Protea Obtusifolia, Variety Jester, Family Proteaceae         Photo by the author.

A bold solution to 'Darwin's Mystery' is offered here

Dr. Harry Levin proposes a bold solution to Darwin's "abominable puzzle" or the famous "abominable mystery" about the origins of flowering plants. This is one of "Ten Essays on the Basic Reexamination of the Late Paleozoic and the Mesozoic Eras of Earth History." Dr. Levin, whose Essay #1 The Evolution of Proteaceae, in Flower and Leaf and whose  wildflower photos enhance our pages, seeks reconsideration of the current thought that flowering plants had origins 150 million years ago or less.

Dr. Harry Levin
His latest groundbreaking essays include a proposed solution to the puzzling origin of New Zealand, which he suggests had a landlink to South America, the Vast Geological Significance of a Fish, and the The Dominance of the Dinosaur. 

A listing of those and other essays can be found on the contents page.

Theories based on the fossil record suggest there were no flowering plants more than 150 million years ago. However, as other scientists have pointed out, absence of evidence is not evidence of absence. Levin, who holds a doctorate in chemical engineering from Johns Hopkins University,  easily doubles the ante here, with his well-supported hypothesis that flowering plants existed more than 300 million years ago.

Searching broadly across fields of scholarship and adding his own interpretation, Dr. Levin marshals evidence from genetics, plate tectonics, paleoclimatology, current plant family distributions, insect evolution, and the presence of coal deposits. This evidence supports (or is consistent with) his hypothesis that flowering plants were widely distributed over the southern hemisphere supercontinent Gondwanaland prior to its breakup 280 million years ago.
"To biology and geology," writes Dr. Levin, "I bring a third viewpoint – that of the engineer. The engineer, in a best aspect of his profession, strives to resolve situations involving multivariables and their interactions. In ten essays, this engineer presents a startling reexamination of earth history. These essays provide accounts of hitherto unsuspected history of the late Paleozoic and the Mesozoic. They do so by interacting biological and geological events that occurred at the same time. The interweaving here  of geophysics and biophysics – both embracing climatology – brings out clarifying and sometimes incontrovertible evidence – content otherwise  obscure to, or hidden from, either one or the other of these disciplines when it acts alone."

-Michael E. Abrams
with Robert Levin

On the Origins of  the Angiosperm and the Gymnosperm


By Harry Levin               
Essay Number Three


The angiosperm phylum of the plant kingdom – the flowering plant – originated in Gondwanaland during the late Devonian, more than 360 million years ago.

This overarching hypothesis is founded on late-Paleozoic and Mesozoic plate tectonics and climatology here in correlation with the past and present geographic distributions of living plants. It is firmly supported by plant molecular genetics, plant morphology, and paleoentomology.

Following the saga of five plant families – three angiosperms and two gymnosperms – the essay illuminates late-Paleozoic to present-day consociations of angiosperms and gymnosperms in close correlation with late Paleozoic and Mesozoic plate tectonics and climatology. It emphasizes the essential late Paleozoic role of insects in evolution of the bisexual flower. Together with molecular genetics, it finds that angiosperms and gymnosperms are distinct and separate phyla in coeval existence since the Devonian; and it contends that both phyla are monophyletic. Significantly, this essay approximates – with examples – hitherto unknown dates of origin of individual plant and animal families.                 

Throughout this account, the time 280 Ma was momentous – a rapid pulsing of biology and geology:

     • It marked the beginning of the Permian.
     • It marked the end of the Permo-Carboniferous Ice Age, except for a continuation of
       orogenic ice cover extending from southern Antarctica into the New South Wales
       region of Australia until 250 Ma.
     • It marked the water-shed biological disconnect of Australia from Gondwanaland.
     • It marked the drifting away of a superterrane, here called Austosunda, from westernmost
       Australia and northern Gondwanaland – part to eventually settle down as continental
       core of the plateau region above India and part to become most of Myanmar (Burma). 


