SlideShare ist ein Scribd-Unternehmen logo
1 von 12
Downloaden Sie, um offline zu lesen
Fossils Attributed to the Orchidaceae

                     RUDOLF SCHMID AND MARVIN J. SCHMID1

INVESTIGATORS INTERESTED IN the evolutionary history of a family
desire to have information from the fossil record to provide
(hopefully) unequivocal evidence for the early history of that
family. Unfortunately for orchidologists, the known fossil record
of the Orchidaceae is extremely meager. We decided to summarize
what is known of the fossil record of orchids (a) because there are
no accounts available other than the rather superficial ones by
Darrah (15) and Krackowizer (27a, 28) , (b) because there are number
of misconceptions in the literature that should be corrected,
and, most importantly, (c) because a reasonably complete list of
fossils attributed to the Orchidaceae could be prepared since we had
access to the Compendium Index of Paleobotany (see 3, 19 for
accounts of its coverage) and to libraries with extensive holdings
in rather obscure publications.


Most persons seem to favor a relatively great age for the
Orchidaceae (e.g., 1, 5, 6, 12, M, 15, IS, 21-23, 48), but relatively
few have ventured a specific time and/or place of origin. Garay (21-
23), following Stebbins (48, pp. 501-502), postulates an origin in
the early Cretaceous- and in addition proposes Malaysia as the
most likely cradle of orchidhood. Leon Croizat (pers. comm.,
1972) believes that the orchids arose "surely not later than the
earliest Cretaceous. Brieger (5, 6) favors the early Tertiary and the
"united Asiatic-American" tropics (6, p. 329, specific area not
indicated). In contrast, some authors, notably Schultes, believe
that the orchids may be "a comparatively young group" (44, p. 1;
45, p. 1043). While there is some dispute as to the exact time of
origin of the family, nearly all workers (e.g., 5, 16, 17, 21. 40, 44)
seem to agree that currently, and in the immediate past, the
Orchidaceae are in a very active period of evolution.

Most workers have indicated either that there is no fossil record for
the Orchidaceae (e.g., 1, 6, 10, 12, 13, 20, 24) or that there are
only doubtful orchidaceous fossils (e.g., 15, 16, 18, 21, 25, 32-34,
36, 39, 40, 42, 44, 45). In contrast, a few persons (e.g., I I , 26, 28,
35, 38, 43, 49, 50, 52) have stated that valid orchid fossils exist.
Most of these authors apparently based their view concerning the
fossil record of the Orchidaceae chiefly or only on knowledge of the
very dubiously orchidaceous Protorchis monorchis and Palaeorchis
rhizoma described by Massalongo (31-34) from the Eocene of
Italy. 4 Nevertheless, a number of other fossils have also been attributed to
the Orchidaceae, most significantly Straus' (49, 50) three species of
putative orchid fruits (Orchidacites) from the Pliocene of Germany. With
the exception of Gothan and Weyland (26), Kirchheimer (27), and Melchior (35),
however, most recent workers seem unaware of Straus' finds. These and other
taxa will be discussed in detail below.
Table 1. Geologic time scale (pre-Mesozoic omitted). (After Hartland, et.al.
       26a)

Era                        Period               Epoch                   Beginning of interval
                                                                        (in millions of years

                                                Holocene (Recent)       0.005

                           Quaternary           Pleistocene (Glacial)   2.5

                                                Pleiocene               7

                                                Miocene                 26

                                                Oligocene               38

                                                Eocene                  54

Cenozoic                   Tertiary             Paleocene               65

                           Cretaceous                                   136

                           Jurassic                                     190

Mesozoic                   Triassic                                     225

       Orchids are not favorable candidates for fossilization, an obvious conclusion
       that has not escaped previous authors (e.g., 15, 18, 28, 40. 44). The following
       characteristics of most Orchidaceae probably account for their scarcity as
       fossils: (a) predominant occurrence, both in the present and presumably in the
       distant past, in the wet tropics, which are areas of rapid decay; (b) herbaceous
       habit; (c) epiphytic habit, which would generally preclude orchids from the
       conditions (usually aquatic) most conducive to fossilization (see also 15) ; (d)
       production of pollinia (usually) rather than individual pollen grains, and
       dispersion of the former by animal vectors instead of wind: and (e) minute,
       easily degradable seeds.

       Krackowizer (28) and Schimper and Schenk (43), however, apparently
       believed a rather extensive fossil record of the Orchidaceae is to be expected,
       and Darrah (15) and also Krackowizer (28) suggested that fossil orchids might
       eventually be encountered in deposits in tropical areas when these become
       better known.

       Fossils that have been attributed to the Orchidaceae (or to the
       Protorchidaceae) are strictly megafossils (e.g., fruits, leaves, tubers, etc.);
       orchidaceous microfossils (e.g., seeds and pollen) have not been reported in the
       literature. Discoveries of cuticular remains (as those already found of the
       Pliocene Orchidacites wegelei of Straus, HI) perhaps offer the best hope for
       significant additions to the fossil record of (he Orchidaceae.

       Although there is no record of fossil orchid pollen, even if orchid pollen
       were preserved as fossils, it is a moot point that it would be recognizable as
       such. Botanists simply might not recognize the fossilized pollen of those
       orchidaceous forms that had not yet evolved pollinia. Perhaps significantly, the
       Asclepiadaceae, which like the Orchidaceae possess pollinia, are not listed in
       Potonie's (41) recent compendium of fossil pollen and spores. Although
       Chandler's (9, and works cited therein) extensive investigations (initially with
       the late Eleanor M. Reid) of the Tertiary London Clay Flora of England
       over a period of several decades failed to reveal any orchidaceous remains,
she suggested (8, p. 29) that "possibly search for pollen among the finer
sediments and residues may eventually demonstrate the presence of this family"
in the London Clay Flora. Subsequent palynological work (two 1961 Ph.D.
theses by Ma Khin Sein and Jane Pallot at the University of London, both
cited in 9, the latter published as 30) , however, has thus far failed to
substantiate this prediction.

As recently discussed by Eyde (19), there are several paths into the
paleobotanical literature. We checked a variety of sources for records of
orchidaceous fossils, most importantly the index by Andrews (3, including
unpublished cards for additions since 19(>5) and the United States Geological
Survey's Compendium Index of Paleontology, an unpublished file available for
consultation only in the Natural History Building, Room W-300, of the
Smithsonian Institution, Washington, D. C. (see 3, 19) . We checked the
Compendium Index for most of the temperate genera listed in Schultes and
Pease (46). Most tropical genera, however, were not sought in the
Compendium Index due to the minuscule yield that could only result from
such a mountain of effort. As noted above, tropical plants are unlikely
candidates for fossilization. In addition, most tropical orchids are endemics
that presumably evolved during the Quaternary, and hence any fossils of them
would be unlikely to be encountered by paleobotanists, most of whom have
worked (until very recently, at least) in temperate areas. Finally, any
orchidaceous fossils of the pre-Quaternary tropical floras of presently temperate
areas would probably be given generic names not based on living taxa.

                                                         5
                      REPORTS OF ORCHIDACEOUS FOSSILS


Three extinct genera (Palaeorchis, Protorchis and Orchidacites) have been
designated as orchidaceous or protorchidaceous. At least one other fossil taxon,
Antholithes pediloides, has been regarded as an orchid. In addition, fossil
remains from the Quaternary have been attributed to a number of extant,
north temperate orchid taxa. Purists who restrict "fossils" to pie-Quaternary
remains and therefore regard Pleistocene finds as "subfossils" may object to the
inclusion of plant remains from the Quaternary in the following enumera-
tion:
                                    Jurassic

   The Compendium Index of Paleobotany, citing Thurmann (51) , lists the
following extant species as occurring in the Jurassic strata at Porrentruy,
France: Ophrys myodes Jacq., Orchis morio L., O. pyramidalls I,., Satyrium
viride L., and Serapias rubra L. This is incorrect. Thurmann did indeed list
these species, but only as part of the modern vegetation of this region.
                                     Eocene

Protorchis monorchis Massalongo (31-33) and Palaeorchis rhizoma (.Massalongo)
Massalongo (32) (= Protorchis rhizoma Massalongo, 31): These species represent
the first described and also the geologically oldest fossils that might possibly
represent orchids. In 1857 Massalongo (31) listed, without benefit of description
or illustration (hence nomina nuda) , the new generic name Protorchis, with
two new species P. monorchis and P. rhizoma based on specimens from the
calcareous Eocene deposits at Monte Bolca, Italy. Massalongo had only
four specimens at his disposal — three of the former species, one of the latter
(33, 34). In 1858 Massalongo (32) validly published Protorchis monorchis and
also the new combination Palaeorchis rhyzoma (the specific epithet is an
orthographic error) based on Protorchis rhizoma. The next year a more
complete description and also a photograph (see Fig. 1) of Protorchis monorchis
were published (33). In his 1858 work Massalongo dated Protorchis as "1851"
with the added notation "in lit.” et in Musaeo" (32, p. 749). Since the 1854
reference is obviously unpublished, the nomina nuda in the 1857 report (31)
thus represent the actual first (though taxonomically invalid) publication of
the names involved.


Although Massalongo (31) initially listed his new species under the
Orchidaceae, he subsequently (32-34) very carefully indicated the tentatively
orchidaceous nature of his fossil specimens by including them in a new taxon,
the Protorchidaceae ("Protorchidee" in 32, 33; or the Latinized
"Protorchideae" in 33, 34). Massalongo (32, 33) admitted that he was unable
to find in the extant flora counterparts of his fossils. Massalongo (32, 33) noted
a resemblance of both his fossil species to the Araceae, which he apparently
regarded as being rather close to the Orchidaceae (32). A superficial likeness
between Palaeorchis rhizoma and the fossil alga Delesserites was rejected when
he decided on the affinities of the former (32). After additional concern that
Palaeorchis rhizoma might be butomaceous, Massalongo (32) finally decided
to retain this species in his Protorchidaceae. This discussion illustrates
the difficulty Massalongo had in assigning his fossils to an extant plant
group. This fact is apparently realized by very few authors since a number
(e.g., 15, 18, 28, 42) incorrectly state or imply that Massalongo had regarded
his fossils as orchids.
According to Massalongo (32, descriptive terminology below is his) , the
Protorchidaceae are next to the orchids and the aroids and consist of small
herbs with tubers or rhizomes bearing lateral fibrous roots and several very
slender, cuneate-obovate or spathulate leaves with entire margins and fine
midribs. Misstatements to the contrary (15, 25, 26, 28, 43), both species do not
possess tubers. Protorchis monorchis (FIGURE 1) has a round, solitary tuber
whereas Palaeorchis rhizoma (never illustrated by Massalongo) differs chiefly
in having a perpendicularly cylindrical rhizome covered with circular,




papillate leaf scars (32). The two species also differ somewhat in having
spathulate versus oblong to spathulate leaves, respectively (32). Massalongo
(33) subsequently indicated that Protorchis monorchis, of which three speci-
mens were available (33), strictly speaking does not have a true tuber, but rather
a rounded rhizome (FIGURE 1). In the same publication he (33) also added the
following information for Protorchis monorchis: tuber 7-8 mm in diameter;
leaves 3-4 per plant, attenuate into a petiole, 5 cm long, and 12-15-18 mm
wide.11 Although Massalongo (34, p. 133) finally indicated that specimens of
both this species and Palaeorchis rhizoma are seedlings, there is no mention in
his previous descriptions (32, 33) of the probable developmental age of these
fossils.


