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Wednesday, 6 November 2013

Speciation? II

Response to Section 5.1.2, "Animals"


Summary: Animal hybrids are "rare organisms" because hybridization is not a viable mechanism for animal diversification. Animal hybrids are typically unisexual, where part of the genome is not heritable. For example, in female fish hybrid clones, the parent species is always required to provide the male portion of the genome, meaning a truly new independent species is not formed. Darwinian evolution requires heredity, but this does not entail the origin of anything new that is heritable. Asexual animal hybrids are thus often called "evolutionary dead ends" where they do not produce new diversity and instead "[e]xtant asexual ‘species’ are little more than scattered twigs at the tips of major phylogenetic branches." Moreover, hybrid asexual females are so similar to the parent species that "sexual males of the progenitor species are unable to distinguish hybrid females from those of their own species," implying little morphological change arises in this process. This mechanism is largely "irrelevant" to sexually reproducing animals.



The FAQ suggests animals may also speciate through hybridization, though one primary paper cited by the FAQ (Vrijenhoek 1994) admits that animal hybrids are "rare organisms."
20 Gregory and Mable (2005) concur, observing that "recent polyploidy is far less common in animals than in plants."21 Indeed, there are good reasons why animal hybrids are much rarer than plant hybrids: the problems encountered with animal hybridization appear far more severe than those raised by plant hybridization.

Darwinian evolution can only operate when there is variation, selection, and heredity. Vrijenhoek (1994) discusses hybridogenesis within fish but finds that the hybrid genome is not entirely heritable since, "Paternal B genes are expressed in the hybrids but are not heritable. Only the A’ ‘hemiclonal’ genome is transmitted between generations." This is why hybrid fish like
Poecilia formosa are often called "clones," because the heritable portion of the genome is simply cloned from generation to generation, and males from the parent "species" are always required to maintain the line. Thus, this "species" cannot be maintained without the constant presence of the parent line, meaning the hybrids are not a truly independent species. Since hybridization in vertebrates typically involves asexual clonal reproduction, this poses a problem for those who would cite hybridization as a mechanism of animal evolution.22 As the Monterey Bay Aquarium Research Institute website explains regarding hybrid fish:

"In hybridogenesis, the female mates with a male, forming a female offspring with both the maternal and paternal genomes. When that female offspring produces eggs however, the male genome is discarded."
23

Again, in other words, these hybrids cannot persist without both species present. These hybrids always require the parent species to be present in order for them to originate and persist. The review by Vrijenhoek (1994) explains that such animal hybrids are typically "evolutionary dead ends": 15


"Asexual species are often considered evolutionary dead ends because of their presumed genetic inflexibility. Among vertebrates and insects. only 0.1% to 0.2% of species are strictly asexual. This rarity suggests a ‘mutation/selection-like’ balance. New asexual lineages arise infrequently and go extinct rapidly.
Extant asexual ‘species’ are little more than scattered twigs at the tips of major phylogenetic branches. Except for bdelloid rotifers. asexual lineages have not speciated and diversified into rich asexual clades."24

If anything, animal hybridization would seem to entail collapse, loss, and decrease of pre-existing diversity rather than the generation of new diversity. Indeed, in this case these asexual animal hybrids are not genetically viable in the long term:

"Genetic decay provides a final challenge to the persistence of clones. Muller suggested that mutations would accumulate like a ‘ratchet mechanism" in asexual lineages. Recombination in sexual lineages produces offspring with higher and lower mutational loads than the parents and purifying selection effectively maintains a low load. An asexual population cannot reduce its load below that of the ‘least loaded’ clone. If by chance that clone is lost, the load has increased one step. Excluding back mutations. it cannot be reduced."

25

It’s not clear that this can be a viable long-term mechanism for evolution in vertebrates because this new "species" could never exist on its own in the wild without the parent species constantly available to continually generate it. In the long term, animal hybridization appears to be a byproduct of existing species, not a mechanism for producing new species.

Little more is said in the FAQ regarding hybridization in animals, but it is clear that hybrids result from crossing highly similar species within the same genus, and nothing new is created that is heritable. In fact, regarding

Poecilia formosa, Gregory and Mable (2005) observe that "sexual males of the progenitor species are unable to distinguish hybrid females from those of their own species,"26 implying that little morphological change has occurred. This is not a viable mechanism for evolution in animals, as Dobzhansky recognizes: "Sudden emergence of new species by allopolyploidy is … irrelevant to Drosophila and most bisexual animals."27

Response to Section 5.2 "Speciations in Plant Species not Involving Hybridization or Polyploidy"

Response to Section 5.2.2, "Maize (Zea mays)"


Summary: Corn breeders produced "almost complete reproductive isolation" between two "races" or "varieties" within the same species but a new species was not claimed to have emerged. The partial reproductive isolation results from a premating mechanism—changes in flowering timing—not a large-scale change which might produce fundamentally new types of organisms. The isolation was produced via strong artificial selection; whether this could occur in the wild is not established. 16


According to the paper cited by the FAQ (Paterniani, 1969), humans have bred the corn species
Zea mays into many races over the past 4000 years, and "all races cross readily one with the other giving progenies with normal fertility."28 Through selective breeding that destroyed hybrid crosses, this study sought to achieve reproductive isolation between two species of corn. However, even at the end of the experiment some intercrossing between the varieties still occurred. As Jonathan Wells writes, "Paterniani noted ‘an almost complete reproductive isolation between two maize populations’ but did not claim that a new species had been produced."29

So what exactly generated the reproductive isolation? It turns out that by destroying hybrids, the experimenters were selecting for individuals that flowered at different times and did not produce hybrids. Reproductive isolation was probably achieved by little more than slight changes (a few days) in the timing of flowering in the two varieties:

"Data on days to flowering show that some change occurred, thus indicating that this mechanism is playing a role in the isolation obtained. Both original populations flowered in the same days with identical averages for days-to-tassel and days-to-ear flowering. Cycle IV of the two populations, already showing a great degree of reproductive isolation, have a marked difference in days to flowering. The white flint maize became about 5 days earlier, for tassel and ear, than the corresponding original population. The yellow sweet maize changed less, but in the opposite way; it became, on the average, 2 days later. As a result, the two cycle-IV populations have about 1 week of difference in flowering time. This difference is of sufficient magnitude to explain most of the reproductive isolation obtained."

30

The paper concludes: "The data show that the number of days from planting to flowering was probably the main factor."
31 Thus, what the paper shows is that the plants are flowering at different times and thus don’t have the opportunity to pollinate one-another.32 Other than that, there’s no indication of biological changes. This does not amount to the kind of large-scale change which can produce fundamentally new types of organisms. And it was done by controlled artificial selection; whether this could be achieved in the wild is not established.

Response to Section 5.2.3, "Speciation as a Result of Selection for Tolerance to a Toxin: Yellow Monkey Flower (Mimulus guttatus)"


Summary: Two populations within the same species of a flowering plant "developed partial postmating isolation between some races" where "total postzygotic reproductive isolation between two populations, in the sense that inviable zygotes are formed, can be produced by a comparatively simple genetic mechanism." The cause of the reproductive isolation is thought have "a simple genetic basis" entailing changes in "a single gene." Large-scale differences have not evolved and speciation is only claimed to have been "initiated," not complete.



According to the paper cited by the FAQ (Macnair and Christie, 1983), the yellow monkey flower,
Mimulus guttatus, has "developed partial postmating isolation between some races."33 While offspring can be produced between the two populations, they are inviable, as the paper 17

later reports "total postzygotic reproductive isolation between two populations, in the sense that inviable zygotes are formed, can be produced by a comparatively simple genetic mechanism."
34 The investigators do not fully understand the genetic cause of reproductive isolation but believe it may be linked to pleiotropic effects caused by the gene responsible for copper tolerance/intolerance. They suggest that the isolation has "a simple genetic basis" and is caused by changes to "a single gene."

On the one hand this shows that reproductive isolation may be achieved. On the other hand the fact that offspring can be produced shows large-scale biological differences have not evolved. In fact, the paper suggests that speciation was only "initiated" and not complete.

In this case, we have seen two races within the same plant species are essentially identical apart from one being tolerant to copper and other intolerant. The two races are so similar that they can produce offspring, but those offspring are not viable. Speciation is not complete and significant biological change is not observed. The exact genetic mechanisms which are causing such reproductive isolation are unknown, but they might result from "a simple genetic basis" entailing changes in "a single gene." This does not show significant biological change.


