"although usually present, the HG valves in P. siculus may be flexible in their morphology or even in their presence, and they are not fixed features in the Pod Mrčaru lizards." — Wehrle, Herrel, German and colleagues, 20201
Summary
In 1971 ten Italian wall lizards were moved from one Adriatic islet, Pod Kopište, to a smaller one nearby, Pod Mrčaru. When biologists returned in 2004 to 2006, the lizards had turned to eating plants, grown bigger heads and a harder bite, and developed cecal valves, folds across the gut that slow food so microbes can ferment it. The 2008 paper in PNAS was widely reported as a new organ evolving in 36 years. This article reads that paper and every follow-up that could be found, and scores the case by the rules of Measuring Biological Information.
- The observations are sound. Plants made up 34% to 61% of the Pod Mrčaru diet against 4% to 7% on Pod Kopište. Bite force was higher, density about five times higher, and territories gone. The native Dalmatian wall lizard had vanished from the islet.2 2
- The discoverers never claimed the change was genetic. Their paper says that plasticity and maternal effects "are needed" as further studies.2
- The valves are not new to lizards. They occur in plant-eating iguanas, agamids and skinks, in other plant-eating lacertids, and in some mainland Balkan green lizards.3 4
- The valves behave like a response to diet. Twelve of 16 wild Pod Mrčaru lizards had them, and none of 20 kept on insects for 15 weeks did.5 A later study by the same group called them "not fixed features."1
- The head and bite differences did not persist in a common garden. Plasticity was among "the main drivers" at this stage.6 Whether valves were tested in that garden could not be read.
- The genome shows a founder event and a handful of outlying loci, none tied to a trait.7
- The popular accounts left out the open question, and the creationist replies mostly got it right. The press release and Dawkins's book did not mention it. Answers in Genesis predicted in 2008 that the valves would "dwindle" on an insect diet, and they did, although several creationist writers got the anatomy and genetics wrong.8
- Comparable cases are mixed. Florida green anoles kept larger toepads in a common garden, and Bahamian brown anoles' legs proved partly plastic.9 10
Verdict. The case is not yet scoreable as gain, loss, modification or transfer, because no inherited change has been shown. It cannot be cited as an observed new organ, and it cannot be dismissed, because a working valve that appears when needed is a capacity that itself needs explaining. It fits the young-earth claim that animals were made able to adapt, and says nothing yet about inherited diversification. One experiment would settle it: hatch eggs from both islets and raise them on both diets.
Part 1: What was done and what was found
The 1971 move
In the summer of 1971 a team led by Eviatar Nevo was studying competition between two lizards on the small islands of the Adriatic. The Italian wall lizard, Podarcis siculus, and the Dalmatian wall lizard, Podarcis melisellensis, rarely share a small island. Nevo's paper opens with that pattern: "Competetive exclusion between Lacerta sicula and L. melisellensis characterizes the small islands of the Adriatic Sea."11 Earlier introductions had been followed up that summer. On one island "the introduced species appears to be replacing the native form," and the paper closes by announcing "a reciprocal introduction involving the two species on the islands of Pod Kopiŝte and Pod Mrĉaru."11
The experiment was about competition, not diet. Five adult pairs of Italian wall lizards were moved from Pod Kopište to Pod Mrčaru, an islet nearby that was held by the Dalmatian species.2 Nothing about their guts was recorded at the time.
The return, 2004 to 2006
Anthony Herrel, Katleen Huyghe, Bieke Vanhooydonck, Thierry Backeljau, Karin Breugelmans, Irena Grbac, Raoul Van Damme and Duncan Irschick visited both islets in spring and summer of 2004, 2005 and 2006, and published in PNAS in March 2008.2 Their findings, taken from the paper itself:
- The native species was gone. Twice-yearly visits "show that this species has become extinct on Pod Mrčaru."2 The competition experiment had its answer.
- The new lizards descend from the old ones, as far as the test could show. Two stretches of mitochondrial DNA from seven Pod Mrčaru lizards and eight Pod Kopište lizards were "genetically indistinguishable."2 Two short mitochondrial genes cannot rule out a later arrival from elsewhere, and the paper's rate figures say they "assume no additional colonization of the island by P. sicula."2
- The diet had changed. Stomach flushing of 330 lizards showed plants making up 34% of the food by volume in spring and 61% in summer on Pod Mrčaru, against 7% and 4% on Pod Kopište.2 About half the plant matter on Pod Mrčaru was leaves and stems, which are hard to digest.2
- Heads and bites were bigger. Pod Mrčaru lizards had longer, wider and taller heads, and not only because they were larger overall. Males bit with 11.25 newtons against 9.85, females with 6.70 against 4.99.2
- The gut had valves. Dissection of preserved lizards found "cecal valves (Fig. 4) in all individuals, including a hatchling (26.4-mm snout-vent length, umbilical scar present) and a very young juvenile (33.11-mm snout-vent length) examined from Pod Mrčaru."2 A cecal valve is a fold across the start of the large intestine. It slows food down and makes a chamber where microbes can ferment plant fibre.
- The gut had worms. Nematodes "were common in the hindgut but absent from individuals from Pod Kopište."2 The authors read this as a sign of a new partnership with gut microbes.
- There were more lizards and fewer territories. Unbaited traps caught 0.05 lizards per trap per hour on Pod Mrčaru and 0.01 on Pod Kopište, and the Pod Mrčaru lizards "do no longer appear to defend territories."2
What the paper does not say
Three gaps matter for what follows. The paper does not say how many lizards were dissected for valves. "All individuals" could be a handful or dozens. It does not say how many Pod Kopište lizards were opened and found without valves. For the absence of valves in other P. siculus it cites an earlier paper by the same group rather than a count.2 And it gives no basis for its generation count.
Years and generations
The paper's headline is "in only 36 years (≈30 generations)."2 Thirty-six years runs from 1971 to 2007, when the paper was written. The lizards were measured between 2004 and 2006, 33 to 35 years after the move. Thirty generations in 36 years means a new generation every 1.2 years. The paper does not say where that figure comes from. A later genomic study repeats "~30 generations" and also gives no derivation.7 The count may be too high if these lizards breed first in their second year, but no study found gives the age at first breeding on these islets. The arithmetic in Part 6 uses 20 to 30 generations to cover that doubt.
Two numbers that do not agree
Herrel's rates reach "up to 8,593 darwins or 0.049 haldanes."2 Neither figure is the largest in the paper's own Table 1, where female body mass reaches 9,441 darwins and female head width 0.051 haldanes, and no cell reads 8,593.2 The islet areas also disagree between papers. Herrel gives Pod Kopište as 0.09 km² and Pod Mrčaru as 0.03 km².2 Maxime Taverne and colleagues, from the same research group eleven years later, list 7,915 m² and 2,931 m², about a tenth as large.12 The ratio of the two islets agrees. Neither error changes the story, but both show why numbers from this case should be taken from the tables and not from summaries.
What later visits added
Later work by the same group and its partners filled in the picture. Bart Vervust and colleagues, working in April 2008, found wider teeth, longer stomachs and small intestines, slower passage of food and better digestion in Pod Mrčaru lizards.5 Transect counts put the density at about 3,082 lizards per hectare on Pod Mrčaru against 1,045 on Pod Kopište.13 The gut bacteria differ: plant-eating Pod Mrčaru lizards carry more kinds of microbe and more of the methane-making Methanobrevibacter.13 Three-dimensional models of the skull show that small changes in shape and jaw muscle together raise bite force.14 Beck Wehrle, Herrel, Donovan German and colleagues found that Pod Mrčaru lizards digested plant diets better in the laboratory, but found no cellulase activity "anywhere in their guts."1 1
None of this is disputed. A small population, founded by ten animals, changed its diet, its head, its gut, its microbes, its numbers and its social life inside four decades. What is disputed is what kind of change it was.
Part 2: Are cecal valves new?
How rare they are
Herrel's paper stresses how unusual the structure is: "The fact that <1% of all currently known species of squamates have cecal valves (13, 14) illustrates the unusual nature of these structures in this population."2 Squamates are lizards and snakes together. The two works cited for that figure are the group's own 2004 survey of plant-eating lacertids and J. B. Iverson's 1980 study of iguanas. Iverson's abstract gives no percentage, and the 2004 paper could not be read in full, so the figure rests on Herrel's authority.3
Iverson's work shows what the valves are. Every iguana species he examined except the marine iguana has "from one to eleven transverse valves in the proximal colon," and they "apparently evolved as simple infoldings of the medical colic wall."3 "Medical" is the abstract's misprint for medial. In 1980, "Herbivorous species of the Scincidae, Agamidae, and Iguanidae are the only lizards known to exhibit colic partitioning."3 So the valve is a fold of the gut wall, and it goes with plant eating across three families that are only distantly related. That pattern alone says the capacity to make it is widespread in lizards.
