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C.H. Pander's introduction to conodonts, 1856

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It is rare in the annals of science for a single publication to provide a definitive introduction to a previously unknown group of animals. Yet this was the case with the monograph that included the initial descriptions of an important, extinct group of fossil animals: the conodonts. The publication, by Christian Heinrich Pander, appeared in St Petersburg in 1856. The Volume of ninety-one pages, written in German and accompanied by seven beautifully executed plates, was issued by the Imperial Academy of Sciences as part of a series entitled Geognostische Beschreibung der Russisch-Baltischen Gouvernements. For the seventy years following 1856, Pander's description of conodonts provided an essentially unmodified account of this newly discovered animal group. Furthermore, with the discovery in 1926 of the potential value of conodonts in stratigraphic correlation, this zoologically enigmatic group of fossils rapidly assumed the position of importance in Paleozoic and Triassic biostratigraphic work that it enjoys today. So, in recognition of Pander's important work, we offer this tribute in honor of the fortieth anniversary of the Pander Society, an international association of nucropaleontologists formed in his honor. Siegfried and Gross (1971) have described Pander's four monographs on Lower Paleozoic fossils as classic, and Raikov (1964), in an assessment of all of Pander's scientific contributions, regards him as the founder of Russian paleontology. He is also commonly seen as one of the founders of the field of embryology. In this contribution we focus on that part of his 1856 monograph in which he named, discussed, and analyzed conodonts.
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It is rare in the annals of science for a single publication to provide a
denitive introduction to a previously unknown group of animals.
Yet this was the case with the monograph that included the initial
descriptions of an important, extinct group of fossil animals: the con-
odonts. The publication, by Christian Heinrich Pander, appeared in
St Petersburg in 1856. The volume of ninety-one pages, written in
German and accompanied by seven beautifully executed plates, was
issued by the Imperial Academy of Sciences, as part of a series
entitled Geognostische Beschreibung der Russisch-Baltischen Gou-
vernements.
For the seventy years following 1856, Pander's description of con-
odonts provided an essentially unmodied account of this newly dis-
covered animal group. Furthermore, with the discovery in 1926 of the
potential value of conodonts in stratigraphic correlation, this zoologi-
cally enigmatic group of fossils rapidly assumed the position of impor-
tance in Paleozoic and Triassic biostratigraphic work that it enjoys
today. So, in recognition of Pander's important work, we offer this trib-
ute in honor of the fortieth anniversary of the Pander Society, an inter-
national association of micropaleontologists formed in his honor.
Siegfried and Gross (1971) have described Pander's four mono-
graphs on Lower Paleozoic fossils as classic and Raikov (1964), in
an assessment of all of Pander's scientic contributions, regards him
as the founder of Russian paleontology. He is also commonly seen as
one of the founders of the eld of embryology. In this contribution
we focus on that part of his 1856 monograph in which he named, dis-
cussed, and analyzed conodonts.
Biographical Review
Extensive biographical studies of Pander have been published in
German by Erman (1859), Lubosch (1918), Loesch (1920),
Siegfried and Gross (1971), and Knorre (1973). Schmitt (2005),
which emphasizes Pander's contribution to embryology, is a recent
contribution in English. Our review relies heavily on Raikov (1964),
which was written originally in Russian but was issued in a German
edition in 1984 with accompanying English summaries and com-
mentary by W. E. von Herzenberg and P. H. von Bitter. We extract
the following from the latter report.
Christian Heinrich
Pander (Figure 1) was born
in Riga, Latvia (then Liv-
land or Livonia) on 23 July
1794, to a prosperous mer-
chant-banker, Johann Mar-
tin Pander, and his wife
Ursula. He had three broth-
ers and four sisters. Pander's
early training in Riga was in
a German-speaking gymna-
sium, from which he gradu-
ated in 1812, aged eighteen.
Between 1812 and 1817 he
took up the study of medi-
cine in Dorpat (now Tartu,
Estonia), and subsequently
in universities in Berlin,
Göttingen, and Würzburg.
In 1817, he received a doc-
torate in medicine in
Würzburg. His dissertation
concerned the development
of the chick in the egg and is widely regarded as a fundamental con-
tribution to the eld of embryology.