Response to Section 5.3, "The Fruit Fly Literature"

Response to Section 5.3.1, "Drosophila paulistorum"


Summary: This study showed that if you start with "semispecies" within a fruit fly species which are "indistinguishable morphologically," and then subject the strains to artificial breeding experiments, "in none has anything like complete isolation been achieved." Moreover, there is no suggestion that the populations were no longer "indistinguishable morphologically" after the experiments. At best, only a "new race or incipient species" was created. Some authorities have challenged even the partial isolation, claiming the results "may have been due to contamination of cultures by other subspecies."



In this example, the FAQ discusses whether reproductive isolation has been achieved between various strains, or "semispecies," of the fruit fly
Drosophila paulistorum. The paper cited by the FAQ (Dobzhansky and Pavlovsky , 1971) states that in the wild, "The semispecies are indistinguishable morphologically,"35 and "not different enough" to be considered "fully differentiated species." They observed that after a certain amount of time breeding in captivity, crosses between two particular strains only produced sterile males. (Female hybrid crosses were apparently still fertile.) The investigators claim that contamination was "ruled out" but Jonathan Wells notes that those claims may be incorrect, for "Coyne and Orr wrote in 2004, however, that Dobzhansky and Pavlovsky’s result ‘may have been due to contamination of cultures by other subspecies.’"36

After conducting artificial breeding experiments where hybrid crosses between the strains were destroyed, the authors produced some assortative mating. (This process does not mimic natural conditions.) The authors admit that the proportion of hybrids merely "decreased" and "in none has anything like complete isolation been achieved."
37 Jonathan Wells thus notes that in this example: "Dobzhansky and Pavlovsky reported only a ‘new race or incipient species,’ not a new 18

species."
38 Another paper in the FAQ (Halliburton and Gall, 1981) lists this study among various studies where "none has succeeded in establishing complete sexual isolation."39 In any case, speciation is not claimed to have occurred, and there is no suggestion whatsoever that the semispecies are no longer "indistinguishable morphologically." In fact, after reviewing this example, Dobzhansky concludes that sometimes "reproductive isolation and speciation precede differential adaptedness,"40 suggesting the populations had not diverged.

Response to Section 5.3.2, "Disruptive Selection on Drosophila melanogaster"


Summary: Artificial selection for the number of chaetae (hairs) on a population of fruit flies produced partial, though not complete, reproductive isolation. The extent of morphological variation is small-scale changes in the number of chaetae. Later attempts to reproduce these results were unsuccessful. The authors explicitly state that natural speciation has not been demonstrated. This experiment does not show complete reproductive isolation, speciation, or significant morphological change.



In this example, the paper cited by the FAQ (Thoday and Gibson, 1962) reports experiments with a small population of wild fruit flies (
Drosophila melanogaster), which over a series of generations selected those with both the highest and lowest numbers of chaetae, or hairs. Through artificial selection over the course of successive generations, they were able to select for flies with more, or less chaetae than in the original population. This is the extent of the variation bred by this experiment.

Even after multiple generations, the flies with high or low numbers of chaetae could still hybridize. However, hybrids did become less common as the experiment progressed, although the reason for this was not known. The experimenters proposed that perhaps it "arises from mating preferences or from an inability of hybrid flies to compete as larvae."
41 The ability for the high and low chaetae populations to hybridize, although rare, was not impossible. Reproductive isolation was thus not complete. Additionally, since artificial (rather than natural) selection was used to create and maintain the two populations, the authors warn:

"We do not, however, wish it to be thought that we regard this as a demonstration that sympatric speciation occurs in Nature, for such a conclusion cannot be drawn from the results of laboratory selection experiments."
42

At best, this experiment shows there is pre-existing variation among fruit flies for numbers of chaetae, and artificial selection for this trait in a non-natural laboratory setting can result in partial reproductive isolation. But changes in the numbers of hairs was all that was produced: it does not show anything close to large-scale evolutionary change.

Thoday and Gibson (1962) open their paper by admitting that "the key demonstration that a single wildtype population can be converted by selection into two populations that are mutually isolated in the conditions in which they have to maintain themselves has not hitherto been made."

43 But Jonathan Wells observes, "Not only did Thoday and Gibson not claim to have produced a new species, but also other laboratories were unable to replicate their results."44 The 19

FAQ likewise admits, "In the decade or so following this, eighteen labs attempted unsuccessfully to reproduce these results." The summary of Halliburton and Gall (1981) is striking:

"Several attempts to repeat these results have failed (e.g., Scharloo

et al., 1967a; Chabora, 1968; Barker and Cummins, 1969). Similar experiments, selecting for other quantitative characters or using other organisms, have usually failed to produce assortative mating (Scharloo, 1964; Robertson, 1966; Scharloo et al., 1967b; Grant and Mettler, 1969; Bos and Scharloo, 1973), but a few have succeeded (Coyne and Grant, 1972; Soans et al., 1974)."45

As far as this example goes, it would seem this "key demonstration" of speciation has not been made.


Response to Section 5.3.3, "Selection on Courtship Behavior in Drosophila melanogaster"


Summary: This experiment sought to induce changes in the mating preferences of two strains of fruit flies. Only "partial" reproductive isolation was achieved, and the extent of change observed was small changes in courtship initiation behaviors (e.g., licking and vibrations). The two strains were "similar" before the experiments, and apart from slight changes in mating behaviors, remained very similar after the experiments.



This experiment took two pre-existing strains of fruit flies from within the same species—
Drosophila melanogaster—and sought to determine whether changes in mating preferences could be induced. This included artificially killing hybrids between the strains (a process that does not necessarily mimic nature). Incomplete reproductive isolation was observed, which one paper cited by the FAQ (Knight et al. 1956) called only "[p]artial sexual isolation."46 Another paper in the FAQ (Halliburton and Gall, 1981) lists this study among various studies where "none has succeeded in establishing complete sexual isolation."47 The most biological change that this example documented was small-scale behavioral differences pertaining to courtship, specifically changes in the amount of "licking" that males do to females to initiate mating. One paper cited by the FAQ (Crossley, 1974) showed just how unimpressive the sort of change observed in this experiment was:

"Quantitative analysis of male and female behavior revealed the underlying causes of changed mating preferences and faster mating. In the LS experiment male courtship became more stimulating because percentage licking of both males and percentage licking plus vibration of males increased."
48

Thus, all that was observed were changes in the courtship initiation behaviors (licking and vibrations) between the strains. This is small-scale change. The two strains were "similar" before the experiments, and apart from slight changes in mating behaviors, remained very similar after the experiments.

These experiments were conducted in the laboratory, but Crossley (1974) observed why laboratory experiments do not match natural conditions: "One difficulty in relating these results to selection against hybrids in nature is that in the laboratory, selection against hybrids was total 20


but in the wild some hybrids would survive to breed in spite of their disadvantages compared with pure bred offspring."
49

Response to Section 5.3.4, "Sexual Isolation as a Byproduct of Adaptation to Environmental Conditions in Drosophila melanogaster"


Summary: This fruit fly study found partial reproductive isolation after selection experiments on fruit flies. No significant morphological change was reported, and any reproductive isolation which did exist stemmed from premating factors. This paper thus serves as a good example of how speciation need not entail significant morphological or genetic change.



In this experiment, the investigators changed temperature and humidity conditions for populations of fruit flies (
Drosophila melanogaster) in the laboratory. Originally the populations came from the same genetic stock, but after about 5 years of artificially exposing them to different environmental conditions, the experiment found that some reproductive isolation was established. According to the paper cited by the FAQ (Kilias et al., 1980), "The highest isolation index detected was 0.388 ± 0.108,"50 meaning reproductive isolation was far from complete.

But was any significant morphological change found? None was reported, and in fact Kilias
et al. (1980) said the reproductive isolation resulted simply from pre-mating factors:

"Since, females from either population mate equally, the reproductive isolation detected in the present investigation seems to be due to changes in behavior (different mating preferences or discrimination) of our populations. … In the present study we failed to detect significant postmating isolation."
51

The change that was observed pertained to the sexual isolation and changes in ovipositional rhythms, and thus they observed that "speciation" (meaning mere reproductive isolation) "may occur with relatively little genetic change in structural genes."
52 This led them to the final conclusion that genetic divergence between populations often occurs after "speciation" (again, mere reproductive isolation) because such divergence was not observed here: "The genetic distance observed between species probably results from post-speciational divergence."53 This paper thus serves as a good example of how speciation need not entail significant morphological or genetic change.