Prologue: Molecular Genetic Support
of Devonian Angiosperm Origin

First and foremost, the hypothesis of Devonian angiosperm origin is consistent with findings of molecular studies of plant DNA in genetic analysis. Certain of these studies below imply an early divergence of angiosperms from a Devonian plant group (probably a seed fern).

    • A. V. Troitsky et al. (1) indicate (in 1991) by rRNA sequence comparisons that both gymnosperms and angiosperms are monophyletic groups. The genealogical splitting of gymnosperm and angiosperm lineages occurred at least 360 million years ago. Magnoliales were the earliest angiosperms. They are dicotyledons. 

    • W. F. Martin et al. (2) indicate (in 1993), by chloroplast and nuclear sequence data taken together,
that angiosperms and gymnosperms were separate lineages about 330 million years ago and that the
separation of monocotyledonous and dicotyledonous lineages of angiosperms took place about
300 million years ago. 

    • T. Kh. Samigulin et al. (3) indicated (in 1999), with partial sequences of the ropC1 gene, that both angiosperms and gymnosperms are monophyletic and that none of the recent gymnosperms is sister to the angiosperm.

    • S-M Chaw et al. (4) indicate (in 2000), with mitochondrial subunit rRNA sequences, that gymnosperms are monophyletic; that, in conjecture, the angiosperms too are monophyletic and substantially older than the fossil record indicates. 

    • L.M. Bowe et al. (5) indicated (in 2000), with sequence data of evolving mitochondrial genes, cox1 and atpA, that extant gymnosperm genes are monophyletic and that angiosperm origin should be sought among extinct seed plant groups.

The above molecular genetic papers suggest angiosperm monophylesis. They are concordant with the forthcoming hypothesis of this essay that the angiosperm originated in the late Devonian more than 360 million years ago, – a thesis founded here on late Paleozoic and Mesozoic plate tectonics and climatology in correlation with the past and present geographic distributions of living plants.   

The above-cited phylogenetic papers refute the Cretaceous theory of angiosperm origin. The Cretaceous theory professes the abrupt appearance of a diverse angiosperm flora in the Cretaceous fossil record of the northern hemisphere, with fallacious implication of polyphylesis.  Moreover, here are cited specific genetic refutations of the anthophyte theory (6), which would link angiosperms with the gymnosperm order Gnetales (7) (8) and (indirectly) other theories which tentatively link the angiosperm bisexual flowers to conifers. Such theories could imply angiosperm origin in the early Mesozoic.

However, it must be said that there is not as yet (in 2007) full acceptance of the molecular genetic theory of the angiosperm as late-Paleozoic “sister to the gymnosperm.”(9)). Indeed, R.M. Bateman et al. (10) sum up (in 2006) that while most molecular genetic studies indicate that all extant gymnosperms form a natural group, “suggesting early divergence of the lineage that led to angiosperms”; yet “it is possible that extinct gymnosperms gave origin to the angiosperms”. The only extinct gymnosperms may have been the Glossopteridales (Permian). 


Part I: Evidence of Early Origin of Angiosperms

Assemblage of Five Plant Families

Evidence is presented that early angiosperm and gymnosperm families were extant more than 300 million years ago over a vast southern hemisphere supercontinent, Gondwanaland, comprising South America, Africa, India, Antarctica, Australia, and smaller terranes.  

For this endeavor, five plant families are selected, three of angiosperms, two of gymnosperms, which today have remarkably similar highly disjunctive southern hemisphere distributions. These are trees and shrubs that consociate with each other, in southernmost South America as well as in the South Pacific regions of Australia, New Zealand and New Caledonia – on islands and on continents nine thousand kilometers apart, on both flanks of Africa.  A key observation, detailed below, is the near-complete absence of these five exemplar plant families from Africa.