Between 1818 and 1820, Pander toured European museums to
study animal skeletons in preparation for a planned work with
Eduard d'Alton on vertebrate anatomy, and served a brief period in
Tartu as editor of a scientic journal. In 1820, Pander became an
adjunct in zoology in the Academy of Sciences in St Petersburg and,
in that capacity he joined a Russian diplomatic mission to the Emi-
rate of Bukhara in Central Asia. In 1821, the rst of fourteen install-
ments of a series titled 'Comparative Osteology' and co-authored
with d'Alton was published in Bonn. During this period, Pander's
research focused primarily on the comparative anatomy of living and
fossil vertebrates. Within one of the publications that resulted from
this research, Pander (1824) formulated ideas on 'uninterrupted
change or metamorphism' in plants and animals in an essay entitled
'General observation about the outside inuences on the organic
development of animals'. This contribution was later cited by
Charles Darwin as one of the forerunners of his hypothesis that the
evolution of living organisms is guided by natural selection.
In 1825, Pander married Amalie von Scherer, with whom he
ultimately fathered three sons and four daughters. Shortly after his
marriage, while on sick-leave from the Academy, he conducted eld
studies in southern Russia and, upon return to St Petersburg in 1826,
he became a regular member of the Academy. He resigned a year
later, however, discouraged by the Academy's rules and practices.
From 1827 to 1833, Pander conducted independent studies in
the surrounding area of St Petersburg. In 1830, he published at his
own expense, a 165-page report on the geology of the St Petersburg
region. This report, accompanied by a geological map and thirty-two
hand-colored plates illustrating various fossils, was a pioneering
study that established Pander as the leading authority on geology of
this segment of the Baltic region, several years before publication of
the work of Murchison and Sedgwick. It also earned him recognition
(many years later) as the father of paleontology in Russia.
In 1833, Pander was persuaded by his father to return to the
family estate, Zarnikau, near Riga, where he and his growing family
remained until 1844. During this eleven-year period Pander traveled
widely in the Baltic provinces, extending his knowledge of their
geology and making additional collections of fossils from the Lower
Paleozoic strata exposed there. His fame as the authority on the geol-
ogy of this region was conrmed by a visit to Zarnikau in 1841 by
Murchison, Verneuil, and Keyserling on their way home from their
studies in the Urals. During this visit, Pander almost certainly exhib-
ited the microfossils he later named Conodonten, and solicited opin-
ions as to their zoologic affinities.
Pander's father died in 1842, and this greatly altered his son's
nancial situation. Thus, two years later, in 1844, Pander returned to
St Petersburg, where he found employment as a specialist responsi-
ble for 'special tasks' in the scientic section of the Department of
Mines. This period in Pander's life saw the publication of important
reports and papers on a variety of geological and paleontological
subjects, including further information on the bedrock formations of
the St Petersburg region; the possibility of salt in the vicinity of Nov-
gorod; coal on both slopes of the Urals; and, of course, his four
extensive reports on fossil Paleozoic shes. The rst of these mono-
graphs, issued in 1856, included the rst report of conodonts, the
subject of the present paper. Completion of the conodont monograph
was especially creditable as Pander suffered from eye problems that
inhibited his use of a microscope.
Pander died on 22 September 1865, following an operation for
kidney stones. He was buried in the Smolenska Lutheran Cemetery
in St Petersburg, but his grave has not been preserved.
Episodes, Vol. 31, No. 4
Classic Paper 433
C. H. Pander's introduction to conodonts, 1856
Figure 1 C.H. Pander in middle
age, pen and ink drawing from a
photograph; also in Raikov (1964,
p. 15).
The content of Pander (1856)
Following an extended six-page introduction, Pander's 1856 mono-
graph, titled Monographie der Fossilen Fische der Silurischen Sys-
tems der Russisch-Baltischen Gouvernements, was divided into two
major parts. The rst, thirty-four pages long, was titled Untersil-
urische Fische. This section consisted of Pander's detailed interpre-
tation and careful taxonomic treatment of Conodonten. This newly
recognized suprageneric taxon was based on Pander's study of thou-
sands of minute fossils he had collected from what would now be
regarded as Ordovician and Silurian strata in various parts of the
Baltic provinces and from Devonian and Carboniferous rocks as far
away as the environs of Moscow. The second and somewhat longer
section was forty-one pages long and dealt in detail with extended
description and careful taxonomic treatment of a large collection of
mostly fragmentary fossil shes.
The 1856 monograph was accompanied by a full-page text-g-
ure depicting cross-sections of conodont elements and eight large,
foldout plates of which the rst four (see Figures 2–5) were devoted
almost entirely to carefully executed illustrations of the conodont
elements described in the rst part. In a brief addendum, following
page 79, Pander noted the receipt of additional specimens of Silurian
sh from a colleague in Tartu and proposes a new name, Lonchodus,
for the conodont genus Centrodus that he established formally on
Page 31.