Response to Section 5.3.5, "Sympatric Speciation in Drosophila melanogaster"


Summary: After two populations of fruit flies were selected for various food-finding behaviors, incomplete reproductive isolation was observed. The populations could still produce "fertile offspring" and speciation was only claimed to be "incipient. No significant morphological change arose.



This is another study where partial reproductive isolation was established between populations of
Drosophila melanogaster, and the differences between the populations were minor, and of ambiguous importance. 21

The experimenters forced fruit fly pupae to navigate a maze to find food where they might choose to go towards the light/dark, up/down, or choose between two different scents. Flies which made opposite choices were then separated, allowed to breed, and then subsequent generations were selected according to which flies made the same choices. When the fly populations were then allowed to mix, partial reproductive isolation was achieved.

Complete reproductive isolation was not found. As one of the papers cite by the FAQ (Rice and Salt, 1988) reports:

"It might be argued that incipient speciation has not occurred in this experiment for two reasons. First, trace amounts of gene flow occurred between the population using habitats SE and 4L, since a small fraction of flies switched habitats between generations. … Second, forced matings between the two populations produced fertile offspring in the F1 and F2, and thus reproductive isolation was mediated only by habitat-preference behavior. … [I]rreversible reproductive isolation did not occur in this experiment."

54

The fact that gene flow between the populations occurred, and that they could be forced to produce fertile offspring, shows how similar the strains remained. They claim that any speciation is merely "incipient."

But what evolved? At most they might have selected for preferences for when seeking food. As they conclude: "In these experiments, the only barriers to gene flow were gaps that gradually developed in the distribution of spatiotemporal habitat preference."

55 It’s important to realize that again, we have seen no significant morphological change. As Jonathan Wells observes, "Within thirty generations the flies had sorted themselves into two populations that did not interbreed, but Rice and Salt claimed only ‘incipient speciation that we believe to have occurred.’"56

Response to Section 5.3.6, "Isolation Produced as an Incidental Effect of Selection on several Drosophila species"


Summary: Three fruit fly studies were reported: they showed "slight" or "incipient" or "not complete" sexual reproductive isolation, but none showed complete reproductive isolation or speciation. None showed significant morphological change.



In this example, the FAQ discusses a paper (del Solar, 1966) which reported experiments that artificially selected "positively and negatively geotactic" and "positively and negatively phototactic" strains of both
Drosophila melanogaster and Drosophila pseudoobscura.57 The paper reports that this produced what was called "slight" sexual isolation, or "incipient reproductive isolation," due to "changes in sexual behavior."58 The paper thus reports that complete reproductive isolation was not found:

"Whether selection for geotaxis and phototaxis always and necessarily produces a change in the sexual behavior, and whether continued selection may carry the sexual divergence
anywhere near complete isolation, can only be decided by further experiments."59 22

Not only was "anywhere near complete isolation" not achieved, but significant biological change was also not achieved. As the paper reports: "The geotactically and phototactically positive and negative strains appear to be
indistinguishable in external morphology."60

Another example discussed by the FAQ in this section pertains to Dodd (1989) which reported experiments on populations of the fruit fly
Drosophila pseudoobscura. Four populations were given a starch-based medium, and the other four were given a maltose-based medium. The paper reported that individuals raised on the starch medium preferred to mate with other starch-fed fruit flies; likewise flies fed maltose preferred to mate with other maltose-fed individuals. Interestingly, these traits arose independently in each of the four populations in each medium.

Because the experiments controlled for food source rather than mating behavior, they concluded that the sexual isolation was "a pleiotropic by-product of the adaption of the populations to the two media" but "The mechanism of the isolation in this system is as yet unknown."
61 Indeed, it was not that the two populations were incapable of interbreeding or never interbred, it was just that they did so less than would be expected under normal random mating. Another paper cited by the FAQ (Schluter and Nagel, 1995) described the findings by stating that only "some premating isolation evolved," and "Reproductive isolation between divergent lines was not complete."62 Speciation was also not said to have occurred.

Aside from mating and food preferences, there were no claims of biological change between the populations. Again, we see that not only has reproductive isolation not been demonstrated, but significant biological change did not evolve.

Despite the aforementioned underwhelming results, the FAQ then discusses another paper which it says reported "Less dramatic results." According to the paper cited by the FAQ (de Oliveira and Cordeiro, 1980), different populations of

Drosophila willistoni were given food at different pH levels. Like the other studies in this section, some individuals preferred to mate with other individuals fed food at the same pH. But the different populations were still capable of interbreeding. When the offspring were fed alkaline food, "their hybrids are not less fit."63 However, the paper reported that "on the acid substrate the hybrids are inferior to their parents which are adapted to this food."64 The paper thus only claimed to find "incipient isolation,"65 not complete reproductive isolation.

As for the degree of morphological change, aside from a preference for a certain pH level in food, no significant biological change was reported. In fact, the paper notes that among three long-standing natural races of
D. willistoni, "These flies are morphologically indistinguishable." This study certainly did not change that observation: Once again we have not seen complete reproductive isolation, nor have we seen significant biological change.

Response to Section 5.3.7, "Selection for Reinforcement in Drosophila melanogaster"


Summary: Again, fruit fly experiments found only "partial" reproductive isolation and did not report significant biological change. One paper boasted that "[t]he evidence here presented shows … that natural selection can act to strengthen isolation." But since the ‘destroy the hybrid’ experiments simulated processes that would never occur in nature—23


the artificial destruction of all hybrid flies for no biological reason other than experimental curiosity—it obviously confused natural selection with artificial selection.



In this section, the FAQ acknowledges that Rice and Hostert (1993) do not find evidence for the reinforcement model of speciation, where, according to the paper, "the physical barrier breaks down before complete reproductive isolation has evolved in allopatry" but yet "matings between previously separated subpopulations are presumed to produce low-fitness hybrid offspring, and this selects for positive assortative mating."
66 If hybrids cannot survive and reproduce, then arguably the two populations are reproductively isolated.

The FAQ then discusses two older papers which purportedly support the reinforcement model. Keep in mind again that the most important question here is not whether two populations can fail to produce viable hybrids, but whether the two populations show some non-trivial degree of evolutionary change.

The first study cited by the FAQ (Ehrman, 1971) took two strains of

Drosophila melanogaster and sought to test for sexual isolation. The paper never claims that significant morphological change evolved, but it does note that this experiment had results similar to Knight et al. 1956, a paper which only found "[p]artial sexual isolation," and never claimed significant biological change arose. Likewise, Ehrman (1971) reports that after breeding experiments, in males there is only "some sexual isolation," and the author thus hopes that "the degree of reproductive isolation evolved to be enhanced by the passage of time."67 The paper cites no significant changes in the fly populations after the experiments. The experiment did not find cross-mating is impossible, placing limits on the degree of change which arose. Not only was complete reproductive isolation not achieved, but there is no report whatsoever that significant biological change emerged. Another paper in the FAQ (Halliburton and Gall, 1981) lists this study among various studies where "none has succeeded in establishing complete sexual isolation."68

In a similar study, another paper cited by the FAQ (Koopman, 1950) mixed two similar species of
Drosophila, D. pseudoobscura and D. persimilis in an attempt to induce reproductive isolation in the lab. Complete reproductive isolation was not established.

Normally one would think that if the two populations are already classified as members of different species then perhaps they are already completely reproductively isolated; but this is not the case here, as the two species are "closely related" and in fact
D. persimilis was "formerly known as D. pseudoobscura, race B."69 Complete reproductive isolation between the two species does not exist in the lab: the two groups can form hybrids, and "Hybrids seem to have the same viability as the pure species," although hybrid males are sterile and females, when backcrossed with parent species, tend to have eggs with "poor viability."70 While they form hybrids in the lab, however, "in nature, not a single hybrid has been found, even from localities where both species occur together."71

To help investigate whether complete reproductive isolation could emerge, the experimenters used a tactic that would not be present in nature: they artificially killed any hybrids. As the paper stated: 24


"The experiments herein described were made in order to determine whether, in artificial populations consisting of the two closely related species … an increase in the reproductive isolating mechanisms could be detected if in each generation the hybrids between the two species were systematically eliminated."
72

The paper concluded that "[t]he evidence here presented shows … that natural selection can act to strengthen isolation between species."
73 But was it "natural selection" or artificial selection? The paper shows astonishment at how "in a surprisingly short time," increased reproductive isolation was established. But they really should not be so surprised since they recognize that "This change, of course, was aided by the practice of removing the hybrids entirely each generation, in this way simulating complete hybrid inviability."74 Thus, natural selection was not at work; rather artificial selection caused these changes.