Most workers, usually referring only to Protorchis monorchis, have
subsequently concluded that Massalongo's fossils are not truly representative
of the Orchidaceae (e.g., 15, 18, 21, 25, 26, 35, 39, 40, 42). Schimper and Schenk
(43) , however, accepted Massalongo's finds as orchidaceous, and van der Pijl
(39, 40) apparently seems tempted to accept Protorchis monorchis as validly
orchidaceous, no doubt because its Eocene date ties in with his understanding
of the evolution of the bees. Meschinelli and Squinabol (3i) included both of
Massalongo's fossil species under the Protorchidaceae in the order Micro-
spermae (= Orchidales), but these authors noted that Palaeorchis rhizoma is
probably a member of the Butomaceae. Krackowizer (28) 7 accepted the views of
Meschinelli and Squinabol (36) except that he regarded Protorchis monorchis
as a true orchid rather than as a protorchid. Admitting that both of
Massalongo's fossils are doubtfully orchidaceous, Leslie A. Garay (pers. comm.,
1972) nevertheless maintains that of all the fossils attributed to the
Orchidaceae, Protorchis monorchis is perhaps the most likely candidate for
inclusion in the family, largely because of its similarity to Orchis pallens L.

In conclusion, the orchidaceous nature of Massalongo's fossils is clearly very
questionable. As has already been suggested (Chester A. Arnold, pers. comm.,
1967; 15, 18), perhaps the most charitable tiling that can be said about the
affinity of Protorchis and Palaeorchis is that they are monocotyledonous.

                                    Oligocene

Antholithes pediloides Cockerell (1 1): T. D. A. Cockerell, the prolific
describer of fossils from the western United States,8 in 1915 delineated from
the Lower Oligocene (the age according to MacGinitie, 29; incorrectly
regarded as Miocene by Cockerell, I I ) beds at Florissant, Colorado, a new
species in the fossil artificial (or form) genus Antholithes. Cockerell attributed
the fossil (FIGURE 9), A. pediloides, to the Orchidaceae because of its marked
resemblance to the lip of Cypripedium, and he also presumed that the several
small "subhyaline" spots scattered over the surface might represent the work
of some insect. Other than the suggestive outline of the fossil, however, the lack
of significant detail makes Cockerell's determination extremely doubtful. In
his classic flora of the Florissant beds, MacGinitie (29) reached the same
conclusion and disposed of A. pediloides among "species of somewhat doubtful
taxonomic value" (p. 159) under incertae sedis
.
                                       Miocene

Darrah (15) briefly discussed, and then discounted as truly orchidaceous, a
fossil stem (apparently unnamed) from the Miocene of Hungary that had been
described by a Robert Brown (there were several Robert Browns). Since
Darrah provided no references in his note, and since alter considerable
searching we have been unable to locate any additional information
concerning this fossil, we can only quote Darrah (15, p. 149) fully:
        "A third form [besides Massalongo's Protorchis and Palaeorchis] was
     once provisionally accepted as a fossil orchid. This fossil stem, found in
     rocks of Miocene age in Hungary, included a few structurally preserved
     hair-like roots which Robert Brown considered to be of some epiphytic:
     orchidaceous plant. . . .
       As a matter of fact it was with this organ [the pseudobulb] that Robert
     Brown attempted to compare his supposed fossil from Hungary."
Pliocene

Orchidacites orchidioides Straus (49), O. wegelei Straus (49), and O.
cypripedioides Straus (50): In 1954 Straus (49) described from the Upper
Pliocene of Willershausen, Germany, two species of fruits, Orchidacites
orchidioides and O. wegelei which he assigned to the Orchidaceae. The two
species, especially the former (as suggested by i t s name), were thought to
resemble various species of Orchis (49). More recently, Straus (50) provided
for Orchidacites a generic diagnosis, which had been omitted from the 1954
report, and described a third species, O. cypripedioides, with fruits regarded
as similar to those of Cypripedium. These taxa are illustrated in FIGURES
2 to 8, reproduced from Straus' more recent paper (50). Orchidacites
is a form genus proposed for fossil fruits comparable to the capsules
of various extant orchid genera (50). According to Straus (50), the
fossil capsules, 1.5 to 2.5 cm long, are ellipsoidal or narrowly ellipsoidal and
have several longitudinal striae (FIGURES 2-8); the remnants of a corolla often
occur at the fruit apex (FIGURES 2-4).

Straus (50, also pers. comm., 1972) believes that the fossil fruits of
Orchidacites came from epiphytes growing on rotting branches that eventually
were blown into the sediments by wind, and, as a consequence, he has
speculated (50, pers. comm., 1972) that many of the present-day orchids (e.g.
Limodorum, Neottia, Corallorrhiza, and Cypripedium) were primitively
epiphytic and now are terrestrial "secondary relicts." This view, of course, is
dissonant with the conventional one that the terrestrial habit is ancestral and
the epiphytic derived (e.g., 5, 6, 16, 17, 21-23, 39, 42).

The Straus fossils have received little comment from either orchidologists or
paleobotanists. Melchior (35) and Gothan and Weyland (26) accepted the
fossils as unmistakably orchidaceous. Kirchheimer (27, p. 650), however,
remained skeptical, believing that the inferior, wing like, ribbed gynoecium
with a distinct styloid process evokes resemblances to young fruits of Halesia,
Pterostyrax and other Styracaceae (a completely unrelated family in the
dicotyledons). Straus (pers. comm., 1972), in counterargument, however,
believes that the fruits he described are truly orchidaceous since fruits of the
Styracaceae never show floral remains whereas fruits of the Orchidaceae often
do.

On examining the photographs reproduced herein ( FIGURES 2-8), Garay
(pers. comm., 1972) is also disinclined to accept Straus' fossils as orchidaceous
because of the curious 3-pronged floral remnants (interpreted by Garay as
a column) ( FIGURES 2-4) and because of the apparently excessive number of
ribs ( FIGURES 2-8) for true orchid fruits (which have a maximum of six).
Robert L. Dressier (pers. comm., 1972) is of a similar opinion, although he is
less certain in excluding Orchidacites cypripedioides (FIGURES 5-8) from the
orchids since the fossil "looks rather like a Cattleya fruit."

In defense of Straus, we should note that a 3-pronged calyx of fused sepals
occurs in some modern taxa (e.g. Pterostylis. see FIGURE 84 in 40: Masdevallia,
etc.) and that on orchid fruits a greater number of ribs (than six) may be
apparent since these may be variously secondarily divided (e.g., Trichopilia
suavis Lindl. et Paxton). Unfortunately, Straus (50) hurts his own cause by
interpreting the floral remnants (FIGURES 2-4) as a corolla, but it is perhaps
more likely that they represent a calyx.
                                   Quaternary

Quaternary orchid fossils are included here for completeness, although they
are unimportant from the viewpoint of our understanding of the origin and
most of the subsequent evolution of the family. The names of at least 20 extant
species of orchids are listed in the Compendium Index of Paleobotany and are
attributed to both the Pleistocene and Holocene (= Recent or Postglacial) of
the Quaternary. Most of these listings were compiled around the turn of the
century, when the Compendium Index included casual, incidental references
to fossils - a practice long discontinued ( 1 9 ). Unfortunately, a number of
these listings are not applicable because the original works discuss the various
orchid species as components of the contemporary flora and not as fossils. This
is the case with reports of Goodyera repens (L.) R. Br. from the Quaternary of
Denmark, Himantoglossum hircinum (L.) Sprengel and Ophrys aranifera
Huds. from the Postglacial of Switzerland, and Malaxis paludosa (L.) Sw. from
the Holocene of Germany, which the Compendium Index attributes to
Anderson (2), Naegeli (37, as cited in 7), and Becker (4), respectively.

The Compendium Index also attributes the following extant orchid taxa to
Sernander's (47) extensive work on the Quaternary (Wiirm Glacial and
Postglacial) of Gotland, Sweden (names listed as they appear in Sernander):
Anacamptis pyramidalis (L.) Rich., Cephalanthera ensifolia (Sw.) Rich.,
Corallorrhiza innata R. Br., Epipactis palustris (L.) Crantz, Gymnadenia
conopsea (L.) R. Br., G. odoratissima (L.) Rich., Listera cordata (L.) R. Br.,
L. ovata (L.) R. Br., Malaxis monophyllos (L.) Sw., Neottia nidusavis (L.)
Rich., Orchis angustifolia Wimm. et Grab., O. maculata L., O. militaris L., O.
ustulata L., Platanthera bifolia (L.) Rich., and Sturmia loeselii (L.) Reichb.
However, Sernander (47) merely discusses these and other orchid species in
terms of a phytosociological survey of the modern bog vegetation of Gotland.
The bogs Sernander studied did indeed contain identifiable fossils, but none
of these were orchids.

To our knowledge, there is only one report of a Quaternary fossil attributed to
the Orchidaceae. In 1965 Vent (52, p. 200) described leaf and fruit
impressions from the Riss-Wurm Interglacial of Weimar-Ehringsdorf, Germa-
ny, and assigned these to "cf. Epipactis palustris (Mill.) Crantz" in the
Orchidaceae. Garay (pers. comm., 1972) , however, discounted the orchidaceous
nature of these fossils after examining Vent's photographs (FIGURE 10) .
                                   SUMMARY

Fossils dating from the Eocene to the Quaternary have been attributed to the
Orchidaceae, but objections have been raised against the orchidaceous nature
of all of these fossils. The most likely orchid fossils, nevertheless, remain
Massalongo's famous fossils from the Eocene of Italy — Protorchis monorchis
and Palaeorchis rhizoma — and especially Straus' recent finds from the
Pliocene of Germany Orchidacites orchidioides, O. wegelei and O.
cypripedioides. In conclusion, then, the Orchidaceae have no positive fossil
record and in this sense present a striking parallel to two well-known gods of
mythology: Athena, who sprang fully grown and fully armored from the head
of Zeus; and the Aztec Huitzilopochtli, who was borne fully grown and fully
armored from Coatlicue.
Acknowledgements: This study was carried out while the senior author was the
recipient of a Smithsonian Institution postdoctoral fellowship. We thank Norris
H. Williams and Leslie A. Garay for valuable discussions.
                                 REFERENCES