Conodonts, the subjects of the rst part of Pander's 1856 mono-
graph, were considered to be the teeth of an extinct group of shes.
This group was diagnosed (in German) in terms of its fossil remains
as (our translation):
Tiny, lustrous, elongated remains very similar in shape to sh
teeth, [which] extend upward or toward one end into a point;
[which] widen gradually or abruptly downward; [which are]
more or less curved, and most [of which] have sharp margins
(keels), one in front and one in back.
Pander continued to note that the fossils were of very different
shapes; some symmetrical, some not, and were provided with faces
that were smooth or longitudinally striated, and commonly had a
sharp carina that projected laterally from the smooth surface.
Clearly, Pander based his diagnosis on what he termed simple teeth;
and in a later paragraph, he noted that compound teeth consisted of a
central tooth with rows of smaller teeth located on an expanded base.
Conodont specimens were divided by Pander into crown
(Spitze) and base (Basis). The crown was solid, the base hollow. In
line with his interpretation of these fossils as sh teeth, the hollow
base was interpreted as the site of the pulp cavity (cavitas pulpae).
Bases (and their included pulp cavities) were shown to have various
shapes, all of which were illustrated on the plates accompanying the
monograph.
Pander wrote that it was noteworthy that the conodont speci-
mens in his large collection all seemed to be largely unaltered and
retained their original color and luster. They appeared to be com-
posed of pure calcium carbonate (Reinem kohlensauren Kalk) and,
because of their unaltered condition, Pander reported that it was pos-
sible to study their internal structures, of which he recognized (and
illustrated) three types. One type was snow-white and not transparent
but with translucent margins; a second type was yellow and com-
pletely translucent; a third was reddish white, compact and com-
pletely opaque. Pander concluded that the completely translucent
specimens without white matter represented youthful stages of
growth, whereas those with white matter represented later growth
stages. He was evidently puzzled about the signicance of the mostly
compound specimens that exhibited the third type of structure.
Pander then turned to the structure of 'these teeth', which he
noted apparently differed in icthyological character from all previ-
ously known fish. This had initially given rise to the impression
that the fossil remains were not fish teeth. However, after a careful
study of the internal structure of several translucent simple speci-
mens, two of which had been sectioned longitudinally, Pander con-
cluded that the cone-shaped lamellae conformed to the shape of the
pulp cavity and had been added by internal apposition, so that the
outer surface was the oldest. He recognized and illustrated other
internal features, but was unable to see anything that he could iden-
tify as bone.
Indeed in his large collections of conodont elements Pander
found no bone or anything else that he could identify as other parts of
a conodont skeleton. Thus, he concluded that it was highly unlikely
that conodonts had any other hard parts. He suggested that this was
also the case with the teeth of such animals as hagsh and lampreys,
and that these living organisms might somehow be related.
Finally, after a careful comparison of specimens in his large
collection, Pander confessed (on page 18) that we do not know what
types of teeth we have before us, whether they belong to the jaws, the
gums, the lips, or the tongue, nor whether each particular shape
serves as the basis for establishment of its own genus and species, or
whether different forms might have come from the same body.
Unable to solve this puzzle, Pander chose in the following
pages of his monograph, to use a strict, almost mechanical form
taxonomy for the formal description of the specimens in his collec-
tion. That is, each species of conodont was interpreted to have had
teeth of a single morphology. Thus, the systematic portion of the
section of his monograph included the establishment of fifty-five
newly named species in thirteen genera, all taxa being based on the
shapes of typical specimens. Typical representatives of each
species are illustrated on the four conodont plates (see Figures
2–5).
December 2008
434
Figure 2 Plate 1 from Pander (1856) illustrating a variety of the
conodont elements that Pander termed 'simple teeth'.
Figure 3 Plate 2 from Pander (1856) illustrating both 'simple'
and 'compound' teeth. Original is a large fold out measuring
about 23–35cm.