They are known in the lizard's own family
Wall lizards belong to the Lacertidae. In 1980 no lacertid was known to have valves. By 2008 Herrel's group could write that the Pod Mrčaru valves "are similar in overall appearance and structure to those found in herbivorous lacertid, agamid, and iguanid lizards."2 The same paper calls the structure "rare in this family."2 Rare is not absent.
A second genus makes the point plainly. Sagonas, Pafilis and Valakos dissected Balkan green lizards, Lacerta trilineata, from islands and from the mainland in 2015. "The dissection of the hindgut revealed an unexpected finding, the presence of cecal valves that were more frequent in island lizards."4 More frequent on islands means present on the mainland too. In that species the valve is a variable trait that some individuals have and others lack, and its frequency tracks where the lizards live.
The question list for this article named the Canary Island lizards, Gallotia, and the Balearic wall lizards, Podarcis lilfordi and P. pityusensis, as places to look for valves. All of them eat a great deal of plant matter.15 No report of valves in them was found in the sources read here. The lacertid valves that Herrel's 2004 survey describes may include some of them, but that paper was read only in abstract.16 This is a gap in this article, not evidence of absence.
Plant eating is common on small islands
Pod Mrčaru is not the only island where P. siculus turned to plants. Taverne's survey of twenty Adriatic populations found diets "ranging from strictly insectivorous populations (e.g., Brusnik or Mala Palagruža) to almost completely herbivorous populations (e.g., Sušac or Pod Mrčaru)," and named Sušac, Pijavica and Pod Mrčaru as the plant-eating P. siculus islands.12 Valentin Pérez-Mellado and Claudia Corti wrote in 1993 that "Among lacertid lizards, plant consumption is fairly common in insular populations."15 A small island with few insects pushes lizards toward plants again and again.
What this means
The valve on Pod Mrčaru is new to that population and was not seen in its source. It is not new to lizards, not new to lacertids, and not new as a kind of structure. It is a fold of the gut wall of a type that appears wherever lizards eat leaves. That is consistent with two readings. Either the ability to make the fold is an old developmental capacity that a plant diet switches on, or a similar mutation has been selected many times. The first reading needs no new genetic information on Pod Mrčaru. The second would. The experiments in Part 3 were designed to tell them apart.
Part 3: Genetic change or plasticity?
What the discoverers said
Herrel's team did not claim to have shown a genetic change. Their paper says so in its discussion:
"Although the presence of cecal valves and large heads in hatchlings and juveniles suggests a genetic basis for these differences, further studies investigating the potential role of phenotypic plasticity and/or maternal effects in the divergence between populations are needed."2
Phenotypic plasticity is the ability of one set of genes to build different bodies in different conditions. A maternal effect is a trait a mother passes on through the egg or her body rather than through her genes. The hatchling with valves is the paper's best evidence for inheritance, and it is weak. A hatchling on Pod Mrčaru has a Pod Mrčaru mother, eats Pod Mrčaru food from its first days, and picks up Pod Mrčaru gut microbes. One animal cannot separate those causes.
Test 1, 2010: take the plants away
Bart Vervust, Panayiotis Pafilis, Efstratios Valakos and Raoul Van Damme ran the first test, partly by accident. They caught lizards on both islets in April 2008. In freshly caught animals, "In 12 of the 16 individuals (75%) examined from Pod Mrčaru, the hindgut contained cecal valves. In sharp contrast, we found no such valves in any of the 27 individuals from Pod Kopište."5 So even in the wild, one Pod Mrčaru lizard in four had no valve.
Twenty Pod Mrčaru lizards, eight males and twelve females, were taken to Antwerp and fed only arthropods. After 15 weeks a power cut killed them: "These 20 lizards had died of oxygen deficiency following a power cut in the animal house at Antwerp University."5 The team dissected them. "In addition, we found no evidence of cecal valves in any of the twenty specimens that had been fed arthropods."5 At the wild rate of 75%, about fifteen of the twenty should have carried valves. None did. Their small intestines and hindguts were also lighter.5
The authors drew the conclusion carefully:
"Many comparative studies have tacitly assumed that the distinctive features of plant-eating lizards (large body size, skull dimensions, special dentition, gut morphology) are a product of genetic adaptation to the special demands of a plant-based diet (e.g., Van Damme 1999; Cooper and Vitt 2002; Espinoza et al. 2004; Herrel et al. 2008). Our results suggest that in P. sicula, at least some of the changes associated with a dietary shift toward a higher proportion of plant material may be plastic."5
Four limits apply. The same animals were not examined before and after, so the valves are inferred to have gone, not watched going. The text says 15 weeks, while the notes to two of the paper's tables say 6 months.5 Some differences survived the diet, because the laboratory Pod Mrčaru lizards still had longer guts than wild Pod Kopište lizards.5 And only one direction was tested. Nobody has fed plants to Pod Kopište lizards to see whether valves grow.
Test 2, 2020: count again, across years
Beck Wehrle and colleagues opened more guts over several years. They found valves in 28 Pod Mrčaru lizards "spanning four seasons over three additional years," and none in 29 from Pod Kopište. But three Pod Mrčaru lizards examined in summer 2013 "did not have clear epithelial folds (valves) in their HGs."1 HG is their abbreviation for hindgut. They concluded that "although usually present, the HG valves in P. siculus may be flexible in their morphology or even in their presence, and they are not fixed features in the Pod Mrčaru lizards."1 Herrel is a co-author of that sentence.
On the digestive differences as a whole, the same paper states the open question in plain terms: "We currently do not know whether the observed changes represent flexibility (Piersma and Drent 2003) or evolved differences with underlying genetic changes (Garland and Adolph 1991)."1 It adds that better digestion of plants "may be mediated through the microbiome of the HG (i.e., not necessarily an evolved difference in the lizards themselves; Garland and Adolph 1991)."1
Test 3, 2024: raise them side by side
The experiment that the 2008 paper asked for has now been done for the head and the bite. Iva Sabolić, Anamaria Štambuk, Herrel and twelve others combined genome data with a crossing experiment and a common garden, which means raising animals from both populations under the same conditions.6 Their abstract reports:
"Diverged traits related to head size and shape or bite force showed moderate heritability in a crossing experiment, but between-population differences in these traits did not persist in a common garden environment. Our results confirm the existence of sufficient additive genetic variance for traits to evolve under selection while also demonstrating that phenotypic plasticity and/or genotype by environment interactions are the main drivers of population differentiation at this early evolutionary stage."6
The head and bite differences that made the 2008 paper famous did not persist when lizards from both islets grew up in the same place. Head size can be inherited within a population, but the difference between the islands is mainly a response to how the animals live. Only the abstract of this paper could be read. It does not say whether guts or valves were scored in the common garden. The full text sits behind a publisher paywall, and Sabolić's 2021 doctoral thesis, which reports the same work, sits behind an automated robot check on the University of Zagreb repository. Reading one of them is the most important open task for this article.
Test 4, 2023: look at the genome
Stéphanie Sherpa and colleagues read about 82,000 short stretches of DNA from 24 lizards on Sušac, Pod Kopište and Pod Mrčaru. They confirmed the founder event: Pod Mrčaru has the least variation and the most alleles fixed by chance.7 Their scan "identified just a handful of loci showing large allelic shifts between ecologically divergent populations," and they drew the conclusion that the differences "are determined by a small number of large‐effect loci and/or that phenotypic plasticity plays a major role."7 Of 29 outlier loci across all their comparisons, seven lay in or near known genes. Six were inside introns and one lay upstream of a gene.7 None has been tied to the valves or to any measured trait. They also found one Pod Mrčaru lizard of mixed ancestry, "potentially indicating very recent gene flow with Sušac."7 Sušac is a larger island whose wall lizards also eat plants. So Pod Mrčaru may not be a closed population.
Guts are flexible in other animals
A gut that remodels itself with diet would be no surprise. J. Matthias Starck's review opens: "The vertebrate gastrointestinal tract is a flexible system that can individually be modified to account for changes in amount and quality of food as well as changes in internal demands."17 In reptiles, he adds, enlarging the gut lining "does not involve cell proliferation, thus it is presumably an energetically cheap process."17 Kohl and colleagues ran the forward experiment in a South American lizard, Liolaemus ruibali. Animals fed 90% plants grew "significantly longer small intestines and larger hindguts, demonstrating that gut morphology is phenotypically flexible."18 Across ten wall-lizard populations in the Skyros islands of Greece, islet lizards had longer guts and slower digestion, and the authors called "digestive plasticity" a key mechanism.19 None of these studies examined valves. Among the sources read here, the Pod Mrčaru valve is the only one whose loss on an animal diet has been reported.