 (1) Ames, O. and D. S. Correll. 1952-53. Orchids of
 Guatemala, Heldiuna: Rot. 26:-i-xiii. 1-727.
 (2) Andersson, G. 1906. Die Entwicklungsgeschichte der
 skandinavischen Flora. Pp. 45-97 in Resultats Set. Congr.
 Int. Dot., Vienne, 1905.
 (3) Andrews, H. N., Jr. 1970. Index o£ generic names of
 fossil plants, 1820-1965. U.S. Geol. SURV. Bull.. 1300.
 (4) Becker. G. 1874. Botanische Wanderungen durch die
Sümpfe und Torfmoore der Niederrheinischen Ebene. Verh.
  Nalurhist. Vereines Preus* Rheinl. Westphallens 31:137-158.
  (5) Brieger, F. G. 1958. On the phytogeography of orchids.
  Pp. 189-200 in Proc. 2nd World Orchid Conf., Honolulu,
  1957.
  (6) Brieger, F. (1960. Geographic distribution and
  phyllogeny [sic] of orchids. Pp. 328-333 in Proc. 3rd World
  Orchid Conf.. London. I960.
  (7) Brockmann-Jerosch, H. 1910. Die Änderungen des Klimas
  seit der grösstcn Ausdehnung der letzten Eiszeit in der Schweiz.
  Pp. 55-71 in Die Veränderungen des Klimas seit dem
  Maximum der letzten Eiszeit, Ber., Exekutivkomitee
  I I . Int. Geol.-Kongr., Stockholm. 1910.
   (8) Chandler. M. E. J. 1951. The Lower Tcrtiary floras
  of southern England. I. Palaeocene floras: London Clay
  Flora (supplement). London: British Museum (Natural
  History). [Text and plates separately bound.]
  (9) Chandler, M. E. J. 1964. Idem. IV. A summary and
  survey of findings in the light of recent botanical
  observations. London: British Museum (Natural
  History).
 (10) Chesters, K. I. M., F. R. Gnauck and X. F. Hughes.
 1967. Angiospermae. Pp. 269-288 in V. B. Harland et al.
 [eds.], The fossil record. London: Geological Society of
 London.
 (11) Cockerell. T. D. A. 1915. Notes on orchids. Bot. Gaz.
 59:331-333.
(12) Correll, D. S. 1950. Native orchids of North
America. Waltham, Mass.: Chronica Botanica Co.
 (13) Coulter, J. M. and C. J. Chamberlain. 1903.
 Morphology of angiosperms. New York: II. Appleton and
 Co.
 (14) Croizat, L. 1961. Principia botanica. Caracas: The
 Author. [1 vol. in 2.] [Published 1961.]
 (15) Darrah. W. C: 1940. Supposed fossil orchids. Amer.
 Orchid Soc. Bull. 9:149-150.
 (16) Dodson. C. H. and R. J. Gillespie. 1967. The biology
 of the orchids. The Mid-America Orchid Congress. [No city
 of publication given.]
 (17) Dressler. R. L. and C. H. Dodson. 1960.
 Classification and phytogeny in the Orchidaceae. Anu.
 Missouri Bot. Gard. 47:25-68.
 (18) Dunsterville, G. C. K., and L. A. Garay. 1959.
 Venezuelan Orchids Illustrated. Vol.1. London: Andre
 Deutsch. [Also Introduction in Spanish in Vol. 2. 1961.]
 (19) Eyde. R. H. 1972. Note on geologic histories of
 flowering plants, Brittonia 24:111-116.
 (20) Andreanszky, G. 1954. Osnövénytan. Budapest.
 Akadémiai Kiado.
(21) Garay, L. A. 1960. On the origin of the
Orchidaceae. Bot. Mus. Leaflets Harvard Univ. 19:57-96
[Also in Proc. 3rd World Orchid Conf., London. 1960, pp.
172-196.]
(22) Garay, L. A. [1964.] Evolutionary significance of
geographical distribution of orchids. Pp. 170-187 in Proc.
4th World Orchid Conf,.Singapore, 1963.
(23) Garay, L. A. 1972 On the origin of the Orchidaceae,
II. J. Arnold Arb. 53:202-215.
(24) Godfrey. M. J. 1933. Monograph & Iconograph of
native British Orchidaceae. Cambridge: University Press.
(25)    Gothan, W. 1921. H. Potonié’s Lehrbuch der
Paläobotanik. 2. Aufl. Berlin: Gebröder Borntraeger.
(26) Gothan. W. and H. Weyland. 1964. Lehrbuch der
Paläobotanik. 2. Aufl. bv H. Weyland. Berlin: Akademie-
Verlag.
(26a) Harland. W. B., A. G. Smith and B. Wilcock [eds.]
 1964. The Phanerozoic time-scale. London: Geological
 Society of London. [Issued as a supplement to vol. 120 of
 Quart. J. Geol. Soc. London.]
(27) Kirchheimer, F. 1957. Die Laubgewächse der
Braunkohlenzeit. Halle (Saale): Veb Wilhelm Knapp.
(27a) Krackowizer, F. 1953. Orquídeas fosséis. Revista do
 Circulo Paulista de Orquidofilos 10(3):36-38.
(28) Krackowizer, F. J. 1964. Orquídeas fosséis. Orquidea
(Rio de Janeiro) 26:39-40.
(29) MacGinitie. II. D. 1953. Fossil plants of the
Florissant beds, Colorado. Carnegie Inst. Washington Pub.
599:i-iii. 1-198.
(30) Machin (ńee Pallot). J. 1971. Plant microfossils
from Tertiary deposits of the Isle of Wight. New. Phytol.
70:851-872.
(31) Massalongo. A. B. 1857. Vorläufige Nachricht
über die neueren paläontologischen Entdeckungen am
Monte Bolca, Neues Jahrb. Mineral., Geognosie 1857:775-
778.
(32) Massalongo. A. B. 1858. Palaeophyta rariora
formationis tertiariae agri Veneti. Atti R. Ist. Veneto Sci.,
Ser. 3, 3:729-793.
(33) Massalongo. A. B. 1859a. Specimen photographicum
 animalium quorumdam plantarumque fossilium agri
 Veronensis. Veronae: Vincentini-Franchini. [Dual text in
 Italian and Latin.]
(34) Massalongo. A. B 1859b. Syllabus plantarum
fossilium hucusque in formationibus tertiariis agri Ve ne ti
detectarum. Veronae: A. Merlo.
(35) Melchior. H. 1964. Reihe Microspermae (Orchidales,
Gynandrae). Pp. 613-625 in H. Melchior [ed.]. A. Engler’s
Syllabus der Pflanzenfamilien. 12. Aufl. Bd. 2.
Angiospermen. Berlin-Nikolassee: Gebrüder Borntraeger.
(35a) Menard. H. W. 1 9 7 1 . Science: growth and change.
Cambridge, Mass.: Harvard University Press.
(36) Meschinelli, A., and X. Squinabol. 1893. Flora
tertiaria Ittalica. Patavii: Sumptibus Auctorum Typis
Seminarii.
(37) Naegeli O. 1905. Ueber westliche Florenelemente
in der Nordostschweiz. Ber. Schweiz. Bot. Ges. 15:14-25.
(38) Novak. F. A. 1961. Vyssi rostliny: Tracheophyta.
Praha: Nakladatelství Ceskoslovenské Akademie Véd.
(39) Pijl, L. van der. 1966. Pollination mechanisms in
orchids. Pp. 61-75 in J. G. Hawkcs [ed.]. Reproductive
biology and taxonomy of vascular plants. Oxford: Pergamon
Press.
(40) Pijl, L. van der, and C. H. Dodson. 1966. Orchid flowers:
their pollination and evolution, Coral Cables. Florida:
University of Miami Press.
(41) Potonié. R. 1967. Versuch der Einordnung der
fossilen Sporae dispersae in das phylogenetische System der
Pflanzenfamilien. Forschungsber. Landes Nordrhein-Westfalen Nr.
1761:1-310.
(42) Rolfe. R. A. 1909-12. The evolution of the Orchidaceae.
Orchid Rev. 17:129-132, 193-196. 249-252. 289-292. 353-356;
18:33-36, 97-99. 129-132, 162-166, 289-294. 321-325; 19:68-69.
289-292: 20:204-207, 225-228. 260-264. [General discussion
in 20:225-228, 260-264.]
(43) Schimper, W. P.. and A. Schenk. 1879-90.
Palaeophytologie. Abt. 2 in K. A. Zittel [ed.], Handbuch der
Palaeontologie. Münchcn: R. Oldenbourg. [Also the 1891
translation into French by C. Barrois et al.:
Paleophytologie. Pt. 2 in K. A. Zittel [ed.]. Traité de
paléontologie.]
(44) Schultcs. R. E. 1960. Native orchids of Trinidad and
Tobago. New York: Pergamon Press.
(45) Schultes, R. E. 1966. Orchid [in part]. Pp. 1041-1043
in Encyclopaedia Brittanica. Vol. 16. Chicago: Encyclopaedia
Brittanica. [Also in subsequent editions.]
(46) Schultes. R. E., and A. S. Pease. 1963. Generic
names of orchids: their origin and meaning. New York:
Academic Press.
(47) Sernander, R. 1894. Studier öfver den Gotländska
vegetationens utvecklingshistoria, Ph.D. Thesis, Universitet i
Uppsala. 1 1 2pp. [Privately printed.]
(48) Stebbins, G. L. Jr. 1950. Variation and evolution
in plants. New York: Columbia University Press.
(49) Straus, A. 1954. Beiträge zur Pliocänflora von
Willershausen. IV. Die Monocotyledonen. Palaeontographica
96B:1-11
(50) Straus, A. 1969. Beiträge zur Kenntnis der
    Pliozänflora von Willershausen (VII). Die Angiospermen-
    Früchte und -Samen. Argumenta Palaeobotannica 3:163
    197.
    (51) Thurmann, J. [1833.] Essai sur les soulèvemens
    Jurassiques du Porrentruy, avec une description
    géognostique des terrains secondaires de ce pays, et des
    considérations générales sur les chaines du Jura. Mém. Soc.
    Hist. Nat. Strasbourg 1 (livre. 2, article "L"):l-84.
    (52) Vent, W. 1965. Neue Pflanzenfunde aus den
    interglazialen Ilmtaltiavertinen von Weimar-Ehringsdorf.
    Geologie 14:198-205.
'

Weitere ähnliche Inhalte

Was ist angesagt?

Age related resisitance in plants
Age related resisitance in plantsAge related resisitance in plants
Age related resisitance in plantsManjappa Ganiger
 
role of endophytes.pptx
role of endophytes.pptxrole of endophytes.pptx
role of endophytes.pptxAnjuShukla11
 
Disease forecasting
Disease forecastingDisease forecasting
Disease forecastingDrTSuthinRaj
 
Biochemical plant defences(HR)
Biochemical plant defences(HR)Biochemical plant defences(HR)
Biochemical plant defences(HR)dev9105
 
Makalah+etika+illegal+logging
Makalah+etika+illegal+loggingMakalah+etika+illegal+logging
Makalah+etika+illegal+loggingAba Abdillah
 
Systemic Acquired Resistance (SAR) and it’s Significance in Plant Disease Ma...
Systemic Acquired Resistance (SAR) and it’s Significance in Plant  Disease Ma...Systemic Acquired Resistance (SAR) and it’s Significance in Plant  Disease Ma...
Systemic Acquired Resistance (SAR) and it’s Significance in Plant Disease Ma...Ankit Chaudhari
 
Management Strategies of Phytopathogenic Prokaryotes
Management  Strategies of Phytopathogenic Prokaryotes Management  Strategies of Phytopathogenic Prokaryotes
Management Strategies of Phytopathogenic Prokaryotes ZeeShanAli679
 
Tugas ppt tumbuhan tingkat tinggi
Tugas ppt tumbuhan tingkat tinggiTugas ppt tumbuhan tingkat tinggi
Tugas ppt tumbuhan tingkat tinggimarwahmoniCha
 