In any case, complete reproductive isolation did not arise as hybrids still formed, albeit at a "low level." So the experiments started off with two similar populations of flies that have partial reproductive isolation, and they ended with two highly similar populations of flies that have "partial"
75 (though slightly more) reproductive isolation. The paper makes no report of significant morphological differences between the populations at the beginning of the experiment, and the end, so this experiment once again shows that (1) complete reproductive isolation was not achieved, and (2) significant biological change did not arise.

Response to Section 5.3.8, "Tests of the Founder-flush Speciation Hypothesis Using Drosophila"


Summary: Three papers testing the founder-flush model of speciation using fruit flies failed to produce complete reproductive isolation. Reproductive isolation was called "partial" and / or "weak," and no significant morphological change was reported.



The first paper cited by the FAQ in this section (Powell, 1978) investigated a hypothetical "founder-flush" mechanism of speciation where a small number of individuals found a new population, which then goes through various cycles of expansion in population size ("flush"), followed by a "crash," where "[a]t each crash the bottleneck population is small and genetic drift is strong."
76 After the crash, another small group of individuals found the new population, and the cycle repeats. Powell (1978) attempted to simulate this process for strains of fruit flies within a population of Drosophila pseudoobscura.

The paper reported that "neither isolation nor inbreeding by themselves lead to reproductive isolation. Only among populations which were inbred (four founder events) and allowed to flush did reproductive isolation evolve."
77 When reproductive isolation did evolve, it was called "partial"; at one point the author claimed only "some degree of reproductive isolation."78 He claimed to only have observed the "first stages of speciation,"79 not complete speciation.

What is also significant is the type of reproductive isolation that evolved. Powell (1978) reports that "no post-mating factors were detected,"
80 indicating that when crossbreeding between the populations did occur in the experiment, it produced viable and fertile offspring. This implies that significant biological change between the populations did not arise over the course of the 25

experiments. The only type of reproductive isolation that was observed was "pre-mating (ethological) isolation,"
81 where behavioral factors reduce cross-mating.

Finally, the paper observes the extreme degree of repeated flushes and crashes simulated in the experiment would require "rather special circumstances" not necessarily common in nature. Either way, once again neither complete reproductive isolation nor significant biological change was observed.

Another paper cited by the FAQ (Dodd and Powell, 1985) repeated this type of experiment and found very similar results to Powell (1978). Reproductive isolation was called "significant" but was far from complete. The overall finding was that: "in general, it appears that some weak ethological isolation exists."

82

Also like Powell (1978), all reproductive isolation that did arise was due to behavioral (premating) mechanisms and "no post-mating isolation could be detected."
83 This implies that significant morphological or genetic change did not arise between the separated populations over the course of the experiment, because fertile and viable offspring could be produced. The paper thus notes that these results counter the common evolutionary presumption that speciation occurs because populations diverge biologically:

"Many scenarios for the formation of new species envision post-mating isolation factors to evolve before pre-mating isolation evolves (e.g., Dobzhansky, 1940). Pre-mating barriers are thought to be secondary, reinforcing mechanisms of isolation. Here they appear to be primary; that is, they have evolved in the apparent absence of post-mating isolation."
84

Like the others, this experiment did not report that significant biological change evolved.

A final paper cited in this section of the FAQ (Ringo

et al., 1985) used populations of Drosophila simulans test the founder-flush model (which fosters genetic drift) against the classical model of speciation, where certain traits undergo selection, gradually leading to a new species. The lines experiencing selection were artificially selected for various "arbitrary traits."85

While some "partial reproductive isolation" in the various lines did arise, they found it was "much weaker than that typically found between sibling species of
Drosophila."86 They thus hope that:

"More-complete reproductive isolation might be established by the same forces at work over a longer time span, perhaps reinforced by direct selection for premating barriers to gene flow."
87

They thus lament that "A large gap lies between the degree of isolation between any experimental populations and the degree of isolation observed between species."
88

As far as premating isolation goes, the paper reports that "Weak sexual isolation was observed between BASE and the drift lines, for the experiment as a whole."
89 However, the paper did find 26

some postmating isolation. Interestingly, "reproductive isolation was stronger in drift lines than in selection lines" and "postmating isolation increased over time in drift lines but not in selection lines" and overall "there was only a 5% reduction in hybrid fitness." This implies that even artificial selection for certain traits did not produce sufficient biological change to prevent viable and fertile hybrids from forming between the selected lines and the original base population. Significant biological change is not reported.

Even the FAQ admits regarding this study that "only weak isolation was found and that there was little difference between the effects of natural selection and the effects of genetic drift." In other words, even when certain traits are selected for in the laboratory, only weak isolation arises viable and fertile offspring between crossed lines can still be produced. Once again, incomplete reproductive isolation (which the FAQ admits is "weak") and only very limited small-scale change observed, even when there is artificial selection for many traits.

Finally, it’s worth noting that Ringo

et al. (1985) and Powell (1978) "have been criticized (Charlesworth et al. 1982) because the base populations were derived from geographically diverse stocks, ‘so it is not clear whether their results are representative of what might happen in a natural population.’"90 Other critics (Meffert and Bryant, 1991) observed that in Ringo et al.’s experiment, "Because the populations contributing to the base population exhibited differences in mating activity that were presumably genetic, the experimental protocol created an artificially high genetic variance for traits affecting mating behavior and exaggerated the divergence among experimental lines."91

The response from the authors to such criticisms is that they wanted to study "the extent and the speed of establishment of reproductive isolation under optimal conditions; that is, we hoped to maximize reproductive isolation among lines by maximizing genetic variation"
92 in the initial population. They did this because when more realistic natural conditions are modeled in experiments, the results might have "[n]egative."93 In other words, even when they gave speciation mechanisms their best shot—better than would likely exist in nature—complete reproductive isolation was not achieved and significant biological change was not observed.

In any case, Meffert and Bryant (1991) observe that due to weaknesses in Ringo
et al. 1985:

"Hence the critical issue in the founder-flush theory of speciation has not been addressed: can bottlenecks in a natural population cause permanent alteration of courtship behavior in founder lines that would lead to premating isolation."
94

Would Meffert and Bryant thus argue that the papers cited in this section of the FAQ don’t even establish what the FAQ claims they do?

Speciation?

The accompanying is a reproduction of the discovery institute's article





Specious Speciation:

The Myth of Observed Large-Scale

Evolutionary Change


A Response to TalkOrigins’

"Observed Instances of Speciation" FAQ


(Updated Sept. 9, 2013)


by Casey Luskin


Casey Luskin is Research Coordinator at the Center for Science and Culture at Discovery Institute. He holds both a B.S. and M.S. in Earth Sciences from the University of California, San Diego and has conducted geological research at the Scripps Institution for Oceanography. He is also a licensed attorney.



PART I: EXECUTIVE SUMMARY

The TalkOrigins Speciation FAQ, titled "Observed Instances of Speciation"
1 (herein "FAQ"), claims it "discusses several instances where speciation has been observed." For years, this FAQ has been cited by pro-Darwin internet debaters as allegedly demonstrating that neo-Darwinian evolution is capable of producing significant biological change. However, an analysis of the technical literature regarding many of the examples discussed in the FAQ2 reveals that such claims are clearly incorrect. This assessment finds:


NOT ONE of the examples demonstrates the origin of large-scale biological change.

The vast majority of the examples do NOT even show the production of new species, where a "species" is defined according to the standard definition of a "reproductively isolated population." Only one single example shows the production of a new species of plants via hybridization and polyploidy, but this example does not entail significant biological change.

 Only one of the examples purports to document the production of a reproductively isolated population of animals—
however this example is overturned by a later study not mentioned in the FAQ.

2


 

Thus, not a single bona fide example of speciation in animals—e.g., the establishment of a completely reproductively isolated population—is given in the FAQ.