What has been shown, and what has not
| Claim | Status | Source |
|---|---|---|
| Diet, head, bite, gut, density and behaviour differ between the islets | Shown, repeatedly | Herrel 2008; Vervust 2010; Wehrle 2020 |
| The Pod Mrčaru lizards descend from the 1971 founders | Shown by mitochondrial and genome data; one migrant from Sušac detected | Herrel 2008; Sherpa 2023 |
| The valves occur in most, not all, Pod Mrčaru lizards | Shown: 12 of 16, then 28 of 31 | Vervust 2010; Wehrle 2020 |
| The valves disappear on an animal-only diet | Shown once, in 20 adults, without before-and-after data | Vervust 2010 |
| Plants make valves grow in lizards that lack them | Not tested | None |
| Head and bite differences persist in a common garden | Tested: they did not | Sabolić 2024 |
| The valves persist in a common garden | Unknown from the abstract | Sabolić 2024 |
| Some allele frequencies changed by more than drift | Suggested by a handful of outliers | Sherpa 2023 |
| A mutation causes any trait difference | Not shown | None |
The experiment that would settle the valve question is simple to state. Hatch eggs from both islets in one laboratory. Raise half of each clutch on insects and half on plants. Open the guts at the same age. If valves appear only in plant-fed lizards from both islets, the valve is plastic and the capacity is shared. If they appear in plant-fed Pod Mrčaru lizards but never in plant-fed Pod Kopište lizards, the populations differ genetically in the ability to make them. If they appear in Pod Mrčaru lizards on either diet, the trait has become fixed. The third result is the only one that would make this a case of an inherited new structure.
Part 4: How the case is used
The popular account
The university press release that went out with the paper set the tone. ScienceDaily ran it on 18 April 2008 under the headline "Lizards Undergo Rapid Evolution After Introduction To A New Home." It quotes Irschick: "Our data shows that evolution of novel structures can occur on extremely short time scales."20 It does not mention that the genetic basis was untested. It also contains two errors. It says DNA tests showed the lizards "were genetically identical to the source population," when two short mitochondrial fragments were compared. And it calls the gut worms "microorganisms called nematodes that break down cellulose."20 Nematodes are worms. Oddly, its own description of the valves is a plasticity claim: "Eating more plants caused the development of new structures called cecal valves."20
Kimberly Johnson's National Geographic story three days later ran the deck "The changes should take millions of years-but these creatures are doing it in mere decades," and described the valves as "muscles between the large and small intestine," which they are not.21 To its credit, it is the one popular piece found that reports the open question. It quotes Andrew Hendry of McGill University, who was not involved: "All of this might be evolution," and "The logical next step would be to confirm the genetic basis for these changes."21
P. Z. Myers, on his blog Pharyngula, headed his post "Still just a lizard" and answered the expected creationist reply before it came: "yes, this population of Podarcis sicula is still made up of lizards, but they’re a different kind of lizard now. Evolution works."22 The post does not mention plasticity or the untested genetics.
Richard Dawkins gave the lizards a section of chapter 5 of The Greatest Show on Earth in 2009, "The lizards of Pod Mrcaru." He wrote that "those valves have actually started to evolve in the population of P. sicula on Pod Mrcaru," and called the story "evolution before our very eyes."23 He did not mention the untested genetic basis. He also dated the return visit to 2008, when it ran from 2004 to 2006, and he counted generations differently from the paper: "The average generation turnover of those lizards is about two years, so the evolutionary change observed on Pod Mrcaru represents only about eighteen or nineteen generations."23 Dawkins wrote a year before the plasticity test was published. Nothing in the later literature has been folded into the popular account, and in 2026 Christoph Adami's book The Evolution of Biological Information was still presenting the valve as the evolution of a new organ, according to a review of it by Winston Ewert.24 Adami's book was not read for this article.
Jerry Coyne's blog was searched for the case and returned no post on it.
The creationist and design replies
The young-earth and design replies mostly argued, before any test had been run, that the capacity was already there and the diet switched it on.
Answers in Genesis, 2008. Jean Lightner, a veterinarian writing for AiG, called the study "an excellent test—and an astounding confirmation—of the creation model."25 Her account of the paper is accurate. She reports the diet figures correctly, notes that "only mitochondrial DNA was sequenced," and says the authors themselves call for work on plasticity and maternal effects. On the valves she wrote that the structure "is present in other herbivorous lizards in this family (Lacertidae)" and "may reappear if conditions change and it then becomes important to survival."25 She also wrote that "36 years is hardly enough time to suspect that a beneficial random mutation might have occurred."25 Part 6 prices that claim. It holds for any one named mutation and fails for the population as a whole.
Answers in Genesis, News to Know. An unsigned AiG piece quoted the anatomist David Menton. He said the lizards "did have the ability to digest plant material," that the valve "is simply an enlargement of muscles already present in the gut wall at this juncture," and that "if the lizards were returned to their original habitat, the cecal valve feature may dwindle as the lizards returned to an insectivorous diet."8 The anatomy is wrong in detail. Menton took "muscles" from National Geographic. Iverson describes colic valves as infoldings of the gut wall, and Wehrle's team describes them as folds of the lining.3 1 But the prediction came true in substance. A year later Vervust's twenty lizards on an insect diet had no valves. Menton predicted it for a return to the old island, and the test was a laboratory diet, but the result is the one he forecast.
Creation Ministries International, 2014. Calvin Smith quoted CMI's Don Batten: "It is significant that the cecal valve is present in other herbivorous lizards in this family (Lacertidae), so it is not surprising that this particular species has the ability to produce cecal valves under certain conditions."26 Batten's point comes straight from Herrel's paper and is sound. Smith's own summary overstates twice. The lizards did not go "from predominantly carnivorous to vegetarian"; plants were a third to three-fifths of their food. And the inference that "the DNA sequence of the newer lizards seemed identical to the parent population" rests on two mitochondrial fragments, which say nothing about the nuclear genes that would build a valve.26
Evolution News, 2026. Winston Ewert argued that "The invading species already had the information necessary to produce the valve," switched off in insect eaters and on in plant eaters, whether "due to a genetic mutation, an epigenetic marker, or environmental triggers."24 That conclusion fits the later evidence well. His facts are loose. He says the lizards' "genomes" were sequenced, calls the valve "a muscle," and says "many other members of the same family have this structure," where Herrel says it is "rare in this family."24 He does not cite Vervust, Wehrle or Sabolić, the three papers that most support him.
Uncommon Descent, 2008. A writer using the name PaV used Motoo Kimura's substitution formula to argue that the change "cannot even begin to be explained using the Darwinian account."27 The calculation fails on its own terms. It starts from "12" founders, when there were ten. It asserts "we know from the paper that many more fixed genes were involved," when the paper reports no nuclear genes at all. It treats years as generations. And its arithmetic drops the factor of 4 in its own formula. Solving the formula as written gives 2.3, not 9.3, which is still impossible, because the deeper error is the model. Kimura's formula prices new mutations at one site. A population founded by ten wild animals starts with twenty copies of every gene and the variation among them, and that standing variation, not new mutation, is the ordinary route to fast change.28
Institute for Creation Research. No ICR article on Pod Mrčaru was found. ICR's model of continuous environmental tracking, in which organisms sense conditions and adjust themselves, is the natural home for this case, and the stickleback and speciation articles discuss it. See Sticklebacks and The Creation Model of Rapid Speciation.
How the claims score against the papers
The creationist writers read the 2008 paper more carefully than the press did. Lightner and AiG both reported that the genetic basis was untested, and the press release and Dawkins did not. Their central forecast, that the valves reflect a capacity switched on by diet, has so far held up better than the popular claim of a new organ evolved. Their weaknesses are loose anatomy, overstatement of what identical mitochondrial DNA shows, and a habit of not citing the follow-up papers that help them.
Part 5: How typical is this?
Fast change after animals reach a new place is common. The useful question for each case is the one Pod Mrčaru has not yet answered: was the difference still there when animals from both places were raised side by side?
Bahamian anoles, limb length. Brown anoles, Anolis sagrei, were introduced onto small Bahamian islands from a nearby source. Jonathan Losos, Kenneth Warheit and Thomas Schoener reported in 1997 that the populations "differentiated from each other rapidly over a 10–14-year period," and that "The more different the recipient island's vegetation from that of the source, the greater the magnitude of differentiation."29 Losos's own group then tested plasticity. They raised hatchlings in cages with only broad or only narrow perches, and "lizards in the broad treatment had relatively longer hindlimbs than lizards in the narrow treatment." They concluded that "not only is hindlimb length a plastic trait in these lizards, but that this plasticity leads to the production of phenotypes appropriate to particular environments."10 They added that the plastic range "is limited compared to the diversity shown throughout the anole radiation."10 A later experiment founded seven islands with one male and one female each and found that founder effects "persisted throughout the course of the experiment despite all populations adapting in the predicted direction-shorter hindlimbs."30 Common garden: the limb difference is partly plastic. This is the closest parallel to Pod Mrčaru.