Josephson and Persistent Spin Currents in Bose-Einstein Condensates of Magnons
Josephson and Persistent Spin Currents in Bose-Einstein Condensates of MagnonsJosephson and Persistent Spin Currents in Bose-Einstein Condensates of Magnons
Josephson and Persistent Spin Currents in Bose-Einstein Condensates of MagnonsKouki Nakata
 
Effect of environment on plant diseases
Effect of environment on plant diseasesEffect of environment on plant diseases
Effect of environment on plant diseasesRajbir Singh
 
Hormone crosstalk in plant disease and defense
Hormone crosstalk in plant disease and defenseHormone crosstalk in plant disease and defense
Hormone crosstalk in plant disease and defenseAkankshaShukla85
 
Roshan Chandurkar Bacteriology History, Introduction & Importance
Roshan Chandurkar Bacteriology History, Introduction & ImportanceRoshan Chandurkar Bacteriology History, Introduction & Importance
Roshan Chandurkar Bacteriology History, Introduction & ImportanceRoshanChandurkar
 
Defensins: Antimicrobial peptide for the host plant resistance
Defensins: Antimicrobial peptide for the host plant resistanceDefensins: Antimicrobial peptide for the host plant resistance
Defensins: Antimicrobial peptide for the host plant resistancesnehaljikamade
 
Difraksi Sinar X (3)
Difraksi Sinar X (3)Difraksi Sinar X (3)
Difraksi Sinar X (3)jayamartha
 
BrownRotofPotato.pptx
BrownRotofPotato.pptxBrownRotofPotato.pptx
BrownRotofPotato.pptxDawitGetahun6
 
Plant parasite coevolution
Plant parasite coevolutionPlant parasite coevolution
Plant parasite coevolutionPavan R
 

Was ist angesagt? (20)

Age related resisitance in plants
Age related resisitance in plantsAge related resisitance in plants
Age related resisitance in plants
 
role of endophytes.pptx
role of endophytes.pptxrole of endophytes.pptx
role of endophytes.pptx
 
Stress
StressStress
Stress
 
Disease forecasting
Disease forecastingDisease forecasting
Disease forecasting
 
Biochemical plant defences(HR)
Biochemical plant defences(HR)Biochemical plant defences(HR)
Biochemical plant defences(HR)
 
Makalah+etika+illegal+logging
Makalah+etika+illegal+loggingMakalah+etika+illegal+logging
Makalah+etika+illegal+logging
 
Systemic Acquired Resistance (SAR) and it’s Significance in Plant Disease Ma...
Systemic Acquired Resistance (SAR) and it’s Significance in Plant  Disease Ma...Systemic Acquired Resistance (SAR) and it’s Significance in Plant  Disease Ma...
Systemic Acquired Resistance (SAR) and it’s Significance in Plant Disease Ma...
 
Management Strategies of Phytopathogenic Prokaryotes
Management  Strategies of Phytopathogenic Prokaryotes Management  Strategies of Phytopathogenic Prokaryotes
Management Strategies of Phytopathogenic Prokaryotes
 
Tugas ppt tumbuhan tingkat tinggi
Tugas ppt tumbuhan tingkat tinggiTugas ppt tumbuhan tingkat tinggi
Tugas ppt tumbuhan tingkat tinggi
 
Induksi Medan Magnet
Induksi Medan MagnetInduksi Medan Magnet
Induksi Medan Magnet
 
Josephson and Persistent Spin Currents in Bose-Einstein Condensates of Magnons
Josephson and Persistent Spin Currents in Bose-Einstein Condensates of MagnonsJosephson and Persistent Spin Currents in Bose-Einstein Condensates of Magnons
Josephson and Persistent Spin Currents in Bose-Einstein Condensates of Magnons
 
Effect of environment on plant diseases
Effect of environment on plant diseasesEffect of environment on plant diseases
Effect of environment on plant diseases
 
Pipa organa
Pipa organaPipa organa
Pipa organa
 
Hormone crosstalk in plant disease and defense
Hormone crosstalk in plant disease and defenseHormone crosstalk in plant disease and defense
Hormone crosstalk in plant disease and defense
 
Roshan Chandurkar Bacteriology History, Introduction & Importance
Roshan Chandurkar Bacteriology History, Introduction & ImportanceRoshan Chandurkar Bacteriology History, Introduction & Importance
Roshan Chandurkar Bacteriology History, Introduction & Importance
 
Defensins: Antimicrobial peptide for the host plant resistance
Defensins: Antimicrobial peptide for the host plant resistanceDefensins: Antimicrobial peptide for the host plant resistance
Defensins: Antimicrobial peptide for the host plant resistance
 
Difraksi Sinar X (3)
Difraksi Sinar X (3)Difraksi Sinar X (3)
Difraksi Sinar X (3)
 
BrownRotofPotato.pptx
BrownRotofPotato.pptxBrownRotofPotato.pptx
BrownRotofPotato.pptx
 
Penyakit Bulai Pada Jagung
Penyakit Bulai Pada JagungPenyakit Bulai Pada Jagung
Penyakit Bulai Pada Jagung
 
Plant parasite coevolution
Plant parasite coevolutionPlant parasite coevolution
Plant parasite coevolution
 

Andere mochten auch

Photographer research
Photographer researchPhotographer research
Photographer researchbekkiasquith
 
Film Festivals & Film Industry
Film Festivals & Film IndustryFilm Festivals & Film Industry
Film Festivals & Film IndustryWang Xiuyi
 
Rta Film Festivals
Rta Film FestivalsRta Film Festivals
Rta Film Festivalsmovieboy19
 
Student film festivals
Student film festivalsStudent film festivals
Student film festivalsslideenergy
 
Queens EDC Women's Power Networking Breakfast 11/5/14 - Juggling the Possibil...
Queens EDC Women's Power Networking Breakfast 11/5/14 - Juggling the Possibil...Queens EDC Women's Power Networking Breakfast 11/5/14 - Juggling the Possibil...
Queens EDC Women's Power Networking Breakfast 11/5/14 - Juggling the Possibil...Jen Slaw
 
Networking On The Greens
Networking On The GreensNetworking On The Greens
Networking On The Greenstferriera
 
Powerful Pitches And Presentations Basics
Powerful Pitches And Presentations BasicsPowerful Pitches And Presentations Basics
Powerful Pitches And Presentations Basicssarah_birken
 
Significant or notable film festivals independant
Significant or notable film festivals   independantSignificant or notable film festivals   independant
Significant or notable film festivals independantHG17
 
The television, social media and the Olympic Games
The television, social media and the Olympic GamesThe television, social media and the Olympic Games
The television, social media and the Olympic GamesEmilio Fernández Peña
 
Super Mixer at John Madden's Goal Line
Super Mixer at John Madden's Goal LineSuper Mixer at John Madden's Goal Line
Super Mixer at John Madden's Goal LineMichaelShay
 
Brochure "Think Big Concept "
Brochure "Think Big Concept " Brochure "Think Big Concept "
Brochure "Think Big Concept " NancyBond
 

Andere mochten auch (18)

Kings speech
Kings speechKings speech
Kings speech
 
Kings speech
Kings speechKings speech
Kings speech
 
Kings speech
Kings speechKings speech
Kings speech
 
Kings speech media
Kings speech mediaKings speech media
Kings speech media
 
Photographer research
Photographer researchPhotographer research
Photographer research
 
Film festivals
Film festivalsFilm festivals
Film festivals
 
Film Festivals & Film Industry
Film Festivals & Film IndustryFilm Festivals & Film Industry
Film Festivals & Film Industry
 
Rta Film Festivals
Rta Film FestivalsRta Film Festivals
Rta Film Festivals
 
Student film festivals
Student film festivalsStudent film festivals
Student film festivals
 
Queens EDC Women's Power Networking Breakfast 11/5/14 - Juggling the Possibil...
Queens EDC Women's Power Networking Breakfast 11/5/14 - Juggling the Possibil...Queens EDC Women's Power Networking Breakfast 11/5/14 - Juggling the Possibil...
Queens EDC Women's Power Networking Breakfast 11/5/14 - Juggling the Possibil...
 
Networking On The Greens
Networking On The GreensNetworking On The Greens
Networking On The Greens
 
FIRST team 1511 sponsorship presentation template
FIRST team 1511 sponsorship presentation templateFIRST team 1511 sponsorship presentation template
FIRST team 1511 sponsorship presentation template
 
Powerful Pitches And Presentations Basics
Powerful Pitches And Presentations BasicsPowerful Pitches And Presentations Basics
Powerful Pitches And Presentations Basics
 
Film festivals
Film festivalsFilm festivals
Film festivals
 
Significant or notable film festivals independant
Significant or notable film festivals   independantSignificant or notable film festivals   independant
Significant or notable film festivals independant
 
The television, social media and the Olympic Games
The television, social media and the Olympic GamesThe television, social media and the Olympic Games
The television, social media and the Olympic Games
 
Super Mixer at John Madden's Goal Line
Super Mixer at John Madden's Goal LineSuper Mixer at John Madden's Goal Line
Super Mixer at John Madden's Goal Line
 
Brochure "Think Big Concept "
Brochure "Think Big Concept " Brochure "Think Big Concept "
Brochure "Think Big Concept "
 

Ähnlich wie Fossils attributed to_the_orchidaceae

Evolutionary history of insects
Evolutionary history of  insectsEvolutionary history of  insects
Evolutionary history of insectsBhumika Kapoor
 
Origin and evolution of human dentition
Origin and evolution of human dentitionOrigin and evolution of human dentition
Origin and evolution of human dentitionadatnihc
 
K-T BOUNDARY PROBLEM
K-T BOUNDARY PROBLEMK-T BOUNDARY PROBLEM
K-T BOUNDARY PROBLEMparag sonwane
 
GEOLOGIC HISTORY, EVOLUTIONARY TRENDS OF BIRDS_110959.pptx
GEOLOGIC HISTORY, EVOLUTIONARY TRENDS OF BIRDS_110959.pptxGEOLOGIC HISTORY, EVOLUTIONARY TRENDS OF BIRDS_110959.pptx
GEOLOGIC HISTORY, EVOLUTIONARY TRENDS OF BIRDS_110959.pptxPRABHUPRASAD31
 
Geological time scale and plant life through ages
Geological time scale and plant life through agesGeological time scale and plant life through ages
Geological time scale and plant life through agesRimiRoy6
 
DIVERSITY OF LIFE- ABAO.pptxkyhfijdyujkhfujutd
DIVERSITY OF LIFE- ABAO.pptxkyhfijdyujkhfujutdDIVERSITY OF LIFE- ABAO.pptxkyhfijdyujkhfujutd
DIVERSITY OF LIFE- ABAO.pptxkyhfijdyujkhfujutdAngelicaRocamora1
 
Unit 6 history of life on earth
Unit 6 history of life on earthUnit 6 history of life on earth
Unit 6 history of life on earth9401140607087
 
Evidence For Evolution
Evidence For EvolutionEvidence For Evolution
Evidence For EvolutionMark McGinley
 
Paleobotanyofangiosperms sreeraj-141124044743-conversion-gate02
Paleobotanyofangiosperms sreeraj-141124044743-conversion-gate02Paleobotanyofangiosperms sreeraj-141124044743-conversion-gate02
Paleobotanyofangiosperms sreeraj-141124044743-conversion-gate02ubaid afzal
 