I should note from the outset that my purpose is not to deny that speciation can occur in nature, especially when speciation is defined merely as a reproductively isolated population. When trying to assess the creative power of the Darwinian mechanism, that definition is trivial. Rather, my purpose is to test the FAQ’s claims. In that regard, if the FAQ is correct that "Many researchers feel that there are already ample reports [of speciation] in the literature," then an analysis of the literature cited in the FAQ suggests those researchers are wrong.


While most of the FAQ’s discussions of the papers it cites are reasonably accurate, these papers amount to citation bluffs if one is claiming to "discus[s] several instances where speciation has been observed." People who believe this FAQ demonstrates that Darwinian processes can produce large-scale biological change have been badly misled. The examples in the FAQ are ultimately used to make inaccurate claims, and the FAQ’s title, "Observed Instances of Speciation," is unwarranted.

PART II: "SPECIATION?" ITS ALL IN THE DEFINITION.

The TalkOrigins "Observed Instances of Speciation" FAQ, claims to discuss "several instances where speciation has been observed." The most important question is whether these examples show significant biological change has occurred. But this analysis shows:


(1) As a primary finding, none of the examples demonstrate that Darwinian evolution is capable of causing large-scale evolutionary change.

(2) As a secondary finding, the vast majority of the examples do not even meet the standard definition of "speciation."

To understand why both (1) and (2) are the case, we first need to understand the implications of how evolutionary biologists typically define "species."

Evolutionary biologists typically define "species" as a reproductively isolated population of individuals. For example, the FAQ quotes the great neo-Darwinian evolutionary biologist Ernst Mayr defining a species as "groups of actually or potentially interbreeding natural populations which are reproductively isolated from other such groups." This classical definition is called the biological species concept. Under this standard definition, speciation entails the origin of such a reproductively isolated population. But does it entail anything else?

Not necessarily. Such definitions say nothing about the degree of morphological, behavioral, or genetic change that has evolved. Thus,

such a definition of "species" does not necessarily imply that significant biological change has taken place between the two populations. In many cases, two populations may be termed different "species" under the biological species concept, but yet the differences between the populations are small-scale and trivial. Indeed: 3

 

One of the papers cited by the FAQ (Dodd, 1989) clearly states that speciation is reduced to mere reproductive isolation, "According to the biological species concept, speciation is basically a problem of reproductive isolation."3 Another paper (Schluter and Nagel, 1995) cited by the FAQ notes that under this definition species are "defined by the criterion of reproductive isolation rather than morphological criteria."4

Yet even the notable evolutionary biologist Theodosius Dobzhansky (1972) admits that under this view, "speciation may occur without rearrangement of the genetic materials in the chromosomes" and "Reproductive isolation evidently can arise with little or no morphological differentiation."5

Putting these quotes, claims, and definitions together, papers cited by the FAQ admit that under the biological species concept, "speciation" does not require any morphological change.

As noted, the FAQ is often cited to explicitly or implicitly claim that Darwinian evolution is capable of producing significant biological change. But the FAQ’s definition of "speciation" seems contrived when used to demonstrate the grander claims of Darwinian evolution that fundamentally new biological structures, body plans, and higher taxa can evolve. Even if we do find reproductively isolated populations that document "speciation," that might provide virtually no evidence that Darwinian processes can produce new complex biological features or large-scale change.
Indeed, the primary finding of this analysis is that the examples in the FAQ do not report the kind of change which shows Darwinian processes can produce fundamentally new types of organisms, new complex biological structures, or higher taxa.

What is more, the vast majority of the examples in the FAQ don’t even document "speciation" under the biological species concept. One paper cited by the FAQ (Rice and Hostert, 1993) notes that "Once pre- and/or postzygotic isolation is complete, speciation has occurred."
6 But in the vast majority of the instances cited by the FAQ, pre- or postzygotic isolation was not complete, and thus speciation did not occur. Thus, a secondary finding of this report is that only one single paper analyzed in the entire FAQ actually reported complete reproductive isolation, and thus "speciation" under the biological species concept.

This raises an irony: the title of the FAQ is "Observed Instances of Speciation," yet the vast majority of the examples analyzed show that complete reproductive isolation was not achieved. Thus, the FAQ is overselling the evidence, not just for significant morphological change, but also for true speciation (e.g., complete reproductive isolation).
If these are some of the best examples for "speciation" that evolutionists can muster, then the evidence for Darwinian evolution must be meager indeed.

Before discussing this analysis of the FAQ, I must again reiterate that my purpose has never been to deny that speciation can occur in nature, especially when speciation is defined by the trivial definition of a mere reproductively isolated population. Rather, my purpose is to test the FAQ’s claims. 4


PART III: ANALYZING THE FAQ

Section 5.0 of the FAQ purports to provide "examples of observations of speciation," making this the section that needs to be studied to confirm if the evidence backs the FAQ’s claims.

Many of the references cited by the FAQ are old and were difficult to obtain. I downloaded whatever papers I could find online from my local university library and analyzed those examples.


A. Summary of Findings


The following table summarizes the findings of this review for the examples analyzed:
FAQ Section:
Summary:

5.1.1.3, "Tragopogon"

Two plant species could hybridize into populations showing small-scale changes compared to the "parent species"—the greatest of which is color changes of the kind well known within plants. Since hybrids are "extremely sterile," it does not seem that speciation has occurred.

5.1.1.5 "Hemp Nettle (Galeopsis tetrahit)"

Two highly similar species of flowering plants within the same genus were crossed in the laboratory to produce a polyploid plant virtually identical to a known species in nature. This confirms the long-known fact that plants can hybridize to form new polyploid forms. But speciation by polyploidy does not produce new morphological characteristics, and the tetraploid daughter species showed only small-scale changes—the greatest of which is color changes of the kind well known within plants—from the parent species. Speciation by hybridization and polyploidy is not a viable mechanism for the vast majority of evolution because: (1) it occurs only within flowering plants, (2) it does not produce new morphological characteristics, and (3) polyploid hybrids cannot arise without pre-existing parent species, meaning it entails a collapse—not gain—of existing diversity.

5.1.1.8, "Maidenhair Fern (Adiantum pedatum)"

An abnormal sporophyte ultimately led to tetraploid, rather than normal diploid sporophytes in a species of fern. No morphological change was reported and tetraploid sporophytes were "less vigorous" than normal. At best this evidence only "provided the opportunity to characterize the first step of one possible route to polyploid formation" as speciation was merely "incipient." Complete speciation was not observed and large-scale morphological change was not observed.

5.1.2, "Animals"

While hybrids can occasionally occur between very closely related animal species, generally speaking animal hybrids are "rare organisms" because hybridization is not a viable

Monday, 4 November 2013

On Jehovah's Firstborn.

This article is reproduced from Examiningthetrinity.blogspot.com


Some trinitarians insist that the literal "firstborn of all creation" describing Jesus at Col. 1:15 really means "the pre-eminent one over all creation."

"Firstborn" (prototokos) is an integral part of the Father = Source idiom so common in the Bible and is closely associated with "beginning" (arkhe) and "only-begotten" (monogenes).

Jehovah (who is the Father alone) is never referred to as "first-born" in any sense. Any person, animal, or thing who is ever called "first-born" in the Bible is always a part of Jehovah's creation and is literally the very first one born in a family with other children following (or figuratively the very first one in a line of others who share the quality or attribute under consideration).

If we are to understand the literal "first-born" to sometimes figuratively mean "pre-eminent" (as some trinitarians attempt to do - primarily to avoid the literal meaning of Col. 1:15: Jesus "the first-born of all creation" - see RSV), certainly it should, occasionally at least, be applied to the truly pre-eminent one of all, The Most High. But this never happens in the entire Bible!

The Most High (Pre-eminent One) is always the Father (Jehovah only) alone- see Mark 5:7; Luke 1:32; Luke 8:28; the parallel accounts of Luke 6:30-35 and Matt. 5:42-45; Ps. 83:18; and Ps. 7:17. But He is never called "firstborn" (or "only-begotten")!

The source or originator of all creation is the Father as the very title itself, "Father," tells us. Prototokos or "firstborn" is nearly always used, as the word literally tells us, to mean one who is the beginning of his Father's creative (or procreative) power. And, in fact, arkhe (obviously meaning "beginning") is often used in conjunction with prototokos. For example, the Greek Septuagint says at Gen. 49:3, "Ruben, thou art my first-born [prototokos] ... and the first [arkhe - 'beginning'] of my children." - Septuagint Version.