These are:

    • the angiosperm family Proteaceae
    • the angiosperm family Winteraceae
    • the angiosperm family Nothofagaceae
    • the gymnosperm family Araucariaceae
    • the gymnosperm family Podocarpaceae

The angiosperms Proteaceae, Winteraceae, and Nothofagaceae have basal dicotyledons, indicative of a dicot nature of very early angiosperms. Interestingly, the foliage of Proteaceae, Araucariaceae, and Podocarpaceae reveals a transition from broad-leafed to needle species. These trees and shrubs display a sequence of evolutionary curtailment of leaf form and structure indicative of defensive adaptation to gradual but varied encroachments of ice fields as probable cause. Phylogenetic information and present-day geographical distribution of these southern hemisphere exemplar families are provided in Table I (at end of essay). 

Supercontinental Continuity

In hypothesis, all five floras more than 300 million years occupied the whole of the southern hemisphere supercontinent Gondwanaland which was in continuity from South America to Australia.

The first to discern this continuity was J. D. Hooker (11) about 1860. A notable contemporary of Darwin, Hooker believed that the evidence of  similar flora on widely separate lands could not be explained without supposing that these lands once formed a continuous expanse (on the basis of his 1839-1843 voyages and observations with the H. M. Discovery ships Erebus and Terror). Gondwanaland had not been conceived of by 1860. By 1970, V.K. Rao said of Proteaceae, on observing their similarity in far-apart locations (12):

“The similarity in plant associations and the component floristic elements in the two [Australia and Africa] widely separated geographical regions presupposes the existence of similar edaphic and climatic conditions not only at the present time but also in the historic past and through the period of their evolution. …. The conclusion that Proteaceae originated on a connected southern continent which subsequently fragmented, therefore, seems to be unassailable.” 

Throughout earth history, it was almost axiomatic that continents did not move. It was not until 1971 that the concept of continental drift emerged  from heresy to first principles. It was then that the concept of a unified  Gondwanaland – consisting of South America, Africa, Arabia, India,  Australia, and Antarctica, as well as smaller tectonic units – became  generally acceptable. Today sequences of rock that were laid down over a 500-million-year period – before Africa and South America separated –  are almost identical on both sides of the rift, in continents now some 5,000 km apart. At the start of the Permian 280 million years ago,  Madagascar was attached to India and was near to Africa. Moreover, India, Africa, and Antarctica were either attached or in close proximity to each other through Australia.

To emphasize, all of the above five flora more than 300 million years ago are inferred here to have occupied the whole of the southern hemisphere supercontinent Gondwanaland, which was continuous from South America through Australia.


The Permo-Carboniferous Ice Age

The presence of the three angiosperms and the two gymnosperms 300 million years ago throughout Gondwanaland is strongly indicated by the great effect of a momentous glaciation, the Permo-Carboniferous Ice Age, also termed the Gondwanaland Ice Age. In hypothesis, this glaciation is a prime cause of the similarly disrupted distributions of these five families.

Beginning 300 million years ago, and lasting 50 million years, the Permo-Carboniferous Ice Age dwarfed recent glaciations, like those of the Pleistocene. The incursions of the Permo-Carboniferous ice fields destroyed all life in their paths without trace. The movements of the ice fields are described by D.H. Tarling and M.P. Tarling (13). In initial onslaughts, Southern Africa and Southeastern South America were engulfed. A huge white salient reached into Central Africa; and in South America, ice cover extended into the lower part of Brazil. Only the flora to the north, beyond the ice fields, survived. Not all of Gondwanaland was glaciated. The absence of glacial rubble today in Northeastern Australia and in some areas of Southwestern Australia indicates that these areas had eluded the polar ice sheet. This absence could account for the survival of Australian Proteaceae and other Gondwanaland angiosperms and gymnosperms despite the pervasive cold. Today, more than half of all genera of Proteaceae are in Eastern Australia, and about a third of are in Southwestern Australia.

In hypothesis, the three angiosperm and two gymnosperm families cited above were present during the Carboniferous prior to the ice invasion, and before 300 million years ago, over a continuous Gondwanaland expanse stretching from Australia to South America. Africa had been a main link along that route of propagation.