Dinosaur Extinction Research
Dinosaur Extinction ResearchDinosaur Extinction Research
Dinosaur Extinction ResearchKim Johnson
 
Ehret et-al.-2012 carcharodon-hubbelli
Ehret et-al.-2012 carcharodon-hubbelliEhret et-al.-2012 carcharodon-hubbelli
Ehret et-al.-2012 carcharodon-hubbelli1979aabb
 
The Evolution of Insects: Part THREE [3] / Summary & Anthropocene - Bug Scho...
The Evolution of Insects: Part THREE [3] /  Summary & Anthropocene - Bug Scho...The Evolution of Insects: Part THREE [3] /  Summary & Anthropocene - Bug Scho...
The Evolution of Insects: Part THREE [3] / Summary & Anthropocene - Bug Scho...Bart Coppens
 
Carrizo Wash Watershed Essay
Carrizo Wash Watershed EssayCarrizo Wash Watershed Essay
Carrizo Wash Watershed EssayJennifer Perry
 
The youngest record of phorusrhacid birds (aves, phorusrhacidae) from the lat...
The youngest record of phorusrhacid birds (aves, phorusrhacidae) from the lat...The youngest record of phorusrhacid birds (aves, phorusrhacidae) from the lat...
The youngest record of phorusrhacid birds (aves, phorusrhacidae) from the lat...herculanoalvarenga
 

Ähnlich wie Fossils attributed to_the_orchidaceae (20)

Evolutionary history of insects
Evolutionary history of  insectsEvolutionary history of  insects
Evolutionary history of insects
 
Origin and evolution of human dentition
Origin and evolution of human dentitionOrigin and evolution of human dentition
Origin and evolution of human dentition
 
K-T BOUNDARY PROBLEM
K-T BOUNDARY PROBLEMK-T BOUNDARY PROBLEM
K-T BOUNDARY PROBLEM
 
GEOLOGIC HISTORY, EVOLUTIONARY TRENDS OF BIRDS_110959.pptx
GEOLOGIC HISTORY, EVOLUTIONARY TRENDS OF BIRDS_110959.pptxGEOLOGIC HISTORY, EVOLUTIONARY TRENDS OF BIRDS_110959.pptx
GEOLOGIC HISTORY, EVOLUTIONARY TRENDS OF BIRDS_110959.pptx
 
Geological time scale and plant life through ages
Geological time scale and plant life through agesGeological time scale and plant life through ages
Geological time scale and plant life through ages
 
DIVERSITY OF LIFE- ABAO.pptxkyhfijdyujkhfujutd
DIVERSITY OF LIFE- ABAO.pptxkyhfijdyujkhfujutdDIVERSITY OF LIFE- ABAO.pptxkyhfijdyujkhfujutd
DIVERSITY OF LIFE- ABAO.pptxkyhfijdyujkhfujutd
 
Paleocene-Oligocene
Paleocene-OligocenePaleocene-Oligocene
Paleocene-Oligocene
 
Unit 6 history of life on earth
Unit 6 history of life on earthUnit 6 history of life on earth
Unit 6 history of life on earth
 
Evidence For Evolution
Evidence For EvolutionEvidence For Evolution
Evidence For Evolution
 
Geological time scale
Geological time scaleGeological time scale
Geological time scale
 
Paleobotanyofangiosperms sreeraj-141124044743-conversion-gate02
Paleobotanyofangiosperms sreeraj-141124044743-conversion-gate02Paleobotanyofangiosperms sreeraj-141124044743-conversion-gate02
Paleobotanyofangiosperms sreeraj-141124044743-conversion-gate02
 
Dinosaur Extinction Research
Dinosaur Extinction ResearchDinosaur Extinction Research
Dinosaur Extinction Research
 
Ehret et-al.-2012 carcharodon-hubbelli
Ehret et-al.-2012 carcharodon-hubbelliEhret et-al.-2012 carcharodon-hubbelli
Ehret et-al.-2012 carcharodon-hubbelli
 
The Evolution of Insects: Part THREE [3] / Summary & Anthropocene - Bug Scho...
The Evolution of Insects: Part THREE [3] /  Summary & Anthropocene - Bug Scho...The Evolution of Insects: Part THREE [3] /  Summary & Anthropocene - Bug Scho...
The Evolution of Insects: Part THREE [3] / Summary & Anthropocene - Bug Scho...
 
Geologic time table- GRADE 11
Geologic time table- GRADE 11Geologic time table- GRADE 11
Geologic time table- GRADE 11
 
fossil angioosperms.pptx
fossil angioosperms.pptxfossil angioosperms.pptx
fossil angioosperms.pptx
 
D3 human evolution
D3 human evolutionD3 human evolution
D3 human evolution
 
Carrizo Wash Watershed Essay
Carrizo Wash Watershed EssayCarrizo Wash Watershed Essay
Carrizo Wash Watershed Essay
 
The youngest record of phorusrhacid birds (aves, phorusrhacidae) from the lat...
The youngest record of phorusrhacid birds (aves, phorusrhacidae) from the lat...The youngest record of phorusrhacid birds (aves, phorusrhacidae) from the lat...
The youngest record of phorusrhacid birds (aves, phorusrhacidae) from the lat...
 
Chase 2013
Chase 2013Chase 2013
Chase 2013
 

Kürzlich hochgeladen

Artificial Intelligence & SEO Trends for 2024
Artificial Intelligence & SEO Trends for 2024Artificial Intelligence & SEO Trends for 2024
Artificial Intelligence & SEO Trends for 2024D Cloud Solutions
 
COMPUTER 10: Lesson 7 - File Storage and Online Collaboration
COMPUTER 10: Lesson 7 - File Storage and Online CollaborationCOMPUTER 10: Lesson 7 - File Storage and Online Collaboration
COMPUTER 10: Lesson 7 - File Storage and Online Collaborationbruanjhuli
 
Bird eye's view on Camunda open source ecosystem
Bird eye's view on Camunda open source ecosystemBird eye's view on Camunda open source ecosystem
Bird eye's view on Camunda open source ecosystemAsko Soukka
 
Using IESVE for Loads, Sizing and Heat Pump Modeling to Achieve Decarbonization
Using IESVE for Loads, Sizing and Heat Pump Modeling to Achieve DecarbonizationUsing IESVE for Loads, Sizing and Heat Pump Modeling to Achieve Decarbonization
Using IESVE for Loads, Sizing and Heat Pump Modeling to Achieve DecarbonizationIES VE
 
UiPath Studio Web workshop series - Day 7
UiPath Studio Web workshop series - Day 7UiPath Studio Web workshop series - Day 7
UiPath Studio Web workshop series - Day 7DianaGray10
 
IaC & GitOps in a Nutshell - a FridayInANuthshell Episode.pdf
IaC & GitOps in a Nutshell - a FridayInANuthshell Episode.pdfIaC & GitOps in a Nutshell - a FridayInANuthshell Episode.pdf
IaC & GitOps in a Nutshell - a FridayInANuthshell Episode.pdfDaniel Santiago Silva Capera
 
How Accurate are Carbon Emissions Projections?
How Accurate are Carbon Emissions Projections?How Accurate are Carbon Emissions Projections?
How Accurate are Carbon Emissions Projections?IES VE
 
UiPath Platform: The Backend Engine Powering Your Automation - Session 1
UiPath Platform: The Backend Engine Powering Your Automation - Session 1UiPath Platform: The Backend Engine Powering Your Automation - Session 1
UiPath Platform: The Backend Engine Powering Your Automation - Session 1DianaGray10
 
COMPUTER 10 Lesson 8 - Building a Website
COMPUTER 10 Lesson 8 - Building a WebsiteCOMPUTER 10 Lesson 8 - Building a Website
COMPUTER 10 Lesson 8 - Building a Websitedgelyza
 
Igniting Next Level Productivity with AI-Infused Data Integration Workflows
Igniting Next Level Productivity with AI-Infused Data Integration WorkflowsIgniting Next Level Productivity with AI-Infused Data Integration Workflows
Igniting Next Level Productivity with AI-Infused Data Integration WorkflowsSafe Software
 
Basic Building Blocks of Internet of Things.
Basic Building Blocks of Internet of Things.Basic Building Blocks of Internet of Things.
Basic Building Blocks of Internet of Things.YounusS2
 
AI Fame Rush Review – Virtual Influencer Creation In Just Minutes
AI Fame Rush Review – Virtual Influencer Creation In Just MinutesAI Fame Rush Review – Virtual Influencer Creation In Just Minutes
AI Fame Rush Review – Virtual Influencer Creation In Just MinutesMd Hossain Ali
 
Apres-Cyber - The Data Dilemma: Bridging Offensive Operations and Machine Lea...
Apres-Cyber - The Data Dilemma: Bridging Offensive Operations and Machine Lea...Apres-Cyber - The Data Dilemma: Bridging Offensive Operations and Machine Lea...
Apres-Cyber - The Data Dilemma: Bridging Offensive Operations and Machine Lea...Will Schroeder
 
Designing A Time bound resource download URL
Designing A Time bound resource download URLDesigning A Time bound resource download URL
Designing A Time bound resource download URLRuncy Oommen
 
IESVE Software for Florida Code Compliance Using ASHRAE 90.1-2019
IESVE Software for Florida Code Compliance Using ASHRAE 90.1-2019IESVE Software for Florida Code Compliance Using ASHRAE 90.1-2019
IESVE Software for Florida Code Compliance Using ASHRAE 90.1-2019IES VE
 
Linked Data in Production: Moving Beyond Ontologies
Linked Data in Production: Moving Beyond OntologiesLinked Data in Production: Moving Beyond Ontologies
Linked Data in Production: Moving Beyond OntologiesDavid Newbury
 
Building Your Own AI Instance (TBLC AI )
Building Your Own AI Instance (TBLC AI )Building Your Own AI Instance (TBLC AI )
Building Your Own AI Instance (TBLC AI )Brian Pichman
 

Kürzlich hochgeladen (20)

Artificial Intelligence & SEO Trends for 2024
Artificial Intelligence & SEO Trends for 2024Artificial Intelligence & SEO Trends for 2024
Artificial Intelligence & SEO Trends for 2024
 
COMPUTER 10: Lesson 7 - File Storage and Online Collaboration
COMPUTER 10: Lesson 7 - File Storage and Online CollaborationCOMPUTER 10: Lesson 7 - File Storage and Online Collaboration
COMPUTER 10: Lesson 7 - File Storage and Online Collaboration
 
201610817 - edge part1
201610817 - edge part1201610817 - edge part1
201610817 - edge part1
 
Bird eye's view on Camunda open source ecosystem
Bird eye's view on Camunda open source ecosystemBird eye's view on Camunda open source ecosystem
Bird eye's view on Camunda open source ecosystem
 
20150722 - AGV
20150722 - AGV20150722 - AGV
20150722 - AGV
 
Using IESVE for Loads, Sizing and Heat Pump Modeling to Achieve Decarbonization
Using IESVE for Loads, Sizing and Heat Pump Modeling to Achieve DecarbonizationUsing IESVE for Loads, Sizing and Heat Pump Modeling to Achieve Decarbonization
Using IESVE for Loads, Sizing and Heat Pump Modeling to Achieve Decarbonization
 
UiPath Studio Web workshop series - Day 7
UiPath Studio Web workshop series - Day 7UiPath Studio Web workshop series - Day 7
UiPath Studio Web workshop series - Day 7
 
IaC & GitOps in a Nutshell - a FridayInANuthshell Episode.pdf
IaC & GitOps in a Nutshell - a FridayInANuthshell Episode.pdfIaC & GitOps in a Nutshell - a FridayInANuthshell Episode.pdf
IaC & GitOps in a Nutshell - a FridayInANuthshell Episode.pdf
 
How Accurate are Carbon Emissions Projections?
How Accurate are Carbon Emissions Projections?How Accurate are Carbon Emissions Projections?
How Accurate are Carbon Emissions Projections?
 