Even if prototokos could be used to mean "pre-eminent one," it's obvious that the terms "Father" (for the person who is the source and the superior of Jesus), "Son" (the person created by his Father, and in a subordinate, intermediary position to his Father), "only-begotten," "first-born," and "beginning of God's creation" all combine (with the most common understanding of those words by those who spoke and read them at that time) to only one possible conclusion: there was a time when only the Father ("the source") existed. Then, at some point, the Father brought another person into existence and this person was the first production of his creative powers, his "firstborn and the beginning of his creation."

Let's look at Col. 1:15,18:

"He [Christ] is the image of the unseen God and the first-born of all creation,.... As he is the beginning [arkhe], he was first to be born [first-born] from the dead, so that he should be first [proteuo] in every way" - The Jerusalem Bible.

Prototokos, used twice in this scripture, literally means "born first" - see Young's Analytical Concordance - or Strong's Exhaustive Concordance. The New Testament in the King James Version and most other trinitarian translations use this meaning throughout. Here are all the instances of prototokos in the NT: Matt. 1:25 (King James only); Luke 2:7; Ro. 8:29; Col. 1:15; Col. 1:18; Heb. 1:5, 6; Heb. 11:28; Heb. 12:23; Rev. 1:5 (compare Col. 1:18). None of them clearly means "pre-eminent" (although you might be able to "interpret" a few of them as either "first-born" or "pre-eminent" if you tried).

Not only do all of these scriptures that use prototokos have either the certain or the most probable meaning of "first-born," but we rarely (if ever) see any Bible translate them as anything but "first-born" or its literal equivalent except at Col. 1:15-18 where the actual meaning would disprove a trinity concept! A few trinitarian translations force an improper interpretation for prototokos at this scripture only (e.g. NIV, NEB).

It is true that being first-born in a family was strongly connected with pre-eminence. The one born first was usually supposed to be the one to receive the birthright and pre-eminence within that family.

But notice the blessings given by Jacob at Gen. 49:3, 8-12, 22-26. The blessings given to Judah and Joseph identify them as the true "pre-eminent ones" of his sons. Reuben, the literal first-born, lost pre-eminence even though he continued to be known as the "first-born" (prototokos in the Septuagint) in the family of Jacob and the "beginning" (arkhe) of Jacob's family - Gen. 49:3, 4; 1 Chronicles 5:1-3 – RSV.

Be careful not to confuse the rights usually given to the first-born with the person of the first-born. The one actually born first (or first in time in any figurative sense) was known as the "first-born." In literal families this first-born was supposed to receive pre-eminence in that family upon the death of his father because of his being born first (in time).

"The first-born son's privileges and responsibilities are known as his `birthright' (bekorah)." - New Bible Dictionary, 1982, p. 378.

At times, however, a first-born would lose his rights (and pre-eminence over the other sons), and they would be given to another son. Even though this person had lost his birthright (and pre-eminence among his brothers), he was still the first-born! - Examine 1 Chronicles 5:1-3 in most Bible translations (e.g., Young's Literal Translation of the Holy Bible, KJV, ASV, RSV, NIV, NAB, JB, etc.) For example, even though Esau lost his birthright to Jacob, he still remained forever Isaac's firstborn.[14]

Yes, the use of the word prototokos in the Bible always means one who has come into existence first in time - before all the rest of his "brothers" - the beginning (arkhe) of his father's creative (or procreative) works. - see pp. 77-88 in Dr. Jason BeDuhn's Truth in Translation, University Press of America, 2003.

Some trinitarians, however, still insist that the Biblical use of the Greek prototokos can, sometimes, mean "pre-eminent" because they dare not admit the obvious, true, literal meaning of Col. 1:15. Their "evidence" for an interpretation of "pre-eminent" for this word boils down to only 7 examples. Five from the Old Testament (Ps. 89:27; Ex. 4:22; Jer. 31:9; Deut. 21:16, 17; and, rarely, Job 18:13) and, sometimes, an appeal to Ro. 8:29 and Col. 1:18 in the New Testament. - See Walter Martin's The Kingdom of the Cults, p. 123.

These 7 examples, then, must be the very best "evidence" possible since there are well over 100 other examples of prototokos found in the Bible, the vast majority of which clearly show by context alone that "firstborn" (in time) is the intended meaning.

The first example, Ps. 89:27, has God saying about "David, my servant" that

"he shall call upon me, saying, `Thou art my Father, my God.... And I will make him firstborn [prototokos], higher than the kings of the earth.... my covenant shall be firm with him." - Septuagint. (Cf., KJV, ASV).

It is true that David was not the firstborn of his father Jesse, nor was he the first king of Israel. However, the first king of Israel, Saul, was rejected by God and removed from God's throne, no longer God's king. The second king of Israel, David, was the first king to remain faithful until his death and, hence, the "firstborn" of all the faithful kings of Israel who will be resurrected by God. He will be "firstborn" (first in time on Israel's throne) among all those kings who will return. However, he certainly will not be pre-eminent over one of those other kings who is his descendent: Jesus Christ.

We also find that David is the first king whom God made a covenant with for an "everlasting kingdom" - 2 Sam. 7:12-16. [He is also the first of the descendants of Judah who are to rule forever (Saul was descended from Benjamin) - Gen. 49:10] He may be considered "firstborn" in this sense, also.

So we can see that Saul was the first king of Israel, but he didn't remain faithful to God ["call upon me, saying `thou art my Father, my God'"]. The very first king of Israel to remain faithful to God was David. In that sense, then, David became "firstborn" [of all succeeding faithful earthly kings of Israel].


However, the later fulfillment of Ps. 89:27 is in the person of Jesus Christ (who is the firstborn of God in another sense) and not the literal David. We see the Messiah being called, figuratively, "David, my servant" at Ezekiel 34:23, 24 just as he is in this Psalm (89:20). We see the final fulfillment of Ps. 89:26-29 in Jesus Christ (Luke 1:32, 33; Heb. 1:5, 6; Jn 20:17).

The second example (Ex. 4:22) is probably the most-used by those trinitarians attempting to prove a "pre-eminent" meaning for prototokos. Here is how it is worded in the Septuagint: God says, "Israel is [the] firstborn [prototokos] son of me." Context reveals that this is the nation of Israel which Jehovah is calling his "firstborn." So in what sense was Israel first in time in relation to Jehovah? It was the first nation to be chosen by him. It has always (since the time of Moses) been the first, but it has certainly never been "pre-eminent" among the nations!

And, of course, we must not change the inspired writer's genitive noun ("of me") in this verse to "over me" as has been done at Col. 1:15 in a few trinitarian Bibles (e.g. NIV). How ridiculous to "interpret" this so that God says: "Israel is the `pre-eminent one' OVER me"! (But, of course, this is precisely what some trinitarians have done with Col. 1:15 - "the pre-eminent one over all creation"!!

God's calling the nation of Israel his "firstborn son" obviously means the first nation he has caused to come into existence to be his own (and others must someday follow).

The third example (Jer. 31:9) is actually found at Jer. 38:9 in the Septuagint). Again God is speaking of the nation of Israel (see context of entire chapter): "I have become a father to Israel, and Ephraim is [the] firstborn [prototokos] OF me."

So how can we understand Ephraim being Jehovah's "firstborn"? - Jer. 31:9.

Here Ephraim is obviously called Jehovah's firstborn in some figurative sense. (The person, Ephraim, was, of course, long dead at this time.) Certainly neither Ephraim, nor even the tribe of Ephraim, was ever Jehovah's "pre-eminent one" or (more parallel to the trinitarian interpretation of Col. 1:15) "the pre-eminent one OVER Jehovah"!

So to explain the use of "firstborn" at Jer. 31:9, the very trinitarian ecumenical study Bible, The New Oxford Annotated Bible, 1977 ed., tells us that

"as [the tribe of] Ephraim is restored, so is all Israel" - p. 954.

This interpretation shows the understanding that the tribe of Ephraim is to be restored first in time ("firstborn"), and then the rest of Israel is to be restored. Notice there is no "pre-eminence" interpretation by these highly respected trinitarian scholars!

Another possibility suggested by trinitarians for "firstborn" at Jer. 31:9 is that, since the land of the tribe of Ephraim is where "the original [first] place of worship [the tabernacle] from the time of Joshua to that of Samuel" - (NAB, St. Joseph ed., p. 902) - was located, in Shiloh, it is God's "firstborn" in that respect (again in the sense of first in time). Or, as explained in Jer. 7:12,

"Go now to my place that was in Shiloh [in `Ephraim'] where I made my name dwell at first" - RSV, NRSV, NIV, and cf. NAB (`91) "in the beginning."