The next 150 years
For some seventy years after the publication of Pander's 1856 mono-
graph, conodonts seem to have been regarded largely as paleonto-
logical curiosities, and there was continued speculation as to their
zoological affinities. One contribution in this period stands out how-
ever. This was a paper by G. J. Hinde (1879) in which he not only
described a number of Ordovician and Devonian conodont speci-
mens that he had collected in North America, but also reviewed Pan-
der's monograph. He noted the lamellar internal structure of con-
odont elements that Pander had described but found specimens in
which the basal body appeared to lack such a structure. Such bodies,
later termed basal llings, were not preserved in the specimens in
Pander's collection and are also absent in the majority of specimens
in other collections. Hinde agreed with Pander's description of the
white structure of some specimens, but interpreted it to be the result
of weathering, not the internal dissolution in late growth stages. In
addition he concluded that a number of differently-shaped speci-
mens found clustered together on a black shale slab represented a
single individual. These he named Polygnathus dubius. Hinde also
reported that he had shown his specimens to Huxley, who agreed
with Pander that myxinoid shes were probably the conodonts' clos-
est relatives. And, nally, Hinde's collections included laterally
expanded, plate-like specimens of a type not described by Pander.
These were termed pectinate specimens.
The next major publication dealing with conodonts, a lengthy
bulletin by E. O. Ulrich and R. S. Bassler, appeared in 1926 and was
based on U. S. National Museum collections from Devonian and
Mississippian rocks in the southeastern United States. In the intro-
duction to this bulletin, the authors noted their agreement with Pan-
der that myxinoid shes were probably the closest relatives of some
conodonts, but suggested that sharks and other sh groups might
also have been involved. They emphatically disagreed with Hinde
that some conodonts might have had 'teeth' of several different
shapes, and based their comprehensive classication of the con-
odonts on a rigorous form taxonomy, which they admitted might be
somewhat articial. The principal contribution of the Ulrich–Bassler
study was its demonstration of the fact that conodonts might be used
successfully in the biostratigraphic correlation of strata.
As they predicted, the study by Ulrich and Bassler sparked an
interest in the biostratigraphic use of conodonts in the United States.
For example, in 1926, Roundy described collections from Mississip-
pian rocks in Texas; Stauffer (1930) illustrated material from the
Middle Ordovician; Gunnell (1931) described conodonts from the
Fort Scott Limestone of Missouri; and Stauffer and Plummer (1932)
reported on collections from the Pennsylvanian of Texas. In 1933,
E. B. Branson and M. G. Mehl of the University of Missouri pub-
lished a series of reports collectively titled 'Conodont Studies', which
documented the occurrence of conodonts from the middle of the
Ordovician through the lower part of the Carboniferous (Mississip-
pian) of eastern and central United States. Furnish (1938) added a
study of Lower Ordovician faunas, and, by mid-twentieth century,
the known stratigraphic range of conodonts was from late Cambrian
to latest Triassic. During this period Pander's name continued to be
remembered by researchers in the proposal of new conodont taxa,
for example in the genera Panderodella, Panderodus, Panderolepis,
Pandorina, and Pandorinellina.
Biostratigraphers, enthused by the correlation potential of con-
odonts, spent little time on studying the composition and structure of
conodont specimens. But ultimately it was determined that speci-
mens are composed largely of francolite, a carbonate apatite, and
include traces of at least thirty-nine other elements. Furthermore,
laminae in element crowns include, or are separated by, thin lms of
un-mineralized tissue that darken gradually as elements are heated
and more of the carbon in this organic material becomes xed. Also,
it has been demonstrated that conodont elements grew by external
apposition so that the outer surface of these specimens is the
youngest, not the oldest as Pander concluded.
The discovery of additional clusters of elements in the 1930s
and 1950s and, in the 1960s, the determination that groups of differ-
ently shaped elements formed recurrent associations, paved the way
to the development of a multi-element taxonomy, which Pander had
considered a possibility back in 1856, and which is now the gener-
ally accepted basis for understanding and describing conodont
species.
In 1856, Pander noted that simple conodont specimens domi-
nated his Lower Ordovician collections but that collections from
younger rocks were composed mostly, or entirely, of compound
forms. This observation has been repeatedly conrmed by post-1856
studies and is, in essence, the basis for recognizing biostratigraphi-
cally useful successions of conodont species.
The especially important discovery, in 1983, of several Scottish
specimens that represent more or less complete conodonts, and the
careful study and description of these specimens by Aldridge et al.
(1986), seems to have conrmed Pander's suspicion that the discrete
specimens in his collection were the only parts capable of preserva-
tion in a group of animals related to myxinoid or lamprey-like shes.
The position of conodonts as extinct chordates, more 'derived' than
living jawless vertebrates, but less so than those with jaws, has been
established by cladistic analysis and has been summarized recently
by Sweet and Donoghue (2001).