UiPath Platform: The Backend Engine Powering Your Automation - Session 1
UiPath Platform: The Backend Engine Powering Your Automation - Session 1UiPath Platform: The Backend Engine Powering Your Automation - Session 1
UiPath Platform: The Backend Engine Powering Your Automation - Session 1
 
COMPUTER 10 Lesson 8 - Building a Website
COMPUTER 10 Lesson 8 - Building a WebsiteCOMPUTER 10 Lesson 8 - Building a Website
COMPUTER 10 Lesson 8 - Building a Website
 
Igniting Next Level Productivity with AI-Infused Data Integration Workflows
Igniting Next Level Productivity with AI-Infused Data Integration WorkflowsIgniting Next Level Productivity with AI-Infused Data Integration Workflows
Igniting Next Level Productivity with AI-Infused Data Integration Workflows
 
20230104 - machine vision
20230104 - machine vision20230104 - machine vision
20230104 - machine vision
 
Basic Building Blocks of Internet of Things.
Basic Building Blocks of Internet of Things.Basic Building Blocks of Internet of Things.
Basic Building Blocks of Internet of Things.
 
AI Fame Rush Review – Virtual Influencer Creation In Just Minutes
AI Fame Rush Review – Virtual Influencer Creation In Just MinutesAI Fame Rush Review – Virtual Influencer Creation In Just Minutes
AI Fame Rush Review – Virtual Influencer Creation In Just Minutes
 
Apres-Cyber - The Data Dilemma: Bridging Offensive Operations and Machine Lea...
Apres-Cyber - The Data Dilemma: Bridging Offensive Operations and Machine Lea...Apres-Cyber - The Data Dilemma: Bridging Offensive Operations and Machine Lea...
Apres-Cyber - The Data Dilemma: Bridging Offensive Operations and Machine Lea...
 
Designing A Time bound resource download URL
Designing A Time bound resource download URLDesigning A Time bound resource download URL
Designing A Time bound resource download URL
 
IESVE Software for Florida Code Compliance Using ASHRAE 90.1-2019
IESVE Software for Florida Code Compliance Using ASHRAE 90.1-2019IESVE Software for Florida Code Compliance Using ASHRAE 90.1-2019
IESVE Software for Florida Code Compliance Using ASHRAE 90.1-2019
 
Linked Data in Production: Moving Beyond Ontologies
Linked Data in Production: Moving Beyond OntologiesLinked Data in Production: Moving Beyond Ontologies
Linked Data in Production: Moving Beyond Ontologies
 
Building Your Own AI Instance (TBLC AI )
Building Your Own AI Instance (TBLC AI )Building Your Own AI Instance (TBLC AI )
Building Your Own AI Instance (TBLC AI )
 