Florida anoles, toepads. When brown anoles reached small islands in Florida in the mid-1990s, the native green anole, Anolis carolinensis, moved higher in the trees. Stuart, Losos and colleagues found that it "adaptively evolved larger toepads after only 20 generations," at 0.091 standard deviations per generation for toepad area and 0.077 for the number of scales on the toe.9 They took pregnant females from four invaded and four uninvaded islands, hatched the eggs in the laboratory and raised the young identically. "The effect of A. sagrei invasion on A. carolinensis toepad characteristics persisted in the common garden," which suggests a genetic difference, "though we cannot rule out trans-generational plasticity."9 Common garden: the difference persisted. This is what a demonstrated case looks like.
Cane toads, legs and speed. Cane toads were introduced to Australia about seventy years before Phillips and colleagues reported in 2006 that the invasion front had sped up about fivefold, and that "toads with longer legs can not only move faster and are the first to arrive in new areas, but also that those at the front have longer legs than toads in older (long-established) populations."31 That 2006 paper did not test inheritance. A 2016 study by Hudson, Shine and colleagues reports that the front-line toads' "offspring inherit that rapid dispersal rate," and that their skeletons have changed shape.32 Common garden: dispersal was shown to be inherited in later work; the leg-length difference itself was not re-read here.
Trinidad guppies, life history. David Reznick, Heather Bryga and John Endler moved a natural population of guppies from a place where predators eat adults to one where they eat the young. They reported in 1990 that this "resulted in significant life-history evolution in the predicted direction after 11 years, or 30–60 generations," and that such population differences "are heritable."33 In 1997 Reznick's team put the rate "up to seven orders of magnitude greater than rates inferred from the paleontological record," and found that males changed faster "largely because males had more genetic variation upon which natural selection could act."34 Common garden: heritability is stated in the abstracts; the rearing design was not read for this article. The selection coefficients for the guppy case are discussed in Haldane's Dilemma.
Sticklebacks and peppered moths. The two best-studied cases in this series show the same pattern. Freshwater sticklebacks lose armour in a few decades by reusing an allele already present at low frequency in the sea, and Barrett's pond experiment followed that allele through the generations. The peppered moth's dark form is caused by an inherited insertion of mobile DNA. Both are inherited changes from existing variation. See Sticklebacks and Peppered Moths. Blind cave fish show both at once: some cave traits appear within one generation by plasticity, and eye loss is inherited. See Blind Cave Fish.
| Case | Years watched | Generations | Trait | Tested in a common garden? | Result |
|---|---|---|---|---|---|
| Pod Mrčaru lizards | about 35 | 18 to 30 | Head, bite | Yes | Did not persist |
| Pod Mrčaru lizards | about 35 | 18 to 30 | Cecal valves | Reverse test only | Lost on an insect diet |
| Bahamian brown anoles | 10 to 14 | not given in the abstract | Hindlimb length | Yes, as rearing on perches | Partly plastic |
| Florida green anoles | about 15 | about 20 | Toepads | Yes | Persisted |
| Cane toads | about 70 | not given in the abstract | Legs, dispersal | Dispersal, in later work | Inherited |
| Trinidad guppies | 11 | 30 to 60 | Age and size at maturity | Stated heritable | Inherited |
Pod Mrčaru is typical in how fast it changed and unusual in how little of the change has been shown to be inherited. The field has known since at least 2000 that a fast change in a lizard's body can be partly plastic, and the one case in this group that passed a common garden, the Florida toepads, is the one that should be quoted as observed inherited change in a lizard.
Part 6: Scoring and the ledger
The score
Measuring Biological Information scores a change by what happened to a coded element: gain, loss, modification, or transfer of function. It also says that "a change that has not been shown to help the organism is not scored at all."35 Every class describes a change in DNA. None of them has been shown here.
| Change | What is known | Score |
|---|---|---|
| Cecal valves | Present in most Pod Mrčaru lizards, absent in Pod Kopište, lost in 20 lizards on an insect diet, not tested in a common garden | Not yet scoreable |
| Head size and bite force | Differences did not persist in a common garden | Not scored: mainly plastic |
| Plant diet, density, lost territories | Behaviour and ecology | Not a genetic change |
| Gut microbes | Differ with diet, island and population of origin | Not in the lizard's genome |
| A handful of outlier loci | Six intronic, one upstream of a gene; no trait or benefit tied to any | Not scored: no benefit shown |
"Not yet scoreable" cuts both ways. The case cannot be cited as the observed evolution of a new structure, because no inherited change has been shown, and the one experiment on the valves showed them disappearing when the diet changed. But it cannot be dismissed as nothing either. A fold of gut wall that forms when a lizard eats leaves, slows food, houses fermenting microbes, and goes away when the leaves stop is a working, conditional piece of engineering. The capacity to build it is real, it was there before 1971, and it needs an explanation of its own. Mainstream biology explains such capacities as the product of older evolution, pointing to valves arising again and again in plant-eating lizards. The design view explains them as built-in flexibility. Pod Mrčaru does not decide between those two, because it shows the capacity being used, not being made.
The ledger
The series asks how many mutations a population had the chance to try, and what it bought. For Pod Mrčaru the inputs are these.
| Input | Value | Source |
|---|---|---|
| Founders | 10 | Herrel 2008 |
| Years to sampling | 33 to 35 | 1971 to 2004–2006 |
| Generations | 18 to 30 | Dawkins; Herrel 2008 |
| Lizards now | about 900 to 9,200 | 3,082 per hectare on 0.29 or 3 hectares |
| Mutation rate | 1.17 × 10-8 per site per generation | Bergeron 2023, reptile mean |
| Genome | 1.5 to 1.6 billion bases | Sherpa 2023 |
| New mutations per hatchling | about 35 to 37 | 2 × rate × genome |
| Hatchlings, at most | about 16,000 to 280,000 | lizards now × generations |
| New mutations in the population, at most | about 600,000 to 10 million | hatchlings × 35 to 37 |
| Chance that one named single-letter change arose even once | about 0.01% to 0.2% | hatchlings × 2 × rate ÷ 3 |
| Inherited adaptive changes documented | none | Parts 3 and 6 |
| Unique functional sequence per lizard-generation | empty | no genetic change identified |
The working is in the note.36 The totals are upper bounds, because the population grew from ten and was smaller for most of the period.
Two things follow. First, Lightner's claim that 36 years "is hardly enough time to suspect that a beneficial random mutation might have occurred" is half right. Any one named mutation very probably never arose on the islet. But the population as a whole produced something like a million new mutations, and some of those may have been useful. More important, fast change in a founded population does not need new mutations. The ten founders brought twenty copies of every gene, with whatever variation Pod Kopište held, and the stickleback case shows that such standing variation is the usual raw material. Second, the empty last cell is not a zero. It is empty because nothing has yet been tied to a gene. If the valves turn out to be inherited, the cell will have to be filled, and the ledger is a lower bound on what the population produced, which is the direction that would cut against the young-earth side. For how a per-lineage count like this one is used across the series, see Benchmarking Evolution.
What the young-earth model wants from this case
Young-earth models need animals to change fast within a created kind, because hundreds of species must come from a few pairs within a few thousand years. See The Creation Model of Rapid Speciation. Pod Mrčaru shows that a lizard population can transform its diet, head, gut, numbers and behaviour within a few decades. That fits the need for speed.
Which mechanism serves the model better is less simple than it looks. Plasticity fits the claim that creatures were made able to adapt, and the creationist writers who predicted it have been borne out so far. But plasticity by itself is not inherited. A plastic change vanishes when the conditions change, as the valves did in Antwerp. Diversification into lasting species after the Flood needs inherited differences that stay put. For that, the model needs change from built-in genetic variation, the stickleback pattern, and Pod Mrčaru has not yet shown any. So the case supports the claim of built-in flexibility and is neutral on the claim of rapid inherited diversification. Mainstream biology has its own route from the first to the second, in which plasticity leads and genetic change follows. Pod Mrčaru, at about 35 years, may be too young to have reached the second step. That is the question the next common garden will answer.
Objections and responses
"The valves were in a hatchling with its umbilical scar. That proves they are inborn." It shows they can form early, not that they are coded differently. A hatchling on Pod Mrčaru comes from a Pod Mrčaru mother, and what she ate while making the egg may matter. Herrel's own paper names maternal effects as a possibility that needed testing.2 Only hatchlings from Pod Kopište eggs raised on plants, or Pod Mrčaru eggs raised on insects, can separate the causes.
"Plasticity is itself evolved, so this is still evolution." Plasticity can evolve, and mainstream biology holds that the ability to grow valves on a plant diet arose in lizard ancestors long ago. But that ancient event was not watched on Pod Mrčaru. What was watched is a population using a capacity it already had. The series counts what was observed, and an old capacity switched on is not an observed origin.
"Genome scans found real genetic change, so something evolved." Allele frequencies did change, mostly by drift after the founding, which is evolution in the population-genetic sense and is not disputed here. Sherpa's team found "just a handful of loci" that stood out and could not tie any to a trait.7 Change in allele frequency is not the same as a new function, and the scoring rules do not score a change with no shown benefit.