But the trinitarian reference work, The New International Dictionary of New Testament Theology, Vol. 2, p. 306, Zondervan, 1986, gives us the most probable explanation: the nation of Israel was also called `Ephraim' "by the contemporary prophets, e.g., Isa. 7:1-9, after the central region associated with the name of the younger of the two sons of Joseph."

So we merely have a parallelism at Jer. 31:9 - (1) "I, Jehovah, am a father (I created it) to the nation of Israel, and (2) `Ephraim' (`Israel') is the first nation I have created (`first-born')." - Compare the parallelism at Hosea 11:8. Again we see a confirmation of Ex. 4:22 (the 2nd "example" above) that Israel was the first nation formed at God's direction, and no hint of "pre-eminence" but only the meaning of first in time for "firstborn"! (This is simply one of the many scriptural uses of "Father," "Son," [or "Firstborn," "onlybegotten," etc.] and "brought forth" [or "begot"] to figuratively describe the CREATOR of something and his CREATION!)

And, again, how absurd it would be to interpret this as "Ephraim is the `pre-eminent one' over me [God]."

The 4th example seems, perhaps, the weakest of all of those cited in the OT, but no less an authority than the very trinitarian Biblical Greek scholar W. E. Vine points to Deut. 21:16, 17 as evidence for a "pre-eminent" interpretation for "first-born." The Septuagint reads:

(15) "And if a man have two wives, the one loved and the other hated, and both the loved [wife] and the hated [wife] should have born him children, and the son of the hated should be first-born [prototokos]; (16) then it shall be that whensoever he shall divide by inheritance his goods to his sons, he shall not be able to give the right of the firstborn to the son of the loved one, having overlooked the son of the hated, which is the firstborn [prototokos]. (17) But he shall acknowledge the firstborn [prototokos] of the hated one [wife] to give to him double of all things which shall be found by him, because he is the first [arkhe: beginning] of his children, and to him belongs the birthright." - The Septuagint Version of the Old Testament, Zondervan Publishing House, 1970.

It is obvious that prototokos here means only "one born first." The birthright itself must not be confused with the one it is usually given to: the firstborn (prototokos)! And to "interpret" verse 17 as "he shall acknowledge the `pre-eminent one' OVER the hated [wife]" is clearly ludicrous!

Another example from the OT sometimes used by trinitarians can be found at Job 18:13 -
"His skin is devoured by disease, The firstborn of death devours his limbs" - NASB.

A few trinitarians attempt to interpret this as an example of "firstborn" meaning "pre-eminent" or "foremost." But there is no honest reason to insist that "the firstborn of death" at Job 18:13 must mean something like "the principal disease" (JB footnote). In the first place, to be parallel with the trinitarian-concocted understanding of Col. 1:15 it would have to be interpreted as "the pre-eminent one over death" - which it clearly does not and can not! In the second place, a careful study will reveal that this scripture is literally calling the disease which ravages the wicked man "the firstborn son of death": the first child (or creation) produced by the "god of death."

The NIV Study Bible (1985 ed.) tells us in the footnote for Job 18:13: "death's firstborn. See 5:7." And the footnote for Job 5:7 says:

"sparks. Lit[erally] `sons of Resheph.' In Canaanite mythology, Resheph was a god of plague and destruction [death]. `(Sons of) Resheph' is used as a poetic image in the OT for fire (SS 8:6), bolts of lightning (Ps 78:48) and pestilence (Dt 32:24; Hab 3:5)."

Obviously the NIVSB has referred "death's firstborn" at Job 18:13 to "Sons of Resheph [Death]" to show that the poetic image used in the OT has more than one son attributed to Resheph. Apparently the firstborn of those sons (the first "son" Resheph created) was pestilence. And it is this pestilence (the 'first born of Death') that "eats away parts of his skin ... [and] devours his limbs" at Job 18:13. (This is why the New English Bible renders Job 18:13 as "Death's eldest child.")
A related interpretation (which I prefer):

"[even] the firstborn of death shall devour his strength; ....

"it signifies not what presides over death, but what death first produces, which are corruption and rottenness, dust and worms; these are the firstborn of death, or the firstfruits and effects of it, and which devour and destroy not the skin only, but the whole body and all its members" - The New John Gill Exposition of the Entire Bible.

The first of the two NT examples sometimes used by trinitarians is Ro. 8:29 -

"Those Christians whom he [God] foreknew he also predestined to be conformed to the image of his Son, in order that he might be the firstborn [prototokos] AMONG many brethren." - RSV.

I have found no Bible (trinitarian or otherwise) which renders this scripture with a meaning of "pre-eminent one over many brothers" in spite of the interpretation a few trinitarian "scholars" attempt to give it. Since the word "among" (en in the NT Greek), not "over," is used here, it cannot honestly be rendered as these trinitarians would like. In fact these trinitarian Bibles translate it as "eldest [prototokos] among many brethren" - NEB, REB, CBW, JB, NJB, AT, and Weymouth. This clearly shows the intended meaning of prototokos as first in time not "pre-eminence" (as a careful study of the context also confirms).

As for the other NT "example," Col. 1:18, "proving" the possibility of a "pre-eminent" interpretation for prototokos, all we have to do is examine Col. 1:15-18 carefully.
We see prototokos clearly meaning "the one born (or reborn) first" at 1:18 where Jesus is the firstborn (or first to return from death to eternal life) from the dead.

To make it even clearer, the trinitarian The Jerusalem Bible (cf. NEB; REB; and Beck) translates it: "first to be born from the dead." - Compare 1 Cor. 15:20, 23 and Rev. 1:5. Surely there is no doubt that first in time is intended here, not "pre-eminence."

"firstborn. Christ was the first to rise from the dead with a resurrection body. .... Others who were raised from the dead ... were raised only to die again." - The NIV Study Bible footnote for Col. 1:18.)

It is also very plain that Paul frequently speaks of certain Christians being resurrected as spirit persons in heaven and that Jesus was the firstborn of these dead, i.e. the first of many persons who are to be resurrected to eternal life in heaven.

1 Cor. 15:20 - "Christ has been raised from the dead, the first fruits [`the first of millions' - LB; `the very first to rise' - Phillips] of those who have fallen asleep." - RSV and many others.

1 Cor. 15:49 - "Just as we [faithful Christians who have the hope of being resurrected to heaven] have borne the image of the man of dust [Adam], we shall also bear the image of the man of heaven [Jesus]."

The Apostle John also writes of this truth and also uses the term "first-born" to describe it: "and Jesus Christ ... the firstborn [prototokos] of the dead" - Rev. 1:5, RSV. Or "first to be raised from death" - GNB; also see LB, Beck, and Beck (NT)). Even the extremely trinitarian The Amplified Bible explains "firstborn of the dead" at Rev. 1:5 as, "that is, first to be brought back to life."

Rev. 2:8 makes a similar statement, but could be phrased as the "only-begotten" from the dead (in the sense that Jesus is the only one to be raised to heavenly life by God Himself. All others are to be raised through Jesus). "The first and the last" simply means the only one in some sense. Adam, for example, was the 'first and the last' in the sense that he was the only one to be created directly from the elements of the earth.

This "firstborn" and "only-begotten" concept for the second creation (resurrection to eternal life) also explains why Jesus can be called the firstborn and the "only-begotten" in another sense: The first of God's creation ("Firstborn") and the only one ('only-begotten') created directly by God Himself.

These examples in Revelation are therefore clearly a repetition of this same well-established truth that Paul is restating at Col. 1:18.

We cannot seriously believe that Paul is telling us at Col. 1:18 that Jesus is the "pre-eminent one" over the dead. Especially since the actual wording by Paul is "the beginning [arkhe], firstborn [prototokos] OUT OF [ek] the dead." - see any interlinear New Testament (or as also confirmed by John "The firstborn OF the dead." - Rev. 1:5). There can be no honest doubt that Col. 1:18 does not mean "pre-eminent one OVER the dead"! It clearly means "the first one resurrected to eternal life in the `new creation'."
Remember, the above 7 examples are the very best "proof" available to desperate trinitarian scholars that prototokos can mean "pre-eminent" in Bible usage! But even they (like the more than one hundred other examples of prototokos in the Bible) show that only firstborn in time is meant.