In Africa, the Permo-Carboniferous Ice Age devastated each of the five   angiosperm and gymnosperm families cited above. This imprint of the glaciation is enumerated below as it persists today:

For Proteaceae in Africa, the phylogenetic composition was altered from 14 tribes to essentially one tribe, the Proteeae.

    • For the Nothofagaceae in Africa, all traces were removed.
    • For the Winteraceae in Africa, all traces were removed.          
    • For the Araucariaceae in Africa, all traces were removed. 
    • The Podocarpaceae in Africa were wiped out; but they made a
       limited comeback along the southeast coast of Africa from Kenya to
       Zimbabwe, probably radiating by land bridge from Madagascar, as
       the climate warmed following the ice age.

The presence of these five families on both flanks of Africa combined with African absences or stark alterations asseverate that these angiosperms and gymnosperms had in the far distant past spread out over the entire Gondwanaland supercontinent; had existed there 300 million years ago; and were largely removed from Africa by the ice age of 300 to 250 million years ago.


Part II: Additional Evidence of Angiosperm Antiquity

Fragmentation and Redistribution of Gondwanaland

Gondwanaland began breaking apart during the late Paleozoic and the Mesozoic. In particular, fragments (tectonostratigraphic terranes or “megashards”) rifted away from northwest Australia and drifted north, ultimately to become attached to southeastern Laurasia. For instance, Ian Metcalfe (14) in 1988 described a group of elongated subcontinental-sized fragments, herein termed “Austosunda,” which had rifted from Northwest Australia. This breakaway occurred during the Permo-Carboniferous. Austosunda would later become part of the continental core of southeastern Asia and Indonesia. The component blocks of Austosunda are termed Lhasa, Changtang, and Sibumasu. The Lhasa and Changtang components of Austosunda became a long southern edge of Laurasia before the coming of the Indian Plate. Today Lhasa and Changtang together comprise the Tibetan Plateau. The Sibumasu component of Austosunda is now the western edge of the Indochina subcontinent. See Figure 1

                                                                                    FIGURE 1

Charting of three continental cores of Southeast Asia: Sibumasu,  lhasa, and Changtang, formerly components of the Austosunda that biolink rifted from Australia and the start of the Permian 280 million years ago to become southeastern margins or Laurasia. The Indian Plate Collided with Laurasia broadly and the Lhasa site to create the Himalayas.

The chart provides selected elements of Figure 1 of I Metcalfe, "Origin and Assembly of South-east Asian Continental Terranes," p. 102 in Gondwanaland and Tethys, ed. by M. G. Audley-Charles and A. Hallam, Oxford University Press (1988).


I. Metcalfe (14) wrote that “by latest early Permian times,” the Sibumasu component of the megashard had broken away from Australia. He noted: “It is considered here that Sibumasu, [and the Lhasa and Changtang regions of Tibet] . . .  were in continuity and rifted together at the same time from Gondwanaland.”

Today flora and fauna of Australia and Sibumasu show similarity along ancient family lines. Metcalfe cited the early-Permian identity of biological  species on both sides of the break as evidence of a previously unified biology. He pointed to “late Carboniferous-early Permian glacial-marine diamictites, cool-water faunas, absence of warm climate floras, and presence of faunas of Gondwanaland (north-west Australian) affinities” – in both Australia and continental core regions (formerly Austosunda) in  southeastern Asia. And in his summary abstract he wrote: “Late Carboniferous and Lower Permian sediments of the Sibumasu, Lhasa, and Changtang blocks . . .  include [these] extensive glacial-marine deposits.”

In hypothesis, separation of a connected group of megashards (Austosunda) from northeastern Gondwanaland occurred at the beginning of the Permian about 280 million years ago; and Austosunda arrived at its destination in southeastern Asia near the end of the Jurassic (about 145 million years ago).