Finally, we should mention that the widespread occurrence of
well-preserved conodont elements, representing distinctive evolu-
tionary sequences, has attracted hundreds of students to the study of
conodonts and, through the establishment of the Pander Society in
Episodes, Vol. 31, No. 4
435
Figure 4 Plate 2A from Pander (1856) illustrating conodont
elements of the type that Pander termed 'compound' teeth. The
original is a large fold out measuring about 23–35 cm.
Figure 5 Plate 3 from Pander (1856). Specimens on the left are
'simple' and 'compound' conodont elements, several showing
internal laminated structures. Specimens on the right side are not
conodonts, but sections of brachiopod shells and one ostracode.
The original is a large fold out measuring about 23–35 cm.
1967 has promoted widespread exchange of ideas about this once
ubiquitous group of marine animals. The Society is an informal
association that presently has 184 members in some thirty-three
countries. It has published and distributed a newsletter annually
since the year of its founding, and on occasions it awards a gold
medal to a member for outstanding contributions to the study of
conodonts. Pander would surely be pleased to read the many reports
of the last forty years that have demonstrated the credibility of a
multi-element taxonomy based on the geographic and stratigraphic
co-occurrence of groups of differently shaped elements, rather than
on the rigorously mechanical taxonomy based on the shape of indi-
vidual elements that he reluctantly accepted in 1856. Pander would
be even more delighted, we believe, in studies that conrm his con-
clusion that conodonts may reliably be classied as an independent
group of primitive vertebrates
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Walter C. Sweet
School of Earth Sciences
The Ohio State University
Columbus, OH 43210
USA
Barry J. Cooper
Primary Industries and Resources South Australia
GPO Box 1671
Adelaide SA 5001
AUSTRALIA
December 2008
436
... A recent paper by Sweet and Cooper (2008), within the classic paper series of Episodes, drew our attention and prompted this response. Their paper concerned the discovery and the concept of conodonts by Christian Heinrich Pander (1856). ...
... Their paper concerned the discovery and the concept of conodonts by Christian Heinrich Pander (1856). He was the first scientist to describe conodonts, which are phosphatic microfossils originally found in the Ordovician-Lower Carboniferous of the East Baltic area (see Sweet and Cooper, 2008). In his seminal paper, Pander (1856, p. 5) gave the first diagnosis of conodonts: " Kleine glänzende, längliche, nach oben oder gegen das eine Ende spitz verlaufende, nach unten sich allmälig oder plötzlich erweiternde, bald mehr bald weniger gekrümmte, meist mit scharfen Rändern (Kielen), einem vorderen und einem hinteren, versehene, den Fischzähnen an Gestalt sehr ähnliche Überreste. ...
... " , that is " Tiny, lustrous, elongated remains very similar in shape [our emphasis] to fish teeth, [which] extend upward or toward one end into a point; [which] widen gradually or abruptly downward; [which are] more or less curved, and most [of which] have sharp margins (keels), one in front and one in back. " (Sweet and Cooper, 2008, p. 434). However, the assertion of Sweet and Cooper (2008, p. 434) that " Conodonts, the subjects of the first part of Pander's 1856 monograph, were exclusively considered to be the teeth of an extinct group of fishes " has to be questioned. ...
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The term vertebrate is generally viewed by systematists in two contexts, either as Craniata (myxinoids or hagfishes + vertebrates s.s., i.e. basically, animals possessing a stiff backbone) or as Vertebrata (lampreys + other vertebrae-bearing animals, which we propose to call here Euvertebrata). Craniates are characterized by a skull; vertebrates by vertebrae (arcualia); euvertebrates are vertebrates with hard phosphatised tissues in the skeleton. The earliest known possible craniate is Myllokunmingia (syn. Haikouichthys) from the Lower Cambrian of Chengjiang, south China. Euvertebrates appear in the Ordovician. C. H. Pander is sometimes thought to have been the first to propose that conodonts are vertebrates, but he did have doubts about the fish affinities of conodonts. This proposal was revived in the 30s and especially in the 80s of the 20th century and given elevated status in 2000 through a cladistic analysis based upon interpretation of conodont mineralized tissues as homologous to those of vertebrates. This analysis resolved conodonts within the clade Vertebrata s.s., and incorporated a 'Total Group Concept' (TGC), including conodonts in the TG Gnathostomes (= jawed vertebrates). This resulted in the unusual scenario in which "teeth" appear before jaws. We reject the TGC nomenclature as applied to early vertebrates. In addition, based on all existing evidence, we consider that conodont hard tissues and several other anatomical structures in conodonts are not homologous with those of vertebrates. Making a revised cladistic analysis, eliminating characters unknown in fossils, conodonts appear stemward (i.e. more basal) to craniates and are thus interpreted as basal chordates at best. To help resolve the phylogenetic relationships of conodonts and chordates, the analysis should be extended to include non-chordate taxa.
