Appalachiosaurus, Arm Length, and Matrices

The Arms of Appalachiosaurus, or how I was again reminded to never trust a cladistic matrix without inspecting every cell...

This post wasn't meant to grow so large, or rambling. It started out simple enough, my dear friend, Dr. Ray Wilhite, sent me photos of the McWane Center tyrannosauroid Appalachiosaurus mount, along with images of the original bones, back in June of 2022. I took one look at those arms and said, "Whaaaaaa?", made the above post and moved on. Or so I thought... thus began a multi-hour journey that I captured below.

Appalachiosaurus montgomeriensis, holotype RMM 6670 [RMM = Red Mountain Museum, however the bones are at the McWane Center in Birmingham, AL], was named in 2005 by Carr, Williamson, and Schwimmer for the most complete "East Coast" tyrannosauroid yet found. Elements of a subadult individual from the Turnipseed Dinosaur Site in the Demopolis Formation of Alabama were discovered in 1982 in marine rocks, indicating it was a "bloat and float." Coming from the Eastern side of the Western Interior Seaway, far from the traditional Albertosaurus, Gorgosaurus, and their ilk, this ~78 million-year-old Campanian theropod rightly stirred up much excitement.

Carr et al. 2005 list eight caudal vertebrae (always start with the tail :-)), many of which have the neurocentral suture present which typically indicates an animal isn't done growing. From the pelvis is a shaft of a pubis (which will become a star of this long missive) and an ischium fragment. Phalanges (not sure how many), three metatarsals, astragalus, calcaneum, fibula, tibia, and femur mean we have a wonderfully complete hind-end to accompany the mostly complete skull. Awesome!

I am always curious to understand a fossil's history, how it came to be known what it is today, and what steps it took along the way. The Carr et al. 2005 paper provides a nice historical overview, but I opted to dig into each of the referenced papers therein as I never know what I'll find.

1986 Lamb

Lamb published an abstract that seems to reference these ones here: Alabama dinosaurs: Part I: Origins (Birmingham Paleontological Society), v, 7, no. 6, p. 1-3., however I have yet to see it (hoping my interlibrary loan request gets filled, if anyone has it please let me know, I'd love to read it!)

King et al. 1988 seem to be the first to mention this skeleton, which was found in 1982 by King in the Demoplis Chalk. They wrote:

The bones are from a single individual and include two tibiae, one fibula, six metatarsals, portions of the pelvic girdle, five ungual phalanges (claws), eight phalanges (toe bones) , two vertebrae, one maxilla, one premaxilla, one dentary,
thirteen teeth, several partial ribs, and several unidentified bones including skull components. The dinosaur bones are catalogued in the Auburn University Museum of Paleontology (AUMP 2366) and are being prepared at the Red Mountain Museum.

The bones formally described in 2005 by Carr et al. were listed in the opening section, and you will read about them again below.  You might note the list of bones from King et al. does not match the list that appears in Carr et al. 2005. James Lamb suggested (pers. comm. 2024) that in 2006 not all of the bones had been prepared, leading him to prepare additional elements. What isn't clear to me is if he did "finish prep" on existing elements (the Carr et al. photos sure look like prep was done!) or if he prepared elements that King et al. 1988 list but don't appear in Carr et al. 2005, like ribs, a second tibia, and additional foot material.

Of course, the second tibia may not have been a tibia, the mismatch in foot elements (King et al. 19 elements: 6 MT, 8 phalanges, 5 claws vs Carr et al. with 3 MT and 7 phalanges) could be because of misidentification. Carr's crew may not have desired to photograph or describe the ribs (if they were truly ribs). Carr's team notes many elements, like the astragalus and calcaneum, not listed in the 1988 list. This happens all the time as elements are prepared their true selves emerge.