Fossils attributed to_the_orchidaceae

  • 1. Fossils Attributed to the Orchidaceae RUDOLF SCHMID AND MARVIN J. SCHMID1 INVESTIGATORS INTERESTED IN the evolutionary history of a family desire to have information from the fossil record to provide (hopefully) unequivocal evidence for the early history of that family. Unfortunately for orchidologists, the known fossil record of the Orchidaceae is extremely meager. We decided to summarize what is known of the fossil record of orchids (a) because there are no accounts available other than the rather superficial ones by Darrah (15) and Krackowizer (27a, 28) , (b) because there are number of misconceptions in the literature that should be corrected, and, most importantly, (c) because a reasonably complete list of fossils attributed to the Orchidaceae could be prepared since we had access to the Compendium Index of Paleobotany (see 3, 19 for accounts of its coverage) and to libraries with extensive holdings in rather obscure publications. Most persons seem to favor a relatively great age for the Orchidaceae (e.g., 1, 5, 6, 12, M, 15, IS, 21-23, 48), but relatively few have ventured a specific time and/or place of origin. Garay (21- 23), following Stebbins (48, pp. 501-502), postulates an origin in the early Cretaceous- and in addition proposes Malaysia as the most likely cradle of orchidhood. Leon Croizat (pers. comm., 1972) believes that the orchids arose "surely not later than the earliest Cretaceous. Brieger (5, 6) favors the early Tertiary and the "united Asiatic-American" tropics (6, p. 329, specific area not indicated). In contrast, some authors, notably Schultes, believe that the orchids may be "a comparatively young group" (44, p. 1; 45, p. 1043). While there is some dispute as to the exact time of origin of the family, nearly all workers (e.g., 5, 16, 17, 21. 40, 44) seem to agree that currently, and in the immediate past, the Orchidaceae are in a very active period of evolution. Most workers have indicated either that there is no fossil record for the Orchidaceae (e.g., 1, 6, 10, 12, 13, 20, 24) or that there are only doubtful orchidaceous fossils (e.g., 15, 16, 18, 21, 25, 32-34, 36, 39, 40, 42, 44, 45). In contrast, a few persons (e.g., I I , 26, 28, 35, 38, 43, 49, 50, 52) have stated that valid orchid fossils exist. Most of these authors apparently based their view concerning the fossil record of the Orchidaceae chiefly or only on knowledge of the very dubiously orchidaceous Protorchis monorchis and Palaeorchis rhizoma described by Massalongo (31-34) from the Eocene of Italy. 4 Nevertheless, a number of other fossils have also been attributed to the Orchidaceae, most significantly Straus' (49, 50) three species of putative orchid fruits (Orchidacites) from the Pliocene of Germany. With the exception of Gothan and Weyland (26), Kirchheimer (27), and Melchior (35), however, most recent workers seem unaware of Straus' finds. These and other taxa will be discussed in detail below.
  • 2. Table 1. Geologic time scale (pre-Mesozoic omitted). (After Hartland, et.al. 26a) Era Period Epoch Beginning of interval (in millions of years Holocene (Recent) 0.005 Quaternary Pleistocene (Glacial) 2.5 Pleiocene 7 Miocene 26 Oligocene 38 Eocene 54 Cenozoic Tertiary Paleocene 65 Cretaceous 136 Jurassic 190 Mesozoic Triassic 225 Orchids are not favorable candidates for fossilization, an obvious conclusion that has not escaped previous authors (e.g., 15, 18, 28, 40. 44). The following characteristics of most Orchidaceae probably account for their scarcity as fossils: (a) predominant occurrence, both in the present and presumably in the distant past, in the wet tropics, which are areas of rapid decay; (b) herbaceous habit; (c) epiphytic habit, which would generally preclude orchids from the conditions (usually aquatic) most conducive to fossilization (see also 15) ; (d) production of pollinia (usually) rather than individual pollen grains, and dispersion of the former by animal vectors instead of wind: and (e) minute, easily degradable seeds. Krackowizer (28) and Schimper and Schenk (43), however, apparently believed a rather extensive fossil record of the Orchidaceae is to be expected, and Darrah (15) and also Krackowizer (28) suggested that fossil orchids might eventually be encountered in deposits in tropical areas when these become better known. Fossils that have been attributed to the Orchidaceae (or to the Protorchidaceae) are strictly megafossils (e.g., fruits, leaves, tubers, etc.); orchidaceous microfossils (e.g., seeds and pollen) have not been reported in the literature. Discoveries of cuticular remains (as those already found of the Pliocene Orchidacites wegelei of Straus, HI) perhaps offer the best hope for significant additions to the fossil record of (he Orchidaceae. Although there is no record of fossil orchid pollen, even if orchid pollen were preserved as fossils, it is a moot point that it would be recognizable as such. Botanists simply might not recognize the fossilized pollen of those orchidaceous forms that had not yet evolved pollinia. Perhaps significantly, the Asclepiadaceae, which like the Orchidaceae possess pollinia, are not listed in Potonie's (41) recent compendium of fossil pollen and spores. Although Chandler's (9, and works cited therein) extensive investigations (initially with the late Eleanor M. Reid) of the Tertiary London Clay Flora of England over a period of several decades failed to reveal any orchidaceous remains,
  • 3. she suggested (8, p. 29) that "possibly search for pollen among the finer sediments and residues may eventually demonstrate the presence of this family" in the London Clay Flora. Subsequent palynological work (two 1961 Ph.D. theses by Ma Khin Sein and Jane Pallot at the University of London, both cited in 9, the latter published as 30) , however, has thus far failed to substantiate this prediction. As recently discussed by Eyde (19), there are several paths into the paleobotanical literature. We checked a variety of sources for records of orchidaceous fossils, most importantly the index by Andrews (3, including unpublished cards for additions since 19(>5) and the United States Geological Survey's Compendium Index of Paleontology, an unpublished file available for consultation only in the Natural History Building, Room W-300, of the Smithsonian Institution, Washington, D. C. (see 3, 19) . We checked the Compendium Index for most of the temperate genera listed in Schultes and Pease (46). Most tropical genera, however, were not sought in the Compendium Index due to the minuscule yield that could only result from such a mountain of effort. As noted above, tropical plants are unlikely candidates for fossilization. In addition, most tropical orchids are endemics that presumably evolved during the Quaternary, and hence any fossils of them would be unlikely to be encountered by paleobotanists, most of whom have worked (until very recently, at least) in temperate areas. Finally, any orchidaceous fossils of the pre-Quaternary tropical floras of presently temperate areas would probably be given generic names not based on living taxa. 5 REPORTS OF ORCHIDACEOUS FOSSILS Three extinct genera (Palaeorchis, Protorchis and Orchidacites) have been designated as orchidaceous or protorchidaceous. At least one other fossil taxon, Antholithes pediloides, has been regarded as an orchid. In addition, fossil remains from the Quaternary have been attributed to a number of extant, north temperate orchid taxa. Purists who restrict "fossils" to pie-Quaternary remains and therefore regard Pleistocene finds as "subfossils" may object to the inclusion of plant remains from the Quaternary in the following enumera- tion: Jurassic The Compendium Index of Paleobotany, citing Thurmann (51) , lists the following extant species as occurring in the Jurassic strata at Porrentruy, France: Ophrys myodes Jacq., Orchis morio L., O. pyramidalls I,., Satyrium viride L., and Serapias rubra L. This is incorrect. Thurmann did indeed list these species, but only as part of the modern vegetation of this region. Eocene Protorchis monorchis Massalongo (31-33) and Palaeorchis rhizoma (.Massalongo) Massalongo (32) (= Protorchis rhizoma Massalongo, 31): These species represent the first described and also the geologically oldest fossils that might possibly represent orchids. In 1857 Massalongo (31) listed, without benefit of description or illustration (hence nomina nuda) , the new generic name Protorchis, with two new species P. monorchis and P. rhizoma based on specimens from the calcareous Eocene deposits at Monte Bolca, Italy. Massalongo had only four specimens at his disposal — three of the former species, one of the latter (33, 34). In 1858 Massalongo (32) validly published Protorchis monorchis and also the new combination Palaeorchis rhyzoma (the specific epithet is an orthographic error) based on Protorchis rhizoma. The next year a more complete description and also a photograph (see Fig. 1) of Protorchis monorchis were published (33). In his 1858 work Massalongo dated Protorchis as "1851" with the added notation "in lit.” et in Musaeo" (32, p. 749). Since the 1854 reference is obviously unpublished, the nomina nuda in the 1857 report (31) thus represent the actual first (though taxonomically invalid) publication of
  • 4. the names involved. Although Massalongo (31) initially listed his new species under the Orchidaceae, he subsequently (32-34) very carefully indicated the tentatively orchidaceous nature of his fossil specimens by including them in a new taxon, the Protorchidaceae ("Protorchidee" in 32, 33; or the Latinized "Protorchideae" in 33, 34). Massalongo (32, 33) admitted that he was unable to find in the extant flora counterparts of his fossils. Massalongo (32, 33) noted a resemblance of both his fossil species to the Araceae, which he apparently regarded as being rather close to the Orchidaceae (32). A superficial likeness between Palaeorchis rhizoma and the fossil alga Delesserites was rejected when he decided on the affinities of the former (32). After additional concern that Palaeorchis rhizoma might be butomaceous, Massalongo (32) finally decided to retain this species in his Protorchidaceae. This discussion illustrates the difficulty Massalongo had in assigning his fossils to an extant plant group. This fact is apparently realized by very few authors since a number (e.g., 15, 18, 28, 42) incorrectly state or imply that Massalongo had regarded his fossils as orchids.
  • 5. According to Massalongo (32, descriptive terminology below is his) , the Protorchidaceae are next to the orchids and the aroids and consist of small herbs with tubers or rhizomes bearing lateral fibrous roots and several very slender, cuneate-obovate or spathulate leaves with entire margins and fine midribs. Misstatements to the contrary (15, 25, 26, 28, 43), both species do not possess tubers. Protorchis monorchis (FIGURE 1) has a round, solitary tuber whereas Palaeorchis rhizoma (never illustrated by Massalongo) differs chiefly in having a perpendicularly cylindrical rhizome covered with circular, papillate leaf scars (32). The two species also differ somewhat in having spathulate versus oblong to spathulate leaves, respectively (32). Massalongo (33) subsequently indicated that Protorchis monorchis, of which three speci- mens were available (33), strictly speaking does not have a true tuber, but rather a rounded rhizome (FIGURE 1). In the same publication he (33) also added the
  • 6. following information for Protorchis monorchis: tuber 7-8 mm in diameter; leaves 3-4 per plant, attenuate into a petiole, 5 cm long, and 12-15-18 mm wide.11 Although Massalongo (34, p. 133) finally indicated that specimens of both this species and Palaeorchis rhizoma are seedlings, there is no mention in his previous descriptions (32, 33) of the probable developmental age of these fossils. Most workers, usually referring only to Protorchis monorchis, have subsequently concluded that Massalongo's fossils are not truly representative of the Orchidaceae (e.g., 15, 18, 21, 25, 26, 35, 39, 40, 42). Schimper and Schenk (43) , however, accepted Massalongo's finds as orchidaceous, and van der Pijl (39, 40) apparently seems tempted to accept Protorchis monorchis as validly orchidaceous, no doubt because its Eocene date ties in with his understanding of the evolution of the bees. Meschinelli and Squinabol (3i) included both of Massalongo's fossil species under the Protorchidaceae in the order Micro- spermae (= Orchidales), but these authors noted that Palaeorchis rhizoma is probably a member of the Butomaceae. Krackowizer (28) 7 accepted the views of Meschinelli and Squinabol (36) except that he regarded Protorchis monorchis as a true orchid rather than as a protorchid. Admitting that both of Massalongo's fossils are doubtfully orchidaceous, Leslie A. Garay (pers. comm., 1972) nevertheless maintains that of all the fossils attributed to the Orchidaceae, Protorchis monorchis is perhaps the most likely candidate for inclusion in the family, largely because of its similarity to Orchis pallens L. In conclusion, the orchidaceous nature of Massalongo's fossils is clearly very questionable. As has already been suggested (Chester A. Arnold, pers. comm., 1967; 15, 18), perhaps the most charitable tiling that can be said about the affinity of Protorchis and Palaeorchis is that they are monocotyledonous. Oligocene Antholithes pediloides Cockerell (1 1): T. D. A. Cockerell, the prolific describer of fossils from the western United States,8 in 1915 delineated from the Lower Oligocene (the age according to MacGinitie, 29; incorrectly regarded as Miocene by Cockerell, I I ) beds at Florissant, Colorado, a new species in the fossil artificial (or form) genus Antholithes. Cockerell attributed the fossil (FIGURE 9), A. pediloides, to the Orchidaceae because of its marked resemblance to the lip of Cypripedium, and he also presumed that the several small "subhyaline" spots scattered over the surface might represent the work of some insect. Other than the suggestive outline of the fossil, however, the lack of significant detail makes Cockerell's determination extremely doubtful. In his classic flora of the Florissant beds, MacGinitie (29) reached the same conclusion and disposed of A. pediloides among "species of somewhat doubtful taxonomic value" (p. 159) under incertae sedis . Miocene Darrah (15) briefly discussed, and then discounted as truly orchidaceous, a fossil stem (apparently unnamed) from the Miocene of Hungary that had been described by a Robert Brown (there were several Robert Browns). Since Darrah provided no references in his note, and since alter considerable searching we have been unable to locate any additional information concerning this fossil, we can only quote Darrah (15, p. 149) fully: "A third form [besides Massalongo's Protorchis and Palaeorchis] was once provisionally accepted as a fossil orchid. This fossil stem, found in rocks of Miocene age in Hungary, included a few structurally preserved hair-like roots which Robert Brown considered to be of some epiphytic: orchidaceous plant. . . . As a matter of fact it was with this organ [the pseudobulb] that Robert Brown attempted to compare his supposed fossil from Hungary."