These dated events (supported throughout these essays) resulted in the introduction of angiosperms into Laurasia (Eurasia) for the first time. Hence, this chronology implies that angiosperms are at least 280 million years old (consistent with the Gondwanaland angiosperm and gymnosperm antiquity presented above); and, furthermore, it explains their rather sudden appearance in the northern hemisphere Cretaceous fossil record.

By middle-to-late Cenomanian, 95 to 91 million years ago, the angiosperms, dicots and monocots, were the prevalent forms of vegetation of the northern hemisphere. In further hypothesis, the early known angiosperms in the  northern hemisphere are consistent with the arrival of the Austosunda terranes 145 million years ago. 

The earliest evidence of the presence of angiosperms in Laurasia was reported by Ge Sun et al. in ~1998 (15). They detail the discovery of fossil angiosperm fruiting axes in the Yixian Formation in Liaoning Province in northeast China. The authors dated their find to be about 140 million years old, consistent with the chronology stated above.

In hypothesis, attachment of the Indian Plate to the Eurasian Plate occurred later, about 85 million years ago during the late Cretaceous, – 60 million years after the arrival of Austosunda on the shores of southeast Laurasia. It brought to Eurasia a second biotic cargo. The Indian Plate became the second center of diversification and radiation of angiosperms into the northern hemisphere. Angiosperms soon became the dominant plants upon the land.


Solution to Wallace Line and Other Disjunction Enigmas

The Sunda Islands are politically a part of Indonesia that extends toward New Guinea from the Malay Peninsula to the Moluccas. They include Sumatra, Java, Borneo, Sulawesi, Bali, and Lombok, and numerous small islands. The Sundas have presented a puzzle of partial-to-total separation of fauna and flora of Sibumasu and Indian Plate derivation. On Borneo the native biota is termed “Cathaysian,” brought by the Indian Plate. To the west, across the straits, the native biota of Sumatra is termed “Australasian,” brought by Austosunda from Gondwanaland. 

As noted above, many angiosperm families reached Laurasia for the first time on the Indian Plate, at its later junction with Laurasia, 85 million years ago. By their examples, some families show phytogeographic disjunction (bimodal distribution patterns) that has resulted from the two separate debarkations – 60 million years apart – on the shores of Laurasia. The present Australasian and Cathaysian flora and fauna are the respective manifestations of this disjunctive geography. Their adjacent presences have often been delineated with no little wonder ever since noted by Alfred Russell Wallace and are termed a “Wallace Line” phenomenon.
These biogeophysical accounts of the migrations of Austosunda and the Indian Plate provide solution to other persistent and troublesome phytogeographic puzzles. Two prominent puzzles in plant geography and their solutions are illustrated as follows:

Puzzle 1: the disjunctive distribution patterns of Nothofagaceae (the southern beech) on the one hand and the Fagaceae (the beech and the oak) on the other. All are of the order Fagales.

Solution:  The Nothofagaceae (now with but one genus, Nothofagus) are a Gondwanaland family that are more than 300 million years old. They  consociated with the Proteaceae in a cold temperate environment before and  during the Gondwanaland Ice Age. Like the Proteaceae, Nothofagaceae were present in Australia and Austosunda when their separations from Gondwanaland occurred about 280 million years ago. The present-day geographic distribution of Nothofagaceae is mainly temperate-zone southern hemispheric. There are 20 species in New Caledonia and 9 species in South America, 9000 km. away.

On the other hand, the Fagaceae (oak and beech) are a younger family, which evolved after Austosunda separation. Oak and beech may have originated on the Indian Plate. They were ferried on the Indian Plate; and  from there 85 million years ago, they radiated throughout Laurasia. The Fagaceae are almost entirely northern hemispheric. None is native to Australasia.

Puzzle 2: the disjunctive distribution patterns of the Magnoliaceae and the Annonaceae on the one hand and the Winteraceae and Eupomatiaceae on the other. All are probably of the order Magnoliales. 