"Vervust's lizards were adults in a laboratory. A lab diet for 15 weeks is not an island." True. The result shows that the valves can disappear, not what would happen over generations. It is one experiment, in one direction, on 20 animals, with a discrepancy between 15 weeks in the text and 6 months in the table notes.5 It is still the only experiment, and its authors and the later Wehrle study both read the valves as flexible.
"Sabolić's common garden did not test the valves, so the valves may still be genetic." Correct as far as the abstract shows. This article therefore scores the valves "not yet scoreable," not "plastic." If the full paper or a new study shows valves forming in Pod Mrčaru lizards raised on insects, the verdict here changes.
"Creationists just say 'it was already there' about everything." In this case they said it in 2008 and 2009, before the first test, and made a checkable forecast: Menton's "the cecal valve feature may dwindle" on an insect diet.8 The next year's experiment came out his way. The claim was testable, and it passed its first test.
"The creationist writers get the anatomy and genetics wrong." Several do. Menton and Ewert call the valve a muscle, Ewert says genomes were sequenced in 2008, Smith reads identical mitochondrial DNA as showing the nuclear information was already present, and PaV's arithmetic is wrong. Those errors are real, and a reader should not take their facts on trust. Their central forecast survives them.
"Other introductions did show inherited change, so this one probably will too." Florida's green anoles kept their larger toepads in a common garden.9 Bahamian brown anoles' legs turned out partly plastic.10 Pod Mrčaru's heads and bites turned out mainly plastic.6 The record in lizards is mixed, so no forecast is safe.
Issues and unanswered questions
- The full text of Sabolić et al. 2024 was not read. Whether the common garden included gut dissection is the most important open fact in this article. The paper is paywalled, and the 2021 Zagreb thesis sits behind a robot check that this project's rules do not allow passing on a non-account site. Help from Eric, or an email to the authors, would settle it.
- Herrel, Vanhooydonck and Van Damme 2004 was read only in abstract. It is the source for valves in other lacertids and for their absence in other P. siculus. Which lacertids, and how many animals, is not known here. The same holds for Sagonas 2015, the Lacerta trilineata valve counts.
- No report of valves in Gallotia, Podarcis lilfordi or P. pityusensis was found. That is a gap in the search, not a finding.
- Nobody has tested the forward direction: whether Pod Kopište lizards fed plants grow valves.
- Generation time. Herrel's about 30 generations and Dawkins's 18 to 19 imply generation times of 1.2 and 2 years. No study found gives the age at first breeding on these islets.
- Island areas differ tenfold between Herrel 2008 and Taverne 2019, and the rate figure in Herrel's text, 8,593 darwins, is not in his table. Nevo 1972's full text, which might give the original areas and counts, was not read.
- The reciprocal half of Nevo's 1971 experiment, P. melisellensis taken to Pod Kopište, is announced in the 1972 summary. No later paper found says what became of it.
- Gene flow. One Pod Mrčaru lizard carried ancestry from Sušac.7 If lizards from Sušac, which also eat plants, have reached Pod Mrčaru, some of the difference could be imported rather than produced.
Verdict
Pod Mrčaru is a real and remarkable case of fast change after an introduction: ten founders, about 35 years, a new diet, bigger heads, gut valves, new microbes, five times the density and no territories. Every one of those observations stands.
What has not been shown is that any of it is inherited. The discoverers said the genetic basis was untested. The only experiment on the valves found them gone after lizards ate insects for 15 weeks. A later study by the same group found the valves "not fixed features." A common garden found that the head and bite differences did not persist. A genome scan found a handful of outlying loci and tied none to a trait. So the case scores as not yet scoreable. It cannot be cited as the observed evolution of a new organ. It cannot be waved away either, because a gut structure that appears when needed and goes away when it is not is a capacity that needs explaining. The honest summary is that Pod Mrčaru shows how much a lizard can change without new genes, and has not yet shown how much it can change with them.
Sources
- 1.
Wehrle, Beck A. et al. “Rapid dietary shift in Podarcis siculus resulted in localized changes in gut function.” Physiological and Biochemical Zoology 93(5):396–415. 2020. Read from the authors' laboratory copy of the printed PDF. Mirror: Hal.science
- 1a.
Page 396: “Pod Mrčaru lizards had higher digestibility of herbivorous and omnivorous diets fed over several weeks in the laboratory than did their source population.” ↩
- 1b.
Page 409: “It should be noted that we attempted to measure cellobiohydrolase activities in P. siculus, but we detected no activity anywhere in their guts.” ↩
- 1c.
Page 409: “For whatever reason, the Pod Mrčaru lizards we examined histologically for HG valves in summer 2013 (n = 3) did not have clear epithelial folds (valves) in their HGs.” “This stands in contrast to all of the other Pod Mrčaru lizards (n = 28, spanning four seasons over three additional years) we examined, which had clearly identifiable folds in the proximal portion to midportion of their HG, dividing the gut in the transverse plane (fig. S4; Wehrle 2018). The Pod Kopište lizards (n = 29) lacked these valves entirely.” ↩ ↩
- 1d.
Page 409: “convince us that, although usually present, the HG valves in P. siculus may be flexible in their morphology or even in their presence, and”. Page 410: “they are not fixed features in the Pod Mrčaru lizards.” ↩ ↩ ↩
- 1e.
Page 412: “We currently do not know whether the observed changes represent flexibility (Piersma and Drent 2003) or evolved differences with underlying genetic changes (Garland and Adolph 1991).” ↩
- 1f.
Page 410: “It is possible that a lifetime of acclimation to their respective wild diets has influenced digestive performance, and this may be mediated through the microbiome of the HG (i.e., not necessarily an evolved difference in the lizards themselves; Garland and Adolph 1991).” ↩
- 1a.
- 2.
Herrel, Anthony et al. “Rapid large-scale evolutionary divergence in morphology and performance associated with exploitation of a different dietary resource.” Proceedings of the National Academy of Sciences 105(12):4792–4795. Mar 25, 2008. Read from the publisher PDF in the Wayback Machine. Page numbers checked against the printed footers. Mirror: PMC ↩
- 2a.
Page 4792: “In 1971 five adult pairs of this species were moved from the small islet of Pod Kopište (0.09 km2) to the nearby Pod Mrčaru (0.03 km2) by Nevo and coworkers (12).” ↩ ↩ ↩
- 2b.
Page 4792: “repeated visits (twice yearly over the past three years, beginning in 2004) show that this species has become extinct on Pod Mrčaru.” ↩
- 2c.
Page 4792: “Genetic mitochondrial DNA analyses indicate that the lizards currently on Pod Mrčaru are indeed P. sicula and are genetically indistinguishable from lizards from the source population”. Page 4794: Methods: 12S and 16S rDNA from 8 Pod Kopište and 7 Pod Mrčaru lizards. ↩
- 2d.
Page 4793: “up to 8,593 darwins or 0.049 haldanes [Table 1; note, however, that these are synchronic rates (1) and assume no additional colonization of the island by P. sicula].” ↩ ↩
- 2e.
Page 4792: “plants composing between 34% (spring) and 61% (summer) of the total volume of the food eaten”. “In contrast, plant consumption was low (7% to 4%) and did not differ seasonally for lizards from the source population”. Page 4795: Methods: 330 lizards stomach-flushed, 119 from Pod Kopište and 211 from Pod Mrčaru. ↩ ↩
- 2f.
Page 4792: “of the plant matter eaten year round by lizards from Pod Mrčaru consists of items with high cellulose content such as leaves and stems”. The sentence begins with about 50%. ↩
- 2g.
Page 4793, table 1: Bite force, newtons, mean and standard deviation: Pod Kopište males 9.85 ± 2.29, females 4.99 ± 1.61; Pod Mrčaru males 11.25 ± 2.42, females 6.70 ± 1.90. Darwins, males/females: body mass 6,384/9,441, bite force 3,690/8,170. Haldanes: head width 0.046/0.051. Plants, % of diet: Pod Kopište 9 ± 7 and 4 ± 5, Pod Mrčaru 60 ± 34 and 48 ± 23. ↩ ↩
- 2h.
Page 4793: “Morphological analysis of preserved specimens shows the presence of cecal valves (Fig. 4) in all individuals, including a hatchling (26.4-mm snout-vent length, umbilical scar present) and a very young juvenile (33.11-mm snout-vent length) examined from Pod Mrčaru.” ↩
- 2i.
Page 4793: “Indeed, in the lizards from Pod Mrčaru, nematodes were common in the hindgut but absent from individuals from Pod Kopište.” ↩
- 2j.
Page 4794: “(0.01 versus 0.05 lizards per trap per hour, caught in unbaited traps, on Pod Kopiste and Pod Mrčaru, respectively)”. “lizards on Pod Mrčaru do no longer appear to defend territories.” ↩ ↩
- 2k.
