (Fach 2 L1A) at the "Bayerische
      Staatssammlung für Paläontologie und historische
      Geologie" in Munich / Germany (fig. 14).
- 23.) A completely preserved damselfly in
      clear amber which is mentioned and figured by BACHOFEN-ECHT
      (1949 (reprinted 1996): p. 78 and fig. 63) (see fig. 15). The specimen is cited as
      Agrionidae (also see WICHARD & WEITSCHAT, 1996: 26), but
      without doubt represents the only known Lestidae in amber,
      since the specimen has wings with an oblique vein and two
      cells beneath the distinctly braced pterostigma (clearly
      visible in the illustration of the original edition, but not
      well visible in the reprint). I could not find the specimen
      in the collection Bachofen-Echt in Munich (BSGPM), thus its
      present deposition unfortunately has to be regarded as
      unknown.
- 24.) WEIDNER (1953) reports about a
      Zygoptera "larva" (Platycnemis ?
      antiqua ?) in a piece of Tertiary Baltic amber in
      the Coll. Scheele (Nr. 1082. Typ. Kat. Nr. 45.) of the
      "Geologisches Staatsinstitut Hamburg" /
      Germany.
- 25.-29.) Mr. Walter Ludwig (Berlin,
      Germany) reported (pers. comm. 1996) about a damselfly wing
      in a piece of Saxonian amber from Bitterfeld in the private
      collection of Mr. Hans Werner Hoffeins (Hamburg), and a
      complete damselfly in Baltic amber in the collection of a
      fossil trader in Berlin (Mr. Kühn). Furthermore he has 3
      specimens in his private collection (Coll. Ludwig, Berlin).
      One specimen is a fragment of a damselfly thorax (open so
      that one can see inside!) with two wing bases and a nearly
      complete wing of a stem-group representative of Euphaeidae
      (see fig. 17). It is very similar to the
      genera Parazacallites and Litheuphaea, and
      is sharing with the latter the complete suppression of any
      secondary antenodals between ScP and RA (contrary to the
      original description of Litheuphaea). Another
      specimen is absolutely unique, since it shows a damselfly
      that is just emerging from its exuvia, both completely
      preserved in a "Schlaube" (see fig. 18). The wings of the imago are
      not yet unfolded and the apex of the abdomen is still inside
      the exuvia. The exuvia has saccoid gills with a long
      filamentous apex, very similar to some Recent tropical
      damselflies (e.g. the two megapodagrionid genera
      Heteragrion and Oxystigma, the platystictid
      genus Palaemnema, the amphipterygid genus
      Diphlebia, and the Euphaeidae). Both mentioned
      specimens are from Baltic amber and will be formally
      described by the author. The third specimen is a piece of
      amber with head (with deeply fissured labium) and one fore
      leg (with cleaning "brush") of a damselfly from the
      Saxonian amber of Bitterfeld (Eastern Germany). According to
      WEITSCHAT (1996) the amber of Bitterfeld is of the same age
      and origin as the Baltic amber, not of Lower Miocene age
      (about 22 mybp) as previously believed.
- 30.) Mr. Hans Lüdicke (Kronberg,
      Germany) has a well preserved and nearly complete damselfly
      (about 4 cm long) from the Baltic amber in his private
      collection. This specimen is currently studied by Prof.
      Rainer Rudolph (Münster, Germany).
- 31.-32.) Two damselflies from the Baltic
      amber are present in the collection of the amber-museum at
      Ribnitz-Damgarten (East Germany). According to Mr. Ulf
      Erichson (pers. comm. 1997) one specimen is a wing and a body
      fragment in a piece of amber that was later manufactured as
      piece of jewellery. The second specimen is a rather complete
      damselfly, however the piece of amber contains some dirt and
      unfortunately had to be glued after it was accidently
      broken.
    All these fossils together make a total of at least 32
    different specimens. The present location of 25 specimens is
    known to me, and except two, all others are preserved in
    Germany, which therefore can be considered as "El
    Dorado" for researches on amber dragonflies. Anyway my
    enumeration will probably still be somewhat incomplete, since
    it is quite likely that at least a few specimens have
    disappeared in private collections without having been noticed
    by scientists. A few small damselflies are rumoured to be
    present in local collections in the Dominican Republic. HAGEN
    (1856: 78) mentions the existence of a further imaginal
    damselfly (Coll. Saturgus / Königsberg) and an odonate
    larva (Kabinett physik.-oekonom. Gesellschaft zu
    Königsberg), both from Baltic amber of course.
    Nevertheless it can not be excluded that these two specimens
    might be identical with specimens already mentioned in this
    enumeration. The same refers to the specimen illustrated in
    WICHARD & WEITSCHAT (1996: 79, Taf. 3) and the specimen
    illustrated in GRIMALDI (1996; also available on a website of
    the American
    Museum of Natural History). The former specimen (see fig. 19) is a pair of damselfly wings
    (apparently Lestidae!) in Baltic amber, while the latter
    specimen is a nearly complete damselfly in Dominican amber (see
    fig. 20), which is in a private collection
    (POINAR, pers. comm. 1997). All the known specimens are in need
    of a thorough revision, because their taxonomic and
    phylogenetic status seems to be more or less unsettled. Such a
    revision is projected by the author.
    
    
    
    
      REFERENCES
    
    
    ANDER, K. (1942): Katalog der Odonaten des
    baltischen Bernsteins. p. 76-77 in: ANDER, K.
    (1942): Die Insektenfauna des baltischen Bernsteins nebst damit
    verknüpften zoogeographischen Problemen. - Acta
    Universitatis Lundensis (= Lunds Universitets
    Årsskrift), N.F. Avd. 2., 38(4): 83
    pp., maps 1-10, tables 1-8 (also published in Kungl.
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    BACHOFEN-ECHT, A. (1949): Der Bernstein
    und seine Einschlüsse. Springer Verlag: Wien. 204
    pp.
    BECHLY, G. (1993): Fossil odonates in
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    BERENDT, G.C. (1830). - Ins.
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    concerning odonates preserved in amber)
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    IL, USA. 806 pp.
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    DONNELLY, T.W. (1993): [untitled note]. -
    Argia 4(4): 13
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    GIEBEL, C.G.A. (1852): Deutschlands
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    HAGEN, H. (1848): Die fossilen Libellen
    Europas. - Stett. ent. Z., 9:
    6-13
    HAGEN, H. (1854). Ueber die Neuropteren der
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      Last Update: 25th July, 2005
      © Günter Bechly,
      Böblingen, 2005