Solution:  The Winteraceae and the Eupomatiaceae  are described as among the most ancient of the angiosperm families, more than 300 million years old. Like the Nothofagaceae and the Proteaceae, they were native to Gondwanaland, including Austosunda and Australia, at the time of the separation of Australia from Gondwanaland, 280 million years ago. Some Winteraceae remained in Gondwanaland and in Australia, while others were ferried on the Austosunda to the margins of Laurasia where it made landfall 145 million years ago. They are today almost entirely indigenous to the southern hemisphere (with a small exception of the Winteraceae genus Drimys), and mainly to Australasia. The Winteraceae have been cited by Whitmore (16) as, “the southern counterpart to Magnoliaceae  . . . The [Magnoliaceae] distribution contrasts strongly with that of Winteraceae.” 

On the other hand, the Magnoliaceae and the Annonaceae are younger families which came into existence after the separation of Australia and  Gondwanaland. The Annonaceae, the custard apple family, are the largest family of the order Magnoliales, with 1100 species. The Magnoliaceae are temperate zone plants, and the Annonaceae, mostly tropical. Both the  Magnoliaceae and the Annonaceae may have originated in India.  

The absence of these latter two families from Australia, New Caledonia, Tasmania, and New Zealand strongly indicates that they had not spread throughout Gondwanaland before the separation of Australia at the start of the Permian some 280 million years ago. It also strongly indicates that they were not “on board” Austosunda at its separation from Australia. Hence, they were not ferried to Laurasia 145 million years ago. The Magnoliaceae and the Annonaceae came to Laurasia on board the Indian Plate about 85 million years ago (along with many other families of angiosperms). 

Thus the evidence is pertinent that biogeophysical events – both separations and new attachments of landmasses– differentiate age of origin. For instance, the facts that the beech and the oak of family Fagaceae and the magnolia of family Magnoliaceae are not native to Australasia point to their origins that occurred less than 280 million years ago. 

Moreover, it is noteworthy that, by similar consideration, the ages of other plant families can be determined, as for example, the family Cactaceae. It was suggested by P. Maxwell in 1990 (17), in “The Rhipsalis Puzzle,” that the species Rhipsalis baccifera is among the oldest of extant cacti. A basis for his suggestion is the unique, wide distribution of that species. He notes that R. baccifera is not only endemic to South America and North America but also to Africa, Madagascar, and Sri Lanka; and he quotes Leon Croizat, 1961 (18): “The Rhipsalidinae certainly yield in antiquity to no other cactus.”

But importantly, in dating the origin of  Rhipsalis, no native species is detected here for Australia or New Zealand. DNA and other evidence point to the family Portulaccaceae as ancestral to Cactaceae. Genera of Portulaccaceae also occur today in South America, North America, Africa, Madagascar, and Sri Lanka.

And furthermore, they occur in Australia and New Zealand, as well. Hence, Cactaceae are less than 280 million years old; Portulaccaceae are older than 280 years.


280 Ma: the Biolink Rift of Australia and Austosunda from Gondwanaland

Each of the following families is less than 280 million years old:

     • Cactaceae
     • Cichlidae (fish, in a companion essay)
     • Fagaceae
     • Magnoliaceae
     • Annonaceae
     • Placental mammal families (in a companion essay).

The criterion is that these families have had no Australian history other than recent; and hence, they are assumed to have been absent from Australia and Austosunda at the beginning of the Permian, 280 million years ago.

Thus, this date, 280 Ma, denotes the earliest date of origin for the genera of the above. On the other hand, Portulaccaceae, Nothofagaceae, Winteraceae, and monotreme and marsupial families, all indigenous to Australia today, are premised to have originated more than 280 million years ago.

Australia and Austosunda are estimated here to have broken the biological link (biolink) to Gondwanaland at the beginning of the Permian about 280 million years ago. In evidence are the early warming and attendant rise in sea level in certain regions of western and northern Australia in the midst of the Permo-Carboniferous glaciation.

In evidence, a momentous sea level rise occurred in the geological history of Australia about 280 million years ago, at the beginning of the Permian.       