1989 Lamb

Carr et al. 2005 note that Lamb informed a Birmingham newspaper in 1989 about the find. I was unable to find a clipping for this reference and they don't include it in their bibliography. I contacted James Lamb and he responded that Carr was told about it by the collections manager at McWane, who misunderstood Lamb's 1989 publication was academic in nature, an abstract in the Alabama Academy of Science, and did not appear in a newspaper article. I am still that abstract, but the best I can tell it was in 1986. There may have been a 1989 newspaper article on it, but my perusing through Alabama's online newspaper offerings from 1989 didn't reveal any (not a lot to choose from, though :-)). Anyone know of the article in question?

1989 Baird

While on a quest to find a suitable taxonomic identification for a fragmentary New Jersey theropod MT II, Baird mentions the animal that would become Appalachiosaurus. Here is what he wrote:

The one dinosaur with which our specimen shows closest affinities
is at present a nondescript [sic]: the partial skeleton of a smallish
carnosaur from the Demopolis Chalk (Late Campanian to
Early Maastrichtian) near Montgomery, Alabama. This
skeleton, for information about which I am greatly indebted to
James Lamb, is currently being prepared and studied at the Red
Mountain Museum in Birmingham (RMM 6670 = AUMP 2366
of Auburn University). Photographs supplied by Lamb show
that the second metatarsal of the Alabama dinosaur is slightly
larger than the New Jersey specimen and is closely similar, although
not identical, to it in configuration. Further discussion
of the Alabama carnosaur would be premature here, except to
observe that it is a long-limbed genus with what appear to be
familial resemblances to Albertosaurus.

Two items jump out at me, the first being he notes it is from the Demopolis Chalk, which will appear again in a bit. The second is he uses the phrase "long-limbed genus." What did Baird mean by long-limbed? Dr. James Lamb says Baird only saw the hind-limb material and was referring to that (Lamb, pers. comm. 2024).

1989 Schwimmer and Best

Schwimmer and Best published a paper on the dinosaur remains of Georgia. They wrote:

Recent discovery of a partial theropod skeleton in central Alabama confirms that Albertosaurus was present in the Southeast during the Late Cretaceous (J Lamb, pers. comm. and DRS pers. obs.). The left metatarsal IV from the undescribed Alabama specimen is also essentially identical to the Georgia specimen (Fig. 3.10, 3.11).

Alas, no mention of the formation the animal-to-become-Appalachiosaurus was excavated from was given. They do believe these southern tyrannosaur elements belonged to Albertosaurus, based on the similarity of their MT IV to numerous specimens of A. libratus, both MT IVs and a radius and ulna found near their Georgian MT.

1992 Carpenter

Carpenter twice briefly mentions the specimen in his 1992 review of tyrannosaurids. I'll quote them both.

1-In the overview of taxa section, he writes "Another Albertosaurus specimen has been collected from the marine Mooreville Chalk of Alabama by Auburn University." He doesn't cite the source of where this information came from.

2-Under the Albertosaurus libratus (Lambe) 1914 section, he notes it was found in the Morreville [sic] Chalk of Alabama, U.S.A. 

Carr et. al 2005 wrote, "Carpenter (1992) made mention of the specimen, then thought to be [emphasis mine] from the Mooreville Chalk...accepted its informal referral to Albertosaurus." 

However, Baird in 1989, and King et al. 1988, correctly noted it was from the Demopolis Chalk Formation, I'm not sure where the "...then thought to be..." originated from. I haven't found any additional sources that note the Mooreville Chalk Formation. I speculate Carpenter had a reason as the Ken I know is sharp as a tack and extremely careful and insightful.

 1993 Schwimmer et al.

Schwimmer et al. 1993 mention the Georgia MT IV again, this time calling it "Albertosaurus?" They go on to discuss the Alabama specimen in two places.

In the Introduction section they note, "Another partial theropod skeleton, not yet
formally described (see King et al., 1988; Baird, 1989), has been
found in upper Campanian age strata in central Alabama."