Page 4793: “are not found in other populations of P. sicula (13) or in P. melisellensis.” Reference 13 is Herrel, Vanhooydonck and Van Damme 2004, Journal of Evolutionary Biology 17:974–984. ↩
- 2l.
Page 4793: “Our data show that in only 36 years (≈30 generations) the experimental introduction of a small propagule of lizards (five males and five females) into a novel environment has resulted in large differences in external morphology”. ↩ ↩
- 2m.
Page 4793: “The fact that <1% of all currently known species of squamates have cecal valves (13, 14) illustrates the unusual nature of these structures in this population.” References 13 and 14 are Herrel, Vanhooydonck and Van Damme 2004 and Iverson 1980. The less-than sign is a mis-encoded glyph in the PDF text layer. ↩
- 2n.
Page 4793: “These valves are similar in overall appearance and structure to those found in herbivorous lacertid, agamid, and iguanid lizards (13, 14)”. ↩
- 2o.
Page 4794: “a structure previously unreported for this species and rare in this family and scleroglossan lizards in general (13, 14, 18).” ↩
- 2p.
Page 4794: “Although the presence of cecal valves and large heads in hatchlings and juveniles suggests a genetic basis for these differences, further studies investigating the potential role of phenotypic plasticity and/or maternal effects in the divergence between populations are needed.” ↩ ↩ ↩
- 2a.
- 3.
Iverson, J. B. “Colic modifications in iguanine lizards.” Journal of Morphology 163(1):79–93. 1980. Abstract only, read from Europe PMC. Full text not obtained.
- 3a.
Section Abstract: “All lizard species of the subfamily Iguaninae except Amblyrynchus cristatus possess from one to eleven transverse valves in the proximal colon.” “Colic valves in iguanine lizards apparently evolved as simple infoldings of the medical colic wall.” ↩ ↩ ↩
- 3b.
Section Abstract: “Herbivorous species of the Scincidae, Agamidae, and Iguanidae are the only lizards known to exhibit colic partitioning, suggesting that the evolution of these structures is intimately related to the evolution of herbivory in these lizards.” ↩ ↩
- 3a.
- 4.
Sagonas, K. et al. “Effects of insularity on digestion: living on islands induces shifts in physiological and morphological traits in island reptiles.” The Science of Nature 102:55. 2015. Abstract only, read from Europe PMC. Full text not obtained.
- 5.
Vervust, Bart et al. “Anatomical and physiological changes associated with a recent dietary shift in the lizard Podarcis sicula.” Physiological and Biochemical Zoology 83(4):632–642. 2010. Read online at JSTOR on a free account. Mirror: Jstor.org
- 5a.
Page 632: “In addition to the differences in skull dimensions and the prevalence of cecal valves previously reported, these two recently diverged populations also differ in aspects of their dentition (teeth width) and the lengths of the stomach and small intestine.” ↩
- 5b.
Page 635: “In 12 of the 16 individuals (75%) examined from Pod Mrčaru, the hindgut contained cecal valves. In sharp contrast, we found no such valves in any of the 27 individuals from Pod Kopište”. Page 633: Fieldwork in April 2008, with measurements in a field laboratory on Lastovo. ↩
- 5c.
Page 634: “The same measurements and observations on gut morphology were carried out on eight males and 12 females of the Pod Mrčaru population after they had been fed an exclusively arthropod diet for 15 wk. These 20 lizards had died of oxygen deficiency following a power cut in the animal house at Antwerp University. None of the animals were killed for the purpose of this study.” ↩
- 5d.
Page 635: “Lizards maintained on an arthropod diet showed a reduction in the weight of the small intestine and the hindgut. In addition, we found no evidence of cecal valves in any of the twenty specimens that had been fed arthropods. This was in sharp contrast to the high proportion (12/16) of individuals with valves among the freshly caught individuals”. ↩ ↩ ↩
- 5e.
Page 637: “Many comparative studies have tacitly assumed that the distinctive features of plant-eating lizards (large body size, skull dimensions, special dentition, gut morphology) are a product of genetic adaptation to the special demands of a plant-based diet (e.g., Van Damme 1999; Cooper and Vitt 2002; Espinoza et al. 2004; Herrel et al. 2008). Our results suggest that in P. sicula, at least some of the changes associated with a dietary shift toward a higher proportion of plant material may be plastic.” ↩
- 5f.
Page 638, table 2: “individuals that were fed an exclusively arthropod diet for 6 mo”. Page 639, table 3: The Table 3 title also says 6 months. The methods on page 634 and the discussion on page 640 say 15 weeks. ↩ ↩
- 5g.
Page 640: “Although parts of their gastroinstestinal systems were still better developed than those of specimens feeding mainly on arthropods in the wild, it seems likely that a prolonged exposure to an animal-based diet would have erased even those differences.” The misspelling is the paper's. ↩
- 5a.
- 6.
Sabolić, I. et al. “Plastic and genomic change of a newly established lizard population following a founder event.” Molecular Ecology 33(2):e17255. 2024. Abstract only, read from Europe PMC. The full text is paywalled, and the author's 2021 University of Zagreb thesis on the same work was not reachable past the repository's robot check. ↩
- 6a.
Section Abstract: “Diverged traits related to head size and shape or bite force showed moderate heritability in a crossing experiment, but between-population differences in these traits did not persist in a common garden environment. Our results confirm the existence of sufficient additive genetic variance for traits to evolve under selection while also demonstrating that phenotypic plasticity and/or genotype by environment interactions are the main drivers of population differentiation at this early evolutionary stage.” ↩ ↩ ↩
- 6a.
- 7.
Sherpa, Stéphanie et al. “Genetic depletion does not prevent rapid evolution in island-introduced lizards.” Ecology and Evolution 13(11):e10721. 2023. Open access, CC BY. Mirror: PMC
- 7a.
Section 2.5: “we were unable to perform robust estimations of past and contemporary effective population size, as both the sample size (between 14 and 16 gene copies) and the short time‐scale at which the demographic events occurred (~30 generations) limit the use of both linkage and coalescent approaches”. ↩
- 7b.
Section Abstract: “Using a genomic approach (~82,000 ddRAD loci), we confirmed a founder effect during introduction and very low neutral genetic differentiation between the introduced population and its source.” “Our genome‐scan analysis identified just a handful of loci showing large allelic shifts between ecologically divergent populations. This low overall signal of selection suggests that the extreme phenotypic differences observed among populations are determined by a small number of large‐effect loci and/or that phenotypic plasticity plays a major role in phenotypic changes.” ↩ ↩ ↩ ↩
- 7c.
Section 3.4: “The seven remaining loci were all functionally annotated and included six loci in intronic regions, and one locus was an upstream variant (~800 bp).” 15 of the 29 outlier loci aligned to the genome of the common wall lizard, Podarcis muralis; seven were intergenic and one lay downstream of an unannotated gene. ↩
- 7d.
Section 3: “A preliminary ADMIXTURE analysis using this dataset revealed one highly admixed individual in Pod Mrčaru, potentially indicating very recent gene flow with Sušac (Figure 2A ).” ↩ ↩
- 7e.
Section 4.3: “the genome size of Podarcis lizards is 1.5–1.6 Gb (Andrade et al., 2019 ; Gabrielli et al., 2023 )”. ↩
- 7a.
- 8.
Answers in Genesis. “Island evolves lizards.” Answers in Genesis, News to Know. Jun 6, 2009. Retrieved Sep 29, 2026. Unsigned. The page date is 2009; the story it answers ran in April 2008.
- 8a.
“But Answers in Genesis anatomist David Menton , after reviewing the research, noted that the original lizards did have the ability to digest plant material; they simply preferred insects for roughly 95 percent of their dietary needs.” “Furthermore, Menton adds, “The ‘new’ muscular valve they found between the small and large intestine is simply an enlargement of muscles already present in the gut wall at this juncture.”” “Menton also suggests that if the lizards were returned to their original habitat, the cecal valve feature may dwindle as the lizards returned to an insectivorous diet.” ↩ ↩ ↩
- 8a.
- 9.
Stuart, Y. E. et al. “Rapid evolution of a native species following invasion by a congener.” Science 346(6208):463–466. 2014. Read from the authors' manuscript in Harvard's repository. Locators are the manuscript's line numbers. Mirror: Dash.harvard.edu
- 9a.
Lines 19–22: “On small islands in Florida, we found that the lizard Anolis carolinensis moved to higher perches following invasion by Anolis sagrei and, in response, adaptively evolved larger toepads after only 20 generations.” Lines 79–80: “islands have diverged at mean rates of 0.091 (toepad area) and 0.077 (lamellae) standard deviations per generation”. ↩
- 9b.
Lines 86–90: “We took gravid A. carolinensis females from four invaded and four un-invaded islands in July 2011, collected their eggs in the lab, and raised the offspring in identical conditions (12). The effect of A. sagrei invasion on A. carolinensis toepad characteristics persisted in the common garden (Fig. 3; Table S4; [(12)]), suggesting genetically based divergence in nature (though we cannot rule out trans-generational plasticity).” ↩ ↩ ↩
- 9a.