P. V. Rich et al (1985) (19) wrote: “Sometime in the early Permian, the ice began to shrink [in Australia]. It probably disappeared earlier from Western Australia than it did from the eastern part of the continent . . .  The seas formed deep bays on the Western Australia coastal margin.” 

In evidence, Figure 2, a version of the Tarlings’ 300-250 Ma geographic chart (13), presents a complete isolation by water of Australia from India and, in fact, from the rest of Gondwanaland. The inference is here taken that at a time during the Permo-Carboniferous Ice Age, 300-250 Ma, Australia  became biologically isolated except for reptiles capable of long-distance travel over water.   


                                                                                                 FIGURE 2

Gondwanaland 300 to 250 Million Years Ago

including Ice Field Incursions during the Gondwanaland Ice Age.  Heavy lines make complete separation by water; broken lines mark incomplete separation. Source: Tarling and Tarling, Continental Drift, a Study of the Earth's Moving Surface (1971).

. . . . . . .
. . . . . . .
. . . . . . .
. . . . . . .
Areas of glaciation between 250 million and 300 million years ago with arrows indicating known directions of ice


Areas of tropical coal forests 300
million years ago

In evidence, M.T. Gibbs et al (2002) (20) wrote: “The Permian Period . . .  contains the most recent transformation from a major glaciation to a generally ice-free state . . . Apparently, the deglaciation was relatively rapid, being mainly confined to the Early Permian Sakmarian [285-280 Ma] Stage . . . [A Glossopteris forest cover replaced ice sheets] and the early and ubiquitous Gandawanan sequence, from tillites to coal swamp deposits, indicates a major climate warming  . . . ” 


Melville’s “Gonophyll Theory” of Angiosperm Origin

Ronald Melville (21) reported in 1960: “The primary diversification of the pre-angiosperm stock appears to have taken place . . . during the late Carboniferous or early Permian. Many angiosperm lineages must date back to this period as distinct lines of evolution . . . ” Melville’s “gonophyll theory” – in explanation of the evolution of  the gynoecium (the female sex organs) and the androecium (the male sex organs) of the angiosperm flower  – holds that flowers were first built of modified seed fern leaves bearing either ovules or microsporangia (21).

Although Melville encountered divergence as far back as he was able to look, he continued: “The great disparity between the floral vascular systems of Ranunculaceae and Magnoliaceae  . . .  implies a very ancient separation of these stocks. A number of other lineages must be as old.”  Thus Melville, sought but was unable to establish monophylesis. He was unable to find the primogenitor, the earliest ancestor of the angiosperm.

The consequences of Melville’s gonophyll theory conflict with the generally accepted doctrine of Cretaceous Origin. Hence, Melville’s gonophyll theory finds itself ignored rather than disproved. However, the time of origin of angiosperms as conjectured by Melville is in concord with the time of origin as provided by the hypothesis stated above.      

A chart by P.M. Hurley (22) on the fossil records of Gondwanaland suggests sufficient time for monophylesis. It shows Gangamopteris (the seed fern cited by Melville as a possible ancestor of Proteaceae) to have been present in Eastern Australia during the Devonian 408 to 360 million years ago. 

Devonian Co-evolution of Angiospems and Insects

Insects are here averred to have been essential to the existence of the angiosperm from the Devonian of more than 360 million years ago until this day. In the late Paleozoic, the Proteaceae had likely evolved their nectar-producing glands and nectar reservoirs to attract and reward pollinators. Such nurturing bounty must well attest that insects even then were active in fashioning the flowers of the Proteaceae to their own advantage. An evolutionary reciprocity ensued over many millions of Paleozoic years in which the Proteaceae induced the pollinating fauna to co-evolve with them in order for both to utilize the floral bounty effectively.

The insects were first among faunal pollinators (along with other arthropods). During the late Paleozoic and thereafter – even in later times of stress that could mean the sudden absence of a major cross-pollinating vertebrate – insects might ubiquitously have been available to carry out pollinations. The earliest traces of insects have been found in Devonian rocks. Some 800 species of cockroaches (including winged ones) were extant during the late Carboniferous; and dragonflies of many sizes were then abundant.