Now notice how the first use of prototokos (in Col. 1:15 - "the first-born of all creation") is used as a complement for the second use of prototokos (in Col. 1:18 - "the first-born from the dead").

That is, being "firstborn of all creation" is equated with and added to being "firstborn from the dead" (or first of the final creation - the ones who will receive eternal life). When these two "firstborns" (the first and the last) are added together the sum is one who is "first [proteuo] in everything" - Col. 1:18, JB, NWT, Living Bible (also known as The Book and The Word), and the New Testament in the Language of Today (Beck).

(Also notice how the ancient Aramaic text renders Col. 1:18 - "he is the beginning, the firstfruits of the resurrection from the dead, that in all things he might be first." - Lamsa. Compare 1 Cor. 15:20 - "But now Christ has been raised from the dead, the first fruits of those who are asleep [in death]." - NASB.) In both cases -'firstborn' and 'firstfruits' - we understand the meaning as the first one produced.)

To insist on a literal "born" (rather than the figurative first "created") in Col. 1:15-18 would mean that we must also understand him as literal vegetation ("fruit") at 1 Cor. 15:20 instead of the figurative 'first one produced or created from the dead'!

So when the distinction of being the very first person to be resurrected to eternal life in the "last creation" is added to the distinction of being the very first of God's original creation, we truly have a person who was the very "first in everything." (Proteuo is often translated "pre-eminent" for this verse. It literally means, however, either `to be first' or 'hold the first place' - Thayer. It derives from protos which means first in time, place, or rank - Thayer.)

"Jesus was the first-born (prototokos) of his mother (Mt. 1:25; Lk. 2:7) .... Jesus is also the first-born of his heavenly Father. He is the first-born of all creation...(Col. 1:15-17). Similarly, he is the first-born in the new creation by being raised first from the dead" - New Bible Dictionary, (2nd ed.), 1982, p. 378, Tyndale House Publ.

This respected trinitarian publication clearly admits, then, that "firstborn" is used in the same way ("very first in time") for each of these scriptural uses of prototokos for Jesus: Jesus was the very first child Mary brought into existence (Matthew 1:25); Jesus was the very first person brought into existence in the new creation (Col. 1:18; 1 Cor. 15:20; Rev. 1:5); and Jesus was the very first thing brought into existence in God's first creation!!

Furthermore, these two "first-borns" of Col. 1:15 and Col. 1:18 must closely parallel each other in meaning to make the comparison (or "addition") sensible and complete! They are of like kind (firstborn in time, not "pre-eminence") so they can be added together and summed up: "So that he should be first in everything." - Col. 1:18. It would be inappropriate to have the first use of prototokos (Col. 1:15) mean "pre-eminent" for the first creation and the second use of prototokos (Col. 1:18) mean (as it so obviously does) "first in time" for the "last creation."

So Paul is probably making a play on words with the two definitions of proteuo when he sums up by saying Jesus is "first [proteuo] in everything." He must intend the meaning of `first in time' to agree with the rest of his metaphor.

There is no proper reason to change the intended meaning of prototokos at Col. 1:15 so that it differs from every other Bible use of the word. The only reason that some trinitarians attempt to interpret it as "pre-eminent one" is that they don't like what it actually, literally says!

Finally, notice that even if Col. 1:15 could be properly translated as "the pre-eminent one of all creation," it could only mean one thing: that he is the highest one of all created things (that is, he IS ONE OF THOSE CREATED THINGS)!

It cannot mean that he is the highest individual over all created things. This is an impossible interpretation for two reasons:

(1) The word "pre-eminent" actually rules out the possibility of a double interpretation. For example: "The leader OF the wolves" is capable of a double interpretation: (A) "The Leader" could be one of the wolves himself. And (B), although unlikely, it is possible that the leader of the wolves is not one of the wolves himself. It is possible that he is a dog, coyote, or even a man. However, the phrase "smartest (or `smartest one') OF the wolves" does not allow for such an ambiguity, and it is, therefore, certain that this "smartest one" IS one of those wolves.

"Pre-eminent" (or "pre-eminent one"), like "smartest" (or "smartest one"), also does not allow for that ambiguity. The "pre-eminent one OF creation" has to be a part of that creation himself!

(2) As we have already seen, the Bible clearly and repeatedly states that Jehovah (the Father alone) is pre-eminent over all creation. Therefore Jesus cannot be the pre-eminent one over all creation but is the first creation of God!

We have also seen that in the Bible the term "firstborn of ..." never means "pre-eminent one over ..."! "Firstborn of Abraham" for example never means the "pre-eminent one over Abraham." Even if we could properly allow "pre-eminent" as a meaning for prototokos, it would mean no more than "pre-eminent one of Abraham's creation (or procreation)"! Whenever anyone calls Jesus the firstborn son of God, it plainly means the first of the "children" God has produced. (Obviously it does not mean "the pre-eminent one over God.") Therefore, when Jesus is called the firstborn of creation, it clearly means that he is the first of that creation that God has produced.

And, again, even if "firstborn" could mean "pre-eminent one," Col. 1:15 would still be saying that Jesus is the "pre-eminent one" of creation. In other words, he is part of that creation, albeit the first and highest part!

We should also consider that those whom God calls son are those whom he has created: Luke 3:38; Rev. 21:7; Gen. 6:2; Job 1:6; Ps. 89:6 [f.n. in RSV and compare LB]; Gal. 3:26. Jesus' very title, the Son of God, indicates that he was created. The International Standard Bible Encyclopedia tells us that angels are called `sons of God' in the Scriptures. Then it states:

"the creation of angels is referred to in Ps 148 2, 5 (cf Col 1 16). They were present [in the beginning] at the creation of the world ... (Job 38 7)." - pp. 132, 133, Vol. 1, Eerdmans, 1984 printing.
In the very same way, those who are said to be the "image" of God are not God himself (he's obviously not his own image) but a part of God's creation! Notice who the image of God is in these scriptures: Gen. 1:26; 9:6; 1 Cor. 11:7; 2 Cor. 3:18.

Therefore, when Col. 1:15 says Jesus is "the image of the invisible God, the firstborn of all creation" (RSV), we know Paul is speaking of someone who is the very first creation made by God (and was created "in his image" long before man was also created "in his image")! - Cf. Gen. 1:26, 27. - See the IMAGE study paper.

There are only two valid, scriptural understandings for "firstborn of ...." If it is used with a single individual (e.g., `firstborn of Moses'), then it means the firstborn has been produced (or created) by that individual. If it is used with a group of individuals (e.g., `firstborn of the herd' - Deut. 12:6; 15:19; `firstborn of our sons' - Neh 10:36), then it always means one who was produced as a part of that group!

That's why, for example, "the firstborn" at 2 Chron. 21:3 may be understood only as either "the firstborn of Jehoshaphat" or "the firstborn of all Jehoshaphat's children." In the first case the firstborn is the first production (or creation) by Jehoshaphat. In the second case the firstborn is the first to be created of all Jehoshaphat's CHILDREN. But in either case it is speaking of the first one created!

Clearly, then, if Jesus is called the firstborn of God, he is being identified as the one first produced by God. And when he is called the firstborn of creation (a group of individuals), he is being identified as one who was produced or created as a part of that group. In other words: Out of all things created by God, Jesus was the very first.
* * * * *

Some anti-Watchtower writer has evidently come up with an idea that I have seen used in letters to fellow Christians here in Ketchikan. The argument usually goes like this: "If Paul had really meant `the first creation by God' at Col. 1:15, he would have used the word protoktistos which means `first creation' instead of prototokos." (Notice the argument here is not that proto doesn't actually mean "first in time" but that ktistos ["creation"] is more appropriate than tokos ["born"]!)

I do know, however, that protoktistos was never used by any inspired NT scripture writer. It should certainly be no surprise, therefore, to learn that it isn't used at Col. 1:15, 18 !
Furthermore, the Bible frequently uses the word for "born" in place of "made" or "created" as would be expected from the common Bible idiom of "Father" as creator or source - Ps. 90:2 ("brought forth" in some translations is the Hebrew word for "born"); Is. 66:8-9; Job 38:28-30; Prov. 8:24-25. So not only was protoktistos not used in the NT at all, it was completely unnecessary because "first-born" could be used with the very same meaning!