Under a section entitled Albertosaurus? sp. they expand upon that sentence:

As noted in the introduction, a partial theropod skeleton was
collected recently in central Alabama from the Demopolis Chalk
in Montgomery County (King et al., 1988, describe the sedimentary
environment of the site). This theropod is presently in
preparation in the Red Mountain Museum, Birmingham, and,
at late Campanian age, is slightly younger than the Blufftown
material. However, comparison of the left fourth metatarsals
from the Blufftown and Montgomery theropods shows they are
indistinguishable in size and overall morphology (James Lamb,
personal commun., and see Baird, 1989, p. 56).

2005 Carr et al.

Carr, Williamson, and Schwimmer provide useful historical references, as you can see from above, all of which were lifted from them :-). They note phalanges are present but they don't state how many, and the link to "Tables 14-16, on-line at http://www.vertpaleo.org/jvp/" doesn't work. How many other early 00s papers have broken links?  I've seen my fair share...

I surmise they figure them all in their FIGURE 20, as the remainder of the bones appear to be all figured (three metatarsals, astragalus, calcaneum, tibia, fibula, femur, ischium, pubis, eight caudal vertebrae, nearly complete skull) but I have no way to know for certain from the paper as there is no "backup" number provided in the text. The answer may be seven as that are how many with measurements that appeared once I was able to track down the aforementioned link. 

Speaking of the link, I logged into the JVP Online Access portal (membership has its privileges :-)) and entered the title of the paper in the search bar (ok, up through tyrannosauroid, it is a long title). A hyperlink appeared which took me to the Taylor and Francis website, where after some searching, I found a link for "Supplementary Data" which allowed me to download a zip file. The zip file was a PDF of all the tables they referenced in their paper.  Hooray! Now if only all other journals in the Early 2000s provided such info. Yes, I had to work at it, but it was rewarding as I found the pot o' gold. Should research be this heard? Not in today's world, but I grew up with manual typewriters and driving to libraries to see 1800s books as photocopies of them didn't exist, so a little extra digital digging is fine by me. I do miss the smell of old books...

If you ever wonder why people get so specific about the weight of Appalachiosaurus, I believe I have the answer. Under the description of the femur Carr et al. note, "Using the equation of Anderson et al. (1985) a mass of 623 kilograms (1,374 pounds) is estimated for RMM 6670." Happily, they provide the methodology used to derive the weight, something that surprisingly doesn't happen as much as you'd like to think it does. Future authors, take note!

The paper includes the 31 characters they used for their cladistic analysis. Fascinatingly to me, 27 of the 31 characters were cranial. I initially thought 30 of 31 were but then carefully read each character and discovered characters 25 and 31 both pertain to the femur, 27 is for the tibia, and 28 for the ischium. I suppose there is a reason they weren't grouped, maybe multiple authors, or characters were remembered at the end of the project?  31 characters, how far Carr has come since 2005 ;-).

James Lamb advised (pers. comm., 2024) he returned to the McWane Science Center in January of 2006 and "I quickly realized there was this one chunk of what had to be a major limb bone based on cross sectional size. The excavation of the specimen had been a complete mish-mash... I had all the pieces from both collections (Auburn and RMM) together, and in about a week fitted pieces together for what I took to be the humerus. I based that on the size, cross sectional shape, etc. It did have a long crest, only partially preserved that I took to be the deltopectoral crest. Neither end of the bone was completely preserved." This would be the first time a "long arm" had been mentioned as far as I can tell. There is one challenge with this narrative, Carr et al. in 2005 published an image of it already assembled, replete with both ends...

2008 The Mount is Completed

McWane mounts their long-armed Appalachiosaurus. Why were its arms considered long? I'd been trying to figure that out for literally hours, then had the idea to ask Triebold. He gave me the quotes above and sent me to James. My hypothesis at that stage of my research was because it was found to be a tyrannosauroid, and Dryptosaurus, has arms longer than tyrannosaurs, maybe someone assumed it'd have longer arms? James' interpretation of the bone being a humerus, plus it being a tyrannosauroid, matched my hypothesis. Hooray!

I found a sign in a photo of the exhibit stating the bone at the top left of the opening image is a humerus. We know it is a pubis, as was noted in the 2005 paper, and I suspect after communicating with James that it was he who made the sign, or advised on its creation.