... A recent paper by Sweet and Cooper (2008), within the classic paper series of Episodes, drew our attention and prompted this response. Their paper concerned the discovery and the concept of conodonts by Christian Heinrich Pander (1856). ...
... Their paper concerned the discovery and the concept of conodonts by Christian Heinrich Pander (1856). He was the first scientist to describe conodonts, which are phosphatic microfossils originally found in the Ordovician-Lower Carboniferous of the East Baltic area (see Sweet and Cooper, 2008). In his seminal paper, Pander (1856, p. 5) gave the first diagnosis of conodonts: " Kleine glänzende, längliche, nach oben oder gegen das eine Ende spitz verlaufende, nach unten sich allmälig oder plötzlich erweiternde, bald mehr bald weniger gekrümmte, meist mit scharfen Rändern (Kielen), einem vorderen und einem hinteren, versehene, den Fischzähnen an Gestalt sehr ähnliche Überreste. ...
... " , that is " Tiny, lustrous, elongated remains very similar in shape [our emphasis] to fish teeth, [which] extend upward or toward one end into a point; [which] widen gradually or abruptly downward; [which are] more or less curved, and most [of which] have sharp margins (keels), one in front and one in back. " (Sweet and Cooper, 2008, p. 434). However, the assertion of Sweet and Cooper (2008, p. 434) that " Conodonts, the subjects of the first part of Pander's 1856 monograph, were exclusively considered to be the teeth of an extinct group of fishes " has to be questioned. ...
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The term vertebrate is generally viewed by systematists in two contexts, either as Craniata (myxinoids or hagfishes + vertebrates s.s., i.e. basically, animals possessing a stiff backbone) or as Vertebrata (lampreys + other vertebrae-bearing animals, which we propose to call here Euvertebrata). Craniates are characterized by a skull; vertebrates by vertebrae (arcualia); euvertebrates are vertebrates with hard phosphatised tissues in the skeleton. The earliest known possible craniate is Myllokunmingia (syn. Haikouichthys) from the Lower Cambrian of Chengjiang, south China. Euvertebrates appear in the Ordovician. C. H. Pander is sometimes thought to have been the first to propose that conodonts are vertebrates, but he did have doubts about the fish affinities of conodonts. This proposal was revived in the 30s and especially in the 80s of the 20th century and given elevated status in 2000 through a cladistic analysis based upon interpretation of conodont mineralized tissues as homologous to those of vertebrates. This analysis resolved conodonts within the clade Vertebrata s.s., and incorporated a ‘Total Group Concept’ (TGC), including conodonts in the TG Gnathostomes (= jawed vertebrates). This resulted in the unusual scenario in which “teeth” appear before jaws. We reject the TGC nomenclature as applied to early vertebrates. In addition, based on all existing evidence, we consider that conodont hard tissues and several other anatomical structures in conodonts are not homologous with those of vertebrates. Making a revised cladistic analysis, eliminating characters unknown in fossils, conodonts appear stemward (i.e. more basal) to craniates and are thus interpreted as basal chordates at best. To help resolve the phylogenetic relationships of conodonts and chordates, the analysis should be extended to include non-chordate taxa.
... Logically this volume commences with Christian Heinrich Pander (1794-1865), long-time resident of modern day Latvia, then part of the Russian Empire and the unchallenged conodont discoverer, who make his remarkable finding sometime around 1848-1850. Pander remains a hero because his 1856 monograph, published in German in St Petersburg, stands as a classic paper that provided much correct assessment despite many subsequent reinterpretations (Sweet & Cooper 2008). Today conodont researchers gather as the "Pander Society". ...