Flying beetles, moths, and butterflies came conspicuously into fossil record during the Jurassic, which began 208 million years ago; but some paleontologists note that insects have undergone little change in the last 200  million years. And the wide diversity of present-day pollination-specialized   insects speaks vividly that long before the Jurassic, lively adaptations in their form and function took place. Today, moths and butterflies flit from flower to flower with body-length proboscises, or sucking tubes, drawn out to drink up nectar. Today, pollination - specialized scarabs like the African Trichostetha fascicularis – whose name means “hairy-chest, with hair in tufts” – move the pollen of Proteaceae effectively from flower to flower. 

Assuredly, in the remote pre-Jurassic past, long before 208 million years ago, the predecessors of the earliest known moths, butterflies, and beetles had been induced by entomophilous flowers to evolve bodies specifically suited to perform cross-pollination. By sustaining the phylogenetic invention of the bisexual flower, insects have been essential in fostering the survival of   almost-eternal flowers. They have been the mainstay in pollination of Proteaceae and other angiosperms from the late Devonian to this day.

In serious error, conventional wisdom abruptly places the origin of the angiosperm at about 150 million years ago. Such doctrine is utterly unable to account for the intricate pollination adaptations of angiosperms and insects – whether it be monocot Orchidaceae with bees or dicot Proteaceae with beetles. The observation rife among paleontologists – that insects have undergone little change in the past 200 million years – stands out in stark contradiction of the doctrine of 150-million-year-ago origin of angiosperm. 

There is no room for doubt that the well-orchestrated, inextricably complementary forms and functions of insects and flowering angiosperms, which are manifest in their intricate pollination adaptations, had co-evolved slowly in the Paleozoic past, long before 150 million years ago. 

Carboniferous Coal Deposits Consistent with Angiosperm Presence

Intimation of the Carboniferous presence of the angiosperm and the gymnosperm in Gondwanaland is given in 1996 by R. Osborne and D. H. Tarling, (23) in describing the formation of late Carboniferous coal fields. They point out that in North America and Northern Europe, forests of Lycopsid trees then dominated the swamps and produced most of the biomass of these coal deposits. (Lycopsids today are the clubmosses, quillworts, and spikemosses.)  On the other hand, they remark that in Gondwanaland: “A different kind of coal-forming forest was developing at the same time  . . . This contained entirely different species of vegetation in a cool temperate climate. The southern flora contained annual growth rings, showing large seasonal variations, unlike the tropical forests of the north.”



This essay pleads phyletic gradualism. Charles Darwin wrote (24) “No complex instinct can possibly be produced through natural selection, except by the slow and gradual accumulation of numerous slight yet profitable variations.”

An overarching fundamental thesis is presented here, namely: the angiosperm originated in Gondwanaland during the late Devonian more than 360 million years ago.

This thesis is arrived at mainly by correlation of late-Paleozoic and Mesozoic plate tectonics and climatology with the present-day geographic distributions of living plants. It is further attested to by phylogenetic, entomological, and fossil fuel evidence. The thesis allows sufficient time for origin, monophyletic diversification, and radiation of angiosperms. It describes late-Paleozoic consociated radiation patterns of specific angiosperm and gymnosperm families which exist today. 

Component tenets of the thesis are that the fragmentation of Gondwanaland began in the late Paleozoic and that subsequently (60 million years apart) two huge fragments joined with Laurasia. This sequence of events offers a criterion hitherto unknown for approximating and distinguishing the dates of origin of plant and animal families. 

Hence, this essay accounts for the sudden appearance of a diverse angiosperm flora in the Cretaceous fossil record of the northern hemisphere.  Thereby, it explains the angiosperm diversity observed in the Cretaceous fossil record, without requiring or implying polyphylesis. 

                              Table 1: Description of Five S. Hemisphere Exemplar Plant Families



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