The long-armed Appalachiosaurus received good press, always great for the museum. Triebold subsequently made a short-armed version for venues that had purchased the long-armed version and wanted a change, or wanted the long-arm for reasons known only to them.  He still offers a three-fingered, long armed version here.

2012 Jovanelly and Lane

Jovanelly and Lane published in a peer-reviewed, now-defunct, Open Access journal, Bentham OPEN a fun study on functional morphology, where they attempted to ascertain the speed and bite force of Appalachiosaurus. Their opening paragraph, however, threw me for a loop of confusion. I'll quote it in its entirety and highlight in bold the part that perplexed me:

Since its discovery in the 1980s, Appalachiosaurus has been the center of debate regarding arm length and the number of digits on the manus. Proponents on both sides offer compelling evidence in the form of cladistical analysis and a supposed fossilized humerus from the large theropod. The humerus, which has yet to be confirmed as belonging to Appalachiosaurus, suggests that this tyrannosaur retained primitive features well into the Cretaceous while others of the same period lost them. 

They provide no citations, no names, nothing about this "debate" at all. Nor do they figure the humerus. In all the literature I combed through in this project I found no reference to a humerus in any paper.

They go on to write (emphasis mine), "Those measurements included: right femur, metatarsal, tibia, astragalus, toe, humerus, jaw depth, skull length, snout width, jaw length, and tooth size (Tables 1 and 2)." Yet nowhere in their tables of measurements does a humerus appear. 

They also wrote, emphasis again mine, "They [Appalachiosaurus] likely took on smaller prey compared to T-rex [sic], which would have been able to take down larger prey such as sauropods alone. Furthermore, the serrations on the teeth would have aided in hunting as they would likely capture food that would later putrefy, making their bite a lethal dose of bacteria like the modern day Komodo dragon [14]." 

Of course I had to track that Komodo dragon reference down, which means I'm now on hour three poking around papers not involving sauropods... 

I did some digging, found the paper, read it (a neat paper about bite forces using the "dry skull" method), and found no reference to Komodo dragons in it at all. Nor bite force, and certainly no "lethal dose of bacteria." I then dug through the rest of their References section, thinking maybe they applied the wrong number in their in-line citation (it happens). Nope, there is no paper that references anything to do with Komodo dragons or bacterial warfare in their bibliography.

As for Tyrannosaurus rex eating a sauropod, I'm not sure where that came from, either. There is a putative Milk River sauropod caudal vertebra that David Evans and Michael Ryan mentioned in an 2018 SVP abstract that the paper is hopefully coming out in 2025 (y'all have a few days left for it to appear!). Their paper would be the first instance of a sauropod from western Canada. 

I'm wondering if they initially included a larger section about the "humerus" and the reviewers didn't think it was funny as they recognized it was a pubis (to reiterate, the pubis is a pubis) and rejected that part of the paper. The authors removed that section but perhaps left their controversial opening paragraph in to thumb their nose? Or forgot? Wild speculation for sure...

Let's all repeat, there is no humerus, the "humerus" is a pubis. However, keep reading as the humerus makes one more appearance, this time in a peer-reviewed academic paper no less!

Downstream Problems aka The Long Arm of the Mount

Having a tyrannosauroid specimen at a museum with long arms, with an actual pubis labeled as humerus, in the "old days" before the internet would cause local problems and erudite papers might feud a bit. With the rise of cell phones, social media, and the fact that most people seem to get their information from Wikipedia, the ability to spread misinformation is massive. Here is a post on Wikipedia which sat for some time (years?) before the disclaimer about it likely had two fingers and shorter arms, appeared.   

Here are images from February 2008 posted by the McWane Science Center of the mount being installed. The long arms are evident. Another shows the McWane Science Center paleontologist James Lamb "...works on a showcase that will display the differences in the T-Rex [sic] and Appalachiosaurus arms."