- 10.
Losos, J. B. et al. “Evolutionary implications of phenotypic plasticity in the hindlimb of the lizard Anolis sagrei.” Evolution 54(1):301–305. 2000. Abstract only, read from Europe PMC.
- 10a.
Section Abstract: “We raised hatchling A. sagrei in terraria provided with only broad or only narrow surfaces. At the end of the experiment, lizards in the broad treatment had relatively longer hindlimbs than lizards in the narrow treatment. These results indicate that not only is hindlimb length a plastic trait in these lizards, but that this plasticity leads to the production of phenotypes appropriate to particular environments.” ↩ ↩ ↩
- 10b.
Section Abstract: “Comparison to hindlimb lengths of other Anolis species indicates that the range of plasticity is limited compared to the diversity shown throughout the anole radiation.” ↩
- 10a.
- 11.
Nevo, Eviatar et al. “Competitive exclusion between insular Lacerta species (Sauria, Lacertidae).” Oecologia 10(2):183–190. 1972. Summary only, read from the publisher's preview page. Full text not obtained.
- 11a.
Section Summary: “Competetive exclusion between Lacerta sicula and L. melisellensis characterizes the small islands of the Adriatic Sea.” “On the third island, the introduced species appears to be replacing the native form.” “A reciprocal introduction involving the two species on the islands of Pod Kopiŝte and Pod Mrĉaru is announced.” Spelling and diacritics as printed on the publisher's page. ↩ ↩
- 11a.
- 12.
Taverne, Maxime et al. “Diet variability among insular populations of Podarcis lizards reveals diverse strategies to face resource-limited environments.” Ecology and Evolution 9(22):12408–12420. 2019. Open access, CC BY 4.0. Read from the HAL copy. Mirror: Hal.science
- 12a.
Page 12411, table 1: Area in square metres: Pod Kopište 7,915; Pod Mrčaru 2,931. ↩ ↩
- 12b.
Page 12415: “Plant consumption is highly variable among islands, ranging from strictly insectivorous populations (e.g., Brusnik or Mala Palagruža) to almost completely herbivorous populations (e.g., Sušac or Pod Mrčaru).” “Insular populations of P. sicula might also shift to a more omnivorous diet, as is seen on the islands Sušac, Pijavica and Pod Mrčaru, where populations include an important proportion of plant items into their diets.” ↩
- 12a.
- 13.
Lemieux-Labonté, Virginie et al. “Proximate drivers of population-level lizard gut microbial diversity: impacts of diet, insularity, and local environment.” Microorganisms 10(8):1550. 2022. Open access, CC BY. Mirror: PMC
- 13a.
Section 2.1: “estimates based on transects range from 3082 lizards per hectare on Pod Mrčaru to 1045 lizards per hectare on Pod Kopište”. The paper cites Vervust, Van Dongen, Grbac and Van Damme 2009, Functional Ecology 23:996–1003, for these figures. ↩ ↩
- 13b.
Section Abstract: “Alpha-diversity analyses showed that the microbial diversity of omnivorous lizards was higher than the microbial diversity of insectivorous lizards. Moreover, omnivorous individuals harbored significantly more Methanobrevibacter.” ↩
- 13a.
- 14.
Taverne, Maxime et al. “Form–function relationships underlie rapid dietary changes in a lizard.” Proceedings of the Royal Society B 290:20230582. 2023. Mirror: PMC
- 14a.
Section Abstract: “Our results show that small differences in shape, combined with variation in muscle architecture, can result in significant differences in performance allowing access to novel trophic resources.” ↩
- 14a.
- 15.
Pérez-Mellado, Valentin, and Claudia Corti. “Dietary adaptations and herbivory in lacertid lizards of the genus Podarcis from western Mediterranean islands (Reptilia: Sauria).” Bonner zoologische Beiträge 44(3–4):193–220. 1993. Read from the Biodiversity Heritage Library scan.
- 16.
Herrel, Anthony et al. “Omnivory in lacertid lizards: adaptive evolution or constraint?” Journal of Evolutionary Biology 17(5):974–984. 2004. Abstract only, read from Europe PMC. Full text not obtained.
- 16a.
Section Abstract: “Our data show that omnivorous lacertid lizards possess modifications of these traits that allow them to successfully exploit plant material as a food source.” ↩
- 16a.
- 17.
Starck, J. Matthias. “Shaping up: how vertebrates adjust their digestive system to changing environmental conditions.” Animal Biology 53(3):245–257. 2003. Abstract only, read from the Crossref record.
- 17a.
Section Abstract: “The vertebrate gastrointestinal tract is a flexible system that can individually be modified to account for changes in amount and quality of food as well as changes in internal demands.” ↩
- 17b.
Section Abstract: “Up-regulation of the mucosa size does not involve cell proliferation, thus it is presumably an energetically cheap process.” ↩
- 17a.
- 18.
Kohl, K. D. et al. “Physiological and microbial adjustments to diet quality permit facultative herbivory in an omnivorous lizard.” Journal of Experimental Biology 219(12):1903–1912. 2016. Abstract only, read from Europe PMC.
- 18a.
Section Abstract: “Additionally, lizards fed the plant-rich diet exhibited significantly longer small intestines and larger hindguts, demonstrating that gut morphology is phenotypically flexible.” ↩
- 18a.
- 19.
Reppa, Aikaterini et al. “Convergent digestive adaptation to resource limitation in an insular lizard across a microgeographic archipelago.” Journal of Animal Ecology 95. 2026. Open access, CC BY-NC-ND. Mirror: PMC
- 19a.
Section Abstract: “These results demonstrate repeated and independent adaptive responses to resource scarcity and highlight digestive plasticity as a key mechanism facilitating persistence in insular habitats.” ↩
- 19a.
- 20.
University of Massachusetts Amherst. “Lizards undergo rapid evolution after introduction to a new home.” ScienceDaily. Apr 18, 2008. Retrieved Sep 29, 2026. Press release, reprinted by ScienceDaily.
- 20a.
“Our data shows that evolution of novel structures can occur on extremely short time scales.” ↩
- 20b.
“Tail clips taken for DNA analysis confirmed that the Pod Mrcaru lizards were genetically identical to the source population on Pod Kopiste.” “Cecal valve evolution probably went hand-in-hand with a novel association between the lizards on Pod Mrcaru and microorganisms called nematodes that break down cellulose, which were found in their hindguts.” ↩
- 20c.
“Eating more plants caused the development of new structures called cecal valves, designed to slow the passage of food by creating fermentation chambers in the gut, where microbes can break down the difficult to digest portion of plants.” ↩
- 20a.
- 21.
Johnson, Kimberly. “Lizards rapidly evolve after introduction to island.” National Geographic News. Apr 21, 2008. Retrieved Sep 29, 2026.
- 21a.
“The changes should take millions of years-but these creatures are doing it in mere decades.” “Researchers found that the lizards developed cecal valves—muscles between the large and small intestine—that slowed down food digestion”. ↩
- 21b.
“What could be debated, however, is how those changes are interpreted—whether or not they had a genetic basis and not a "plastic response to the environment," said Hendry, who was not associated with the study.” ““All of this might be evolution,” Hendry said. “The logical next step would be to confirm the genetic basis for these changes.”” ↩
- 21a.
- 22.
Myers, P. Z. “Still just a lizard.” Pharyngula. Apr 23, 2008. Retrieved Sep 29, 2026.
- 22a.
Paragraph 1: “The title gets the principal objection of any creationist out of the way: yes, this population of Podarcis sicula is still made up of lizards, but they’re a different kind of lizard now. Evolution works.” ↩
- 22a.
- 23.
Dawkins, Richard. The Greatest Show on Earth: The Evidence for Evolution. Free Press. 2009. Read by archive.org search-inside on a lending-restricted copy. Page numbers from the item's page map, checked against the book's index entry "Pod Mrcaru, 113-16".
- 23a.
Chapter 5, page 116: “Now, the fascinating thing is that, although caecal valves don’t normally occur in Podarcis sicula and are rare in the family to which it belongs, those valves have actually started to evolve in the population of P. sicula on Pod Mrcaru, the population that has, for only the past thirty-seven years, been evolving towards herbivory.” “I should repeat that the only thing that is really exceptional about this whole story, and the reason I am telling it here, is that it all happened so extremely rapidly, in a matter of a few decades: evolution before our very eyes.” ↩
- 23b.
Page 113: “Then, in 2008, another group of mainly Belgian scientists, associated with Anthony Herrel, visited the islands”. Page 116: “The average generation turnover of those lizards is about two years, so the evolutionary change observed on Pod Mrcaru represents only about eighteen or nineteen generations.” ↩ ↩
- 23a.
- 24.