  • 7. Pliocene Orchidacites orchidioides Straus (49), O. wegelei Straus (49), and O. cypripedioides Straus (50): In 1954 Straus (49) described from the Upper Pliocene of Willershausen, Germany, two species of fruits, Orchidacites orchidioides and O. wegelei which he assigned to the Orchidaceae. The two species, especially the former (as suggested by i t s name), were thought to resemble various species of Orchis (49). More recently, Straus (50) provided for Orchidacites a generic diagnosis, which had been omitted from the 1954 report, and described a third species, O. cypripedioides, with fruits regarded as similar to those of Cypripedium. These taxa are illustrated in FIGURES 2 to 8, reproduced from Straus' more recent paper (50). Orchidacites is a form genus proposed for fossil fruits comparable to the capsules of various extant orchid genera (50). According to Straus (50), the fossil capsules, 1.5 to 2.5 cm long, are ellipsoidal or narrowly ellipsoidal and have several longitudinal striae (FIGURES 2-8); the remnants of a corolla often occur at the fruit apex (FIGURES 2-4). Straus (50, also pers. comm., 1972) believes that the fossil fruits of Orchidacites came from epiphytes growing on rotting branches that eventually were blown into the sediments by wind, and, as a consequence, he has speculated (50, pers. comm., 1972) that many of the present-day orchids (e.g. Limodorum, Neottia, Corallorrhiza, and Cypripedium) were primitively epiphytic and now are terrestrial "secondary relicts." This view, of course, is dissonant with the conventional one that the terrestrial habit is ancestral and the epiphytic derived (e.g., 5, 6, 16, 17, 21-23, 39, 42). The Straus fossils have received little comment from either orchidologists or paleobotanists. Melchior (35) and Gothan and Weyland (26) accepted the fossils as unmistakably orchidaceous. Kirchheimer (27, p. 650), however, remained skeptical, believing that the inferior, wing like, ribbed gynoecium with a distinct styloid process evokes resemblances to young fruits of Halesia, Pterostyrax and other Styracaceae (a completely unrelated family in the dicotyledons). Straus (pers. comm., 1972), in counterargument, however, believes that the fruits he described are truly orchidaceous since fruits of the Styracaceae never show floral remains whereas fruits of the Orchidaceae often do. On examining the photographs reproduced herein ( FIGURES 2-8), Garay (pers. comm., 1972) is also disinclined to accept Straus' fossils as orchidaceous because of the curious 3-pronged floral remnants (interpreted by Garay as a column) ( FIGURES 2-4) and because of the apparently excessive number of ribs ( FIGURES 2-8) for true orchid fruits (which have a maximum of six). Robert L. Dressier (pers. comm., 1972) is of a similar opinion, although he is less certain in excluding Orchidacites cypripedioides (FIGURES 5-8) from the orchids since the fossil "looks rather like a Cattleya fruit." In defense of Straus, we should note that a 3-pronged calyx of fused sepals occurs in some modern taxa (e.g. Pterostylis. see FIGURE 84 in 40: Masdevallia, etc.) and that on orchid fruits a greater number of ribs (than six) may be apparent since these may be variously secondarily divided (e.g., Trichopilia suavis Lindl. et Paxton). Unfortunately, Straus (50) hurts his own cause by interpreting the floral remnants (FIGURES 2-4) as a corolla, but it is perhaps more likely that they represent a calyx. Quaternary Quaternary orchid fossils are included here for completeness, although they are unimportant from the viewpoint of our understanding of the origin and most of the subsequent evolution of the family. The names of at least 20 extant species of orchids are listed in the Compendium Index of Paleobotany and are
  • 8. attributed to both the Pleistocene and Holocene (= Recent or Postglacial) of the Quaternary. Most of these listings were compiled around the turn of the century, when the Compendium Index included casual, incidental references to fossils - a practice long discontinued ( 1 9 ). Unfortunately, a number of these listings are not applicable because the original works discuss the various orchid species as components of the contemporary flora and not as fossils. This is the case with reports of Goodyera repens (L.) R. Br. from the Quaternary of Denmark, Himantoglossum hircinum (L.) Sprengel and Ophrys aranifera Huds. from the Postglacial of Switzerland, and Malaxis paludosa (L.) Sw. from the Holocene of Germany, which the Compendium Index attributes to Anderson (2), Naegeli (37, as cited in 7), and Becker (4), respectively. The Compendium Index also attributes the following extant orchid taxa to Sernander's (47) extensive work on the Quaternary (Wiirm Glacial and Postglacial) of Gotland, Sweden (names listed as they appear in Sernander): Anacamptis pyramidalis (L.) Rich., Cephalanthera ensifolia (Sw.) Rich., Corallorrhiza innata R. Br., Epipactis palustris (L.) Crantz, Gymnadenia conopsea (L.) R. Br., G. odoratissima (L.) Rich., Listera cordata (L.) R. Br., L. ovata (L.) R. Br., Malaxis monophyllos (L.) Sw., Neottia nidusavis (L.) Rich., Orchis angustifolia Wimm. et Grab., O. maculata L., O. militaris L., O. ustulata L., Platanthera bifolia (L.) Rich., and Sturmia loeselii (L.) Reichb. However, Sernander (47) merely discusses these and other orchid species in terms of a phytosociological survey of the modern bog vegetation of Gotland. The bogs Sernander studied did indeed contain identifiable fossils, but none of these were orchids. To our knowledge, there is only one report of a Quaternary fossil attributed to the Orchidaceae. In 1965 Vent (52, p. 200) described leaf and fruit impressions from the Riss-Wurm Interglacial of Weimar-Ehringsdorf, Germa- ny, and assigned these to "cf. Epipactis palustris (Mill.) Crantz" in the Orchidaceae. Garay (pers. comm., 1972) , however, discounted the orchidaceous nature of these fossils after examining Vent's photographs (FIGURE 10) . SUMMARY Fossils dating from the Eocene to the Quaternary have been attributed to the Orchidaceae, but objections have been raised against the orchidaceous nature of all of these fossils. The most likely orchid fossils, nevertheless, remain Massalongo's famous fossils from the Eocene of Italy — Protorchis monorchis and Palaeorchis rhizoma — and especially Straus' recent finds from the Pliocene of Germany Orchidacites orchidioides, O. wegelei and O. cypripedioides. In conclusion, then, the Orchidaceae have no positive fossil record and in this sense present a striking parallel to two well-known gods of mythology: Athena, who sprang fully grown and fully armored from the head of Zeus; and the Aztec Huitzilopochtli, who was borne fully grown and fully armored from Coatlicue. Acknowledgements: This study was carried out while the senior author was the recipient of a Smithsonian Institution postdoctoral fellowship. We thank Norris H. Williams and Leslie A. Garay for valuable discussions. REFERENCES (1) Ames, O. and D. S. Correll. 1952-53. Orchids of Guatemala, Heldiuna: Rot. 26:-i-xiii. 1-727. (2) Andersson, G. 1906. Die Entwicklungsgeschichte der skandinavischen Flora. Pp. 45-97 in Resultats Set. Congr. Int. Dot., Vienne, 1905. (3) Andrews, H. N., Jr. 1970. Index o£ generic names of fossil plants, 1820-1965. U.S. Geol. SURV. Bull.. 1300. (4) Becker. G. 1874. Botanische Wanderungen durch die
  • 9. Sümpfe und Torfmoore der Niederrheinischen Ebene. Verh. Nalurhist. Vereines Preus* Rheinl. Westphallens 31:137-158. (5) Brieger, F. G. 1958. On the phytogeography of orchids. Pp. 189-200 in Proc. 2nd World Orchid Conf., Honolulu, 1957. (6) Brieger, F. (1960. Geographic distribution and phyllogeny [sic] of orchids. Pp. 328-333 in Proc. 3rd World Orchid Conf.. London. I960. (7) Brockmann-Jerosch, H. 1910. Die Änderungen des Klimas seit der grösstcn Ausdehnung der letzten Eiszeit in der Schweiz. Pp. 55-71 in Die Veränderungen des Klimas seit dem Maximum der letzten Eiszeit, Ber., Exekutivkomitee I I . Int. Geol.-Kongr., Stockholm. 1910. (8) Chandler. M. E. J. 1951. The Lower Tcrtiary floras of southern England. I. Palaeocene floras: London Clay Flora (supplement). London: British Museum (Natural History). [Text and plates separately bound.] (9) Chandler, M. E. J. 1964. Idem. IV. A summary and survey of findings in the light of recent botanical observations. London: British Museum (Natural History). (10) Chesters, K. I. M., F. R. Gnauck and X. F. Hughes. 1967. Angiospermae. Pp. 269-288 in V. B. Harland et al. [eds.], The fossil record. London: Geological Society of London. (11) Cockerell. T. D. A. 1915. Notes on orchids. Bot. Gaz. 59:331-333. (12) Correll, D. S. 1950. Native orchids of North America. Waltham, Mass.: Chronica Botanica Co. (13) Coulter, J. M. and C. J. Chamberlain. 1903. Morphology of angiosperms. New York: II. Appleton and Co. (14) Croizat, L. 1961. Principia botanica. Caracas: The Author. [1 vol. in 2.] [Published 1961.] (15) Darrah. W. C: 1940. Supposed fossil orchids. Amer. Orchid Soc. Bull. 9:149-150. (16) Dodson. C. H. and R. J. Gillespie. 1967. The biology of the orchids. The Mid-America Orchid Congress. [No city of publication given.] (17) Dressler. R. L. and C. H. Dodson. 1960. Classification and phytogeny in the Orchidaceae. Anu. Missouri Bot. Gard. 47:25-68. (18) Dunsterville, G. C. K., and L. A. Garay. 1959. Venezuelan Orchids Illustrated. Vol.1. London: Andre Deutsch. [Also Introduction in Spanish in Vol. 2. 1961.] (19) Eyde. R. H. 1972. Note on geologic histories of flowering plants, Brittonia 24:111-116. (20) Andreanszky, G. 1954. Osnövénytan. Budapest. Akadémiai Kiado.
  • 10. (21) Garay, L. A. 1960. On the origin of the Orchidaceae. Bot. Mus. Leaflets Harvard Univ. 19:57-96 [Also in Proc. 3rd World Orchid Conf., London. 1960, pp. 172-196.] (22) Garay, L. A. [1964.] Evolutionary significance of geographical distribution of orchids. Pp. 170-187 in Proc. 4th World Orchid Conf,.Singapore, 1963. (23) Garay, L. A. 1972 On the origin of the Orchidaceae, II. J. Arnold Arb. 53:202-215. (24) Godfrey. M. J. 1933. Monograph & Iconograph of native British Orchidaceae. Cambridge: University Press. (25) Gothan, W. 1921. H. Potonié’s Lehrbuch der Paläobotanik. 2. Aufl. Berlin: Gebröder Borntraeger. (26) Gothan. W. and H. Weyland. 1964. Lehrbuch der Paläobotanik. 2. Aufl. bv H. Weyland. Berlin: Akademie- Verlag. (26a) Harland. W. B., A. G. Smith and B. Wilcock [eds.] 1964. The Phanerozoic time-scale. London: Geological Society of London. [Issued as a supplement to vol. 120 of Quart. J. Geol. Soc. London.] (27) Kirchheimer, F. 1957. Die Laubgewächse der Braunkohlenzeit. Halle (Saale): Veb Wilhelm Knapp. (27a) Krackowizer, F. 1953. Orquídeas fosséis. Revista do Circulo Paulista de Orquidofilos 10(3):36-38. (28) Krackowizer, F. J. 1964. Orquídeas fosséis. Orquidea (Rio de Janeiro) 26:39-40. (29) MacGinitie. II. D. 1953. Fossil plants of the Florissant beds, Colorado. Carnegie Inst. Washington Pub. 599:i-iii. 1-198. (30) Machin (ńee Pallot). J. 1971. Plant microfossils from Tertiary deposits of the Isle of Wight. New. Phytol. 70:851-872. (31) Massalongo. A. B. 1857. Vorläufige Nachricht über die neueren paläontologischen Entdeckungen am Monte Bolca, Neues Jahrb. Mineral., Geognosie 1857:775- 778. (32) Massalongo. A. B. 1858. Palaeophyta rariora formationis tertiariae agri Veneti. Atti R. Ist. Veneto Sci., Ser. 3, 3:729-793. (33) Massalongo. A. B. 1859a. Specimen photographicum animalium quorumdam plantarumque fossilium agri Veronensis. Veronae: Vincentini-Franchini. [Dual text in Italian and Latin.] (34) Massalongo. A. B 1859b. Syllabus plantarum fossilium hucusque in formationibus tertiariis agri Ve ne ti detectarum. Veronae: A. Merlo. (35) Melchior. H. 1964. Reihe Microspermae (Orchidales, Gynandrae). Pp. 613-625 in H. Melchior [ed.]. A. Engler’s
  • 11. Syllabus der Pflanzenfamilien. 12. Aufl. Bd. 2. Angiospermen. Berlin-Nikolassee: Gebrüder Borntraeger. (35a) Menard. H. W. 1 9 7 1 . Science: growth and change. Cambridge, Mass.: Harvard University Press. (36) Meschinelli, A., and X. Squinabol. 1893. Flora tertiaria Ittalica. Patavii: Sumptibus Auctorum Typis Seminarii. (37) Naegeli O. 1905. Ueber westliche Florenelemente in der Nordostschweiz. Ber. Schweiz. Bot. Ges. 15:14-25. (38) Novak. F. A. 1961. Vyssi rostliny: Tracheophyta. Praha: Nakladatelství Ceskoslovenské Akademie Véd. (39) Pijl, L. van der. 1966. Pollination mechanisms in orchids. Pp. 61-75 in J. G. Hawkcs [ed.]. Reproductive biology and taxonomy of vascular plants. Oxford: Pergamon Press. (40) Pijl, L. van der, and C. H. Dodson. 1966. Orchid flowers: their pollination and evolution, Coral Cables. Florida: University of Miami Press. (41) Potonié. R. 1967. Versuch der Einordnung der fossilen Sporae dispersae in das phylogenetische System der Pflanzenfamilien. Forschungsber. Landes Nordrhein-Westfalen Nr. 1761:1-310. (42) Rolfe. R. A. 1909-12. The evolution of the Orchidaceae. Orchid Rev. 17:129-132, 193-196. 249-252. 289-292. 353-356; 18:33-36, 97-99. 129-132, 162-166, 289-294. 321-325; 19:68-69. 289-292: 20:204-207, 225-228. 260-264. [General discussion in 20:225-228, 260-264.] (43) Schimper, W. P.. and A. Schenk. 1879-90. Palaeophytologie. Abt. 2 in K. A. Zittel [ed.], Handbuch der Palaeontologie. Münchcn: R. Oldenbourg. [Also the 1891 translation into French by C. Barrois et al.: Paleophytologie. Pt. 2 in K. A. Zittel [ed.]. Traité de paléontologie.] (44) Schultcs. R. E. 1960. Native orchids of Trinidad and Tobago. New York: Pergamon Press. (45) Schultes, R. E. 1966. Orchid [in part]. Pp. 1041-1043 in Encyclopaedia Brittanica. Vol. 16. Chicago: Encyclopaedia Brittanica. [Also in subsequent editions.] (46) Schultes. R. E., and A. S. Pease. 1963. Generic names of orchids: their origin and meaning. New York: Academic Press. (47) Sernander, R. 1894. Studier öfver den Gotländska vegetationens utvecklingshistoria, Ph.D. Thesis, Universitet i Uppsala. 1 1 2pp. [Privately printed.] (48) Stebbins, G. L. Jr. 1950. Variation and evolution in plants. New York: Columbia University Press. (49) Straus, A. 1954. Beiträge zur Pliocänflora von Willershausen. IV. Die Monocotyledonen. Palaeontographica 96B:1-11
  • 12. (50) Straus, A. 1969. Beiträge zur Kenntnis der Pliozänflora von Willershausen (VII). Die Angiospermen- Früchte und -Samen. Argumenta Palaeobotannica 3:163 197. (51) Thurmann, J. [1833.] Essai sur les soulèvemens Jurassiques du Porrentruy, avec une description géognostique des terrains secondaires de ce pays, et des considérations générales sur les chaines du Jura. Mém. Soc. Hist. Nat. Strasbourg 1 (livre. 2, article "L"):l-84. (52) Vent, W. 1965. Neue Pflanzenfunde aus den interglazialen Ilmtaltiavertinen von Weimar-Ehringsdorf. Geologie 14:198-205. '