Looking into the 2012 paper, the primary author, Jovanelly, calls the Department of Physics, Astronomy, Geology department of Berry College home, where she appeared to be, in 2011, an Assistant Professor of Geology. Lesley Lane interned at the Tellus Science Museum, and she and another student, Tom Baldvins, presented “My What Long Arms You Have: An Investigation into the Unique Features of Appalachiosaurus, T-Rex’s [sic] Appalachian Cousin" at the museum in Cartersville, Georgia.

Interestingly, on or around Dec 21st, 2012, the Tellus Museum's skeletal mount received shorter arms. Was it coincidence that the undergraduate research about it having long arms that was held at the Tellus?

Yet, even in 2021, I found this quote, "The bones suggest that Appalachiosaurus did have longer and perhaps more useful arms than T-rex [sic], though the arms haven’t actually been found yet." from a newspaper article. The reasoning isn't cited by the author but I presume twas told this by someone. How much of the tyrannosauroid factor comes into play?

Dryptosaurus, the other reasonably well-known East Coast tyrannosauroid, has a reduced humerus, defined as the ratio of the humerus to the femur, coming in at .375. To put that into perspective, my eyeballing yielded humer:femur ratios as follows: Velociraptor .7, Allosaurus MOR 693 .6, the spinosaurid Suchomimus .52, Gorgosaurus .31, Albertosaurus .3, Tyrannosaurus .29, and Tarbosaurus having but a .27.

The new-to-this-sauropod-lad character, phalanx I-1:femur ratio of 0.2 for Dryptosaurus, means it has relatively large hands with a short arm. Compare that with a Tyrannosaurus .05 (presuming I did my math right :-)). I eyeballed "Big Al" at the MOR and came up with a .2, DINO 11541 (A. jimmadseni) I came up with a .14 or .11 (I was working off photos I had taken and not actual measurements, so please be kind :-)). Allosaurus is definitely handsy, so if I did my equations right, Dryptosaurus was a point guard!

Social Media and Peer Reviewed Journals

I never did receive links to his "post-2012 pubis/humerus critiques" but that is understandable, one look at the bone indicates it is not a pubis. And it was on Twitter, he doesn't know me, so why continue in such an exchange?

However, Schwimmer's comments about the journal have stuck with me, especially in the "pay to publish" world many journals have become. There clearly is a perception that "bigger" journals (defined by circulation? cost to publish?) are "more accurate" and that regional journals are less so. I've spent the last week finishing up an Ultrasaurus tabriensis manuscript that will be published in a regional journal, and funny enough, all papers on the Dry Mesa Dinosaur Quarry have been in regional journals. Does that make them "lesser"?

Peer review seems to have started in the 1960s, certainly none of the pre-1960s sauropod papers were peer reviewed! Yet Cope, Hatcher, Holland, Gilmore, Lull, Marsh, and Osborn and Mook are cited in every sauropod paper. Does that make their papers "lesser"? (OK, no comment on most of the Cope and Marsh works :-)) What about the papers I routinely cite from Lull 1919, Gilmore 1936, Hatcher 1901, some of the "classics!" The list could go on for many paragraphs. Are they "lesser" because they weren't peer reviewed?

What about websites like this mine, or SVPOW, or Equatorial Minnesota?  Paleofiles? Mickey Mortimer's site? There are easily a dozen blogs that are as good, or better in many cases, than a peer-reviewed journal article. Especially with the ability to upload as much as one wants, of whatever one wants, in whatever format one wants (website infrastructure permitting). 

One may argue, "how do I know that website will be running in a year? Five years?" and I ask the same of anything online. Any academic reading this paragraph will slowly purse their lips and slowly shake their head vertically when I ask this question, "Have you encountered a paper where the supplementary information was no longer available from a journal article?" Sometimes I can track down the physical copy via interlibrary loan, or someone has a PDF of it, but often I am unable to do so. When the author has passed away where does one gather the info? And that presumes a living author could find their original files from 2005, effectively nine-computer-hard-drives and three email addresses ago...