Ewert, Winston. “Evolution of biological information: lizards and a “new” organ.” Science and Culture Today. Aug 5, 2026. Retrieved Sep 29, 2026. Discovery Institute's news site, formerly Evolution News.
- 24a.
“Adami characterizes this as the evolution of a new organ.” ↩
- 24b.
“There is a more plausible explanation. The invading species already had the information necessary to produce the valve. However, it was turned off as unnecessary in an insect-eating population. As the population switched to a plant diet, it was turned on. This might be due to a genetic mutation, an epigenetic marker, or environmental triggers.” ↩
- 24c.
“However, when the genomes of these lizards were sequenced, they found them to be a match for Podarcis siculus .” “The cecal valve is a muscle that separates the intestine into compartments.” “Indeed, many other members of the same family have this structure.” ↩
- 24a.
- 25.
Lightner, Jean K. “Life: designed by God to adapt.” Answers in Genesis. Jun 4, 2008. Retrieved Sep 29, 2026.
- 25a.
Section Abstract: “The recent discovery of rapid changes in a lizard population has offered an excellent test—and an astounding confirmation—of the creation model.” ↩
- 25b.
Section A Change in the Digestive Tract: “The cecal valve is present in other herbivorous lizards in this family (Lacertidae). This suggests that certain structures important to the survival of an animal in one environment may not be retained in the population if they move to a different environment and the structure is no longer useful. However, the structure may reappear if conditions change and it then becomes important to survival.” ↩
- 25c.
Section Pondering the Source of the Changes: “However, only mitochondrial DNA was sequenced, which was identical in both populations of P. sicula . One wonders what the source of such a genetic change might be; 36 years is hardly enough time to suspect that a beneficial random mutation might have occurred.” ↩
- 25a.
- 26.
Smith, Calvin. “The good, the bad and the ugly ….” Creation Ministries International. Aug 27, 2014. Retrieved Sep 29, 2026.
- 26a.
“As CMI’s Dr Don Batten commented; “It is significant that the cecal valve is present in other herbivorous lizards in this family (Lacertidae), so it is not surprising that this particular species has the ability to produce cecal valves under certain conditions.”” ↩
- 26b.
“food preference (went from predominantly carnivorous to vegetarian)”. “Astonishingly the DNA sequence of the newer lizards seemed identical to the parent population, which means that the genetic information for cecal valves (along with the other genetic information accounting for the other changes) was likely already present in these creatures”. ↩
- 26a.
- 27.
PaV. “Kimura and the Adriatic lizards.” Uncommon Descent. May 8, 2008. Retrieved Sep 29, 2026. Pseudonymous author. The site was archived in April 2023.
- 27a.
“Well, the transplanted population size of the Adriatic lizards was 12.” “Let’s just assume that ONE mutant gene was fixed (although we know from the paper that many more fixed genes were involved), and let’s assume that it took the entire 36 years to become fixed.” “s1=1/36 x 200 x 1.5 x 10^-5 =10^5/36 x 300= 100,000/10,800=9.3.” “Thus, the simple math demonstrates that, using the Modern Synthesis, what was documented to have occurred with the transplanted lizard population cannot even begin to be explained using the Darwinian account.” ↩ ↩
- 27a.
- 28.
PaV's formula is Kimura's rate of substitution, K = 4 Ne s v, with K the substitutions per generation, Ne the effective population, s the selective advantage and v the mutation rate per gamete at the site. He sets K to one substitution in 36 and takes Ne = 200 and v = 1.5 × 10-5. Solving for s gives s = K / (4 Ne v) = (1/36) / (4 × 200 × 1.5 × 10-5) = 0.0278 / 0.012 = 2.3. His line "s1=1/36 x 200 x 1.5 x 10^-5" omits the 4 and gives 9.3. Using generations rather than years makes it worse: at 30 generations s = 2.8. The formula assumes the favoured allele arises by new mutation after the founding. It does not apply to variation the ten founders already carried.27 ↩
- 29.
Losos, Jonathan B. et al. “Adaptive differentiation following experimental island colonization in Anolis lizards.” Nature 387:70–73. 1997. Abstract only, read from the publisher's page.
- 29a.
Section Abstract: “Populations of the lizard Anolis sagrei , introduced onto small islands from a nearby source, differentiated from each other rapidly over a 10–14-year period. The more different the recipient island's vegetation from that of the source, the greater the magnitude of differentiation.” ↩
- 29a.
- 30.
Kolbe, J. J. et al. “Founder effects persist despite adaptive differentiation: a field experiment with lizards.” Science 335:1086–1089. 2012. Abstract only, read from Europe PMC.
- 30a.
Section Abstract: “These founding events generated significant among-island genetic and morphological differences that persisted throughout the course of the experiment despite all populations adapting in the predicted direction-shorter hindlimbs-in response to the narrower vegetation on the small islands.” ↩
- 30a.
- 31.
Phillips, B. L. et al. “Invasion and the evolution of speed in toads.” Nature 439:803. 2006. Abstract only, read from Europe PMC.
- 31a.
Section Abstract: “Here we show that the annual rate of progress of the toad invasion front has increased about fivefold since the toads first arrived; we find that toads with longer legs can not only move faster and are the first to arrive in new areas, but also that those at the front have longer legs than toads in older (long-established) populations.” ↩
- 31a.
- 32.
Hudson, C. M. et al. “Constructing an invasion machine: the rapid evolution of a dispersal-enhancing phenotype during the cane toad invasion of Australia.” PLoS ONE 11(9):e0156950. 2016. Open access. Abstract read. Mirror: PMC
- 32a.
Section Abstract: “As cane toads (Rhinella marina) have spread across tropical Australia over an 80-year period, their rate of invasion has increased from around 15 to 60 km per annum. Toads at the invasion front disperse much faster and further than conspecifics from range-core areas, and their offspring inherit that rapid dispersal rate.” ↩
- 32a.
- 33.
Reznick, David A. et al. “Experimentally induced life-history evolution in a natural population.” Nature 346:357–359. 1990. Abstract only, read from the publisher's page.
- 33a.
Section Abstract: “These differences are heritable, and correspond with theoretical predictions”. “This resulted in significant life-history evolution in the predicted direction after 11 years, or 30–60 generations.” ↩
- 33a.
- 34.
Reznick, D. N. et al. “Evaluation of the rate of evolution in natural populations of guppies (Poecilia reticulata).” Science 275:1934–1937. 1997. Abstract only, read from Europe PMC.
- 34a.
Section Abstract: “The resulting rate of evolution of age and size at maturity was similar to rates typically obtained for traits subjected to artificial selection in laboratory settings and up to seven orders of magnitude greater than rates inferred from the paleontological record. Male traits evolved more rapidly than female traits largely because males had more genetic variation upon which natural selection could act.” ↩
- 34a.
- 35.
The scoring procedure and its rule that an unhelpful change is not scored are set out in Part 2 of Measuring Biological Information, in the section "The scoring procedure". ↩
- 36.
Population now. Transect estimates give 3,082 lizards per hectare on Pod Mrčaru.13 The islet is 2,931 m², or 0.2931 ha, in Taverne's table, and 0.03 km², or 3 ha, in Herrel's paper.12 2 3,082 × 0.2931 = 903; 3,082 × 3 = 9,246. Generations. Herrel gives about 30; Dawkins, using a two-year generation, gives 18 to 19.2 23 Mutations per hatchling. The reptile mean rate is 1.17 × 10-8 per site per generation37 and the Podarcis genome is 1.5 to 1.6 × 109 bases.7 A hatchling receives two genomes, so it carries 2 × 1.17 × 10-8 × 1.5 × 109 = 35.1 to 2 × 1.17 × 10-8 × 1.6 × 109 = 37.4 new mutations. Hatchlings. An upper bound is the present population times the number of generations: 903 × 18 = 16,254 and 9,246 × 30 = 277,380. New mutations. 16,254 × 35.1 = 5.7 × 105; 277,380 × 37.4 = 1.04 × 107. One named change. A given site mutates in a hatchling with probability 2 × 1.17 × 10-8, and one of the three possible new letters is a third of that. The expected number of times one named change arose is 16,254 × 2 × 1.17 × 10-8 ÷ 3 = 1.3 × 10-4 to 277,380 × 2 × 1.17 × 10-8 ÷ 3 = 2.2 × 10-3, or about 0.01% to 0.2%. Cautions. The rate is an average of six reptile species, none of them a lacertid. The hatchling count ignores that the population started at ten. Both make the figures upper bounds. ↩
- 37.
Bergeron, Lucie A. et al. “Evolution of the germline mutation rate across vertebrates.” Nature 615:285–291. 2023. Open access, CC BY. Mirror: PMC
- 37a.
Page 286: “On average, mutation rates per generation are higher in reptiles (average of all species 1.17 × 10−8, 95% CI of the mean = 5.34 × 10−9 to 1.80 × 10−8)”. ↩
- 37a.