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Full-text available
The word vertebrate is generally viewed by systematists in two contexts, either as Craniata (myxinoids or hagfishes + vertebrates s.s., i.e. basically, animals possessing a backbone and a skull) or as Vertebrata (lampreys + other vertebrae-bearing animals, also known as Euvertebrata). Craniates are characterized by a skull; vertebrates by vertebrae (arcualia); euvertebrates are vertebrates with mineralized (phosphatised) tissues in the skeleton. The earliest known possible craniate is Myllokunmingia (syn. Haikouichthys) from the Lower Cambrian Lagerstätte of Chengjiang, south China. Euvertebrates appear in the Ordovician. Christian H. Pander is sometimes thought to have been the first to propose that conodonts are vertebrates, but he had doubts about the fish affinities of conodonts. This proposal was rejuvenated in the 30s and especially in the 80s of the 20th century and given elevated scientific status in 2000 through a cladistic analysis based upon interpretation of conodont mineralized tissues as homologous to those of vertebrates (that is enamel, dentine, bone). This analysis resolved conodonts within the clade Vertebrata s.s., and incorporated a ‘Total Group Concept’ (TGC), including conodonts in the Total Group Gnathostomes (= jawed vertebrates). This resulted in the unusual evolutionary scenario in which ‘teeth’ appear before jaws. The TGC nomenclature as applied to early vertebrates by ‘the British School’ has been rejected. Conodont hard tissues and several other anatomical structures in conodonts cannot be considered as homologous with those of vertebrates. Based upon a revised cladistic analysis, where characters unknown in fossils were rejected, conodonts appear stemward (i.e. more basal) to craniates and are interpreted as basal chordates at best. To help resolve the phylogenetic relationships of conodonts and chordates, the analysis should be extended to include non-chordate taxa.
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Diversified continental, marginal-marine and marine epicontinental deposits in Poland of Early Jurassic age yielded unusually rich palynomacerals. Numerous parameters, such as the presence of acritarchs and dinoflagellate cysts, spore/bisaccate pollen grain ratio, content of terrestrial phytoclasts, degree of oxidation, presence of tetrads and sporangia, degree of palynomorph alteration, presence and character of amorphous organic matter, presence of epibionts on palynomorphs, were collectively found as indicative of certain palynofacies. The ratio of spores and bisaccate pollen grains significantly depends on the climatic conditions where also seasonal changes may influence the local characteristics of palynofacies. However, the ratio also strongly reflects the local depositional environment which may vary regionally and spore/bisaccate pollen grain ratio can be taken as a general indicator of distance from the shore. Early Jurassic palynomacerals from the Polish Basin are strongly dominated by terrestrial elements, marine palynomacerals occur in significant quantities only in Pliensbachian deposits in Pomerania Western Poland. Charcoal is an important component of palynomacerals. Due to its resistance to biogenic degradation and buoyancy, charcoal produced by extensive wildfires was widely re-deposited and concentrated particularly in foreshore to shallow shoreface and delta plain environments. Three types of palynofacies inversions (abnormal palynofacies composition) are discussed. Six main palynofacies types linked to depositional systems previously determined by sedimentological studies have been distinguished providing a robust paleoenvironmental tool for recognition of palynofacies attributed to certain palaeoenvironments.
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Conodonts were mostly small, elongate, eel-shaped marine animals that inhabited a variety of environments in Paleozoic and Triassic seas. Although long enigmatic, conodonts are now regarded as vertebrates and their closely controlled fossil record is not only the most extensive of all vertebrates, but it also makes conodonts the fossils of choice in upper Cambrian through Triassic biostratigraphy. Conodonts were soft-bodied except for a variety of phosphatic elements that formed a distinctive feeding apparatus. Post-mortal dissociation of the apparatus and subsequent jumbling of its elements on the sea floor led, from 1856 to about 1966, to development of an artificial, form-based taxonomy that was utilitarian, but clearly unsatisfactory as a vehicle for understanding the group in biologic terms. Natural assemblages of elements, discovered between 1879 and 1952, have been interpreted as undisturbed skeletal apparatuses, and in the mid-1960s it was determined that original composition of the apparatuses of many species could be reconstructed and statistically evaluated from collections of disjunct elements by various grouping procedures. These determinations led to an emphasis on multielement taxonomy by most (but not all) students of conodonts. Even so, only about a third of the approximately 550 valid conodont genera, have been established (or re-interpreted) in multielement terms and this makes any of the several extant schemes of suprageneric classification phylogenetically suspect. We comment on a recent scheme that recognizes 41 families assigned to some 7 orders, and suggest how it might be modified so as to square with principles of phylogenetic systematics.
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Full-text available
Conodonts were mostly small, elongate, eel-shaped marine animals that inhabited a variety of environments in Paleozoic and Triassic seas. Although long enigmatic, conodonts are now regarded as vertebrates and their closely controlled fossil record is not only the most extensive of all vertebrates, but it also makes conodonts the fossils of choice in upper Cambrian through Triassic biostratigraphy. Conodonts were soft-bodied except for a variety of phosphatic elements that formed a distinctive feeding apparatus. Post-mortal dissociation of the apparatus and subsequent jumbling of its elements on the sea floor led, from 1856 to about 1966, to development of an artificial, form-based taxonomy that was utilitarian, but clearly unsatisfactory as a vehicle for understanding the group in biologic terms. Natural assemblages of elements, discovered between 1879 and 1952, have been interpreted as undisturbed skeletal apparatuses, and in the mid-1960s it was determined that original composition of the apparatuses of many species could be reconstructed and statistically evaluated from collections of disjunct elements by various grouping procedures. These determinations led to an emphasis on multielement taxonomy by most (but not all) students of conodonts. Even so, only about a third of the approx-imately 550 valid conodont genera, have been established (or re-interpreted) in multielement terms and this makes any of the several extant schemes of suprageneric classification phylogenetically suspect. We comment on a recent scheme that recognizes 41 families assigned to some 7 orders, and suggest how it might be modified so as to square with principles of phylogenetic systematics.
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Three new specimens which preserve the soft parts of conodonts are described from the Lower Carboniferous of Granton, Edinburgh. The animals was apparently laterally flattened in life and the somites were V-shaped. The nature of the preserved axial lines is equivocal; some may represent the walls of the gut. The elements of one of the new specimens show that it does not belong to Clydagnathus, to which the other soft-bodied specimens from Granton was tentatively assigned. The possibility of a relationship between the euconodonts and the Chaetognatha is discounted. Nor do the conodonts constitute a phylum, but are a separate group of primitive jawless craniates.-Authors
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Soon after the publication by Dr. Pander, in 1856, of his wellknown monograph, in which he announced the discovery, in the lowest fossiliferous rocks of Russia, of small ‘teeth” named Conodonts, and referred by him to fishes, several discussions arose as to the character of these minute bodies, and various opinions were expressed as to their near relations, without, however, any satisfactory conclusion being arrived at. Since that date Conodonts have been found in several other places; and in this communication I propose to describe a great variety of forms which I have collected within the last two years from several different formations in North America. Though my specimens may not suffice to determine the true position of the organisms to which they were attached, they will at least add something to our previous knowledge, and thus assist in reaching a decision on the subject. The next account, after Dr. Pander's, of the discovery of Conodonts is by Dr. J. Harley, in an article on the Ludlow bone-bed and its crustacean remains. Only two of the specimens described by this gentleman have any resemblance to the Conodonts of Pander; but a comparison of these with other very differently formed bodies in the same beds led him to express the opinion that all the forms were of crustacean origin, and that Conodonts were probably only spines similar to those attached to the margins of the carapace of Limulus and the caudal segment of Squilla. He therefore included all together under the provisional genus Astacoderma.
Article
Christian Heinrich Pander (1794-1865), a Russian scientist of German culture, is known for his epoch-making work in embryology, as well as for his important contributions to palaeontology. Indeed he viewed embryonic development and the history of the earth as two aspects of one and the same essential phenomenon, namely, a perpetual metamorphosis affecting the living world on different scales. He viewed embryology as a gradual, epigenetic transformation (as opposed to preformation) with an intermediary stage, the formation of simple germ-layers. As early as 1821, he argued more generally that species themselves transform under the influence of certain environmental factors. Pander thus embodies the very close link that existed between the triumph of epigenesis and the expansion of transformist theories in the early 19th century.
Monographie der fossilen Fische des silurischen Systems der russisch-baltischen Gouvernements St Petersburg: Buchdruckerei der Kaiserlichen Akademie der Wissenschaften, 91 pp., 9 plates Raikov German Translation with Commentary and English Summaries by
  • C H Pander
Pander, C. H., 1856, Monographie der fossilen Fische des silurischen Systems der russisch-baltischen Gouvernements. St Petersburg: Buchdruckerei der Kaiserlichen Akademie der Wissenschaften, 91 pp., 9 plates Raikov, B. E., 1964, Christian Heinrich Pander, ein bedeutender Biologe und Evolutionist, 1794–1865. German Translation with Commentary and English Summaries by Hertzenberg, W. E. von and Bitter, P. H. von, 1984: Senckenberg-Buch, v. 62, 144 pp.