Here is where I am stuck. If a journal says "peer reviewed" then I have to take it at its word that it was peer-reviewed. Many reviewers opt to remain anonymous. Sometimes I wonder how a paper even made it into publication, the errors are so stunning (a sauropod paper from last year comes to mind...). Just because a paper was peer-reviewed doesn't make it good. 

2013 Loewen, Irmis, Sertich, Currie, and Sampson

HOW?

What isn't speculation is some of the best theropod lads around, in the Lythronax descriptive paper, actually scored an Appalachiosaurus humerus and ulna in 2013!  Mark Loewen, Randy Irmis, Jose Sertich, Phil Currie, and Scott Sampson are wonderful paleontologists, but how did this happen?

Here is the raw link. You'll need to click the "Supplementary Materials" link to download their matrix.

https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3819173/ 

I'll post their Humerus and first Ulna characters here:

Humerus
377. Humerus, form of shaft: (Carpenter et al., 2005; Brusatte et al., 2009:218)
378. Humerus, proximal head, morphology:(Sereno et al. 2009:73; Brusatte et al., 2010:240)
379. Deltopectoral crest form: (Modified from Li et al., 2009:140 and Brusatte el al., 2010:)
380. Humerus, rotation along shaft, orientation of long axis of proximal end relative to that of distal end: (Carr and Williamson 2010:236; Brusatte et al., 2010:242)
381. Humerus, additional muscle attachment tubera at the corner of the anterior and lateral surfaces distal to the deltopectoral crest:  (Brusatte et al., 2010:243)
382. Humerus, form of distal condyles: (Brusatte et al., 2010:245)
383. Humerus, length relative to the femur: (Carr and Williamson 2010: 235; Sereno et al. 2009: 72; Brusatte et al., 2010:239)

384: Ulna, shaft axis, form: (Li et al., 2009; Sereno et al. 2009:75; Brusatte et al., 2010:246)

362: Radius and ulna separation: (Li et al., 2009:214)

Now let's look how they were scored in their matrix:

What humerus?  What ulna?  A forelimb? These are intentionally coded characters. Did they lift them from a previous cladogram? I'm on hour five, digging through the weeds, more perplexed than ever, as I checked Brusatte et al. 2010 and Carr and Williamson 2010, plus older matrices by a number of theropod lads. None have scores for the forelimb material of Appalachiosaurus.

Does having 7 of 9 characters present in a taxon where those bones aren't present impact the final phylogeny?  Stay tuned!

Their results recover Appalachiosaurus outside of Tyrannosauridae, supporting Carr et al. 2005. It "is nested above Alectrosaurus and Xionguanlon [SIC], and a clade of the Asian non-tyrannosaurids Alioramus altai and Alioramus remotus, is the sister taxon to Tyrannosauridae. They close with, "Among all analyses, the reconstructions with the highest relative probability suggest that Appalachiosaurus had either an Asian or Appalachian ancestral range." 

Aside, in the paper they use the spelling Xionguanlon but the actual spelling is Xiongguanlong. A minor typo in a document this large but I thought I'd mention it.

None of the aforementioned taxa are scored for humerus or ulna characters, just '?'. They did all agree Appalachiosaurus is a tyrannosauroid, just outside the tyrannosaurs proper. After blurring my eyes on hour 7 of this sauropod-forsaken project, I see the overall trend is to concur the limbs reduce in size, with the forearms going before the hands. At this point I've come to the conclusion I need to run their matrix with all ???.  So I reach out to my good friend Gunnar Bivens.

Gunnar ran the numbers and... nothing. The tree's topology remained the same, albeit 3 steps shorter when correcting for the limbs. Correcting for the tooth and claw character errors changed nothing at all. So they gifted Appalachiosaurus a forearm, scored it, and nothing changed??? My guess is because none of its closest relatives in their matrix have forelimb material, either, so it was a 'no harm, no foul' scenario. The analysis that included false data returned the same tree, and that is a testament to the system, that the "true characters" swamped out the false characters. But the opposite could have easily occurred here, too, that the system ignored strong phylogenetic pulse because its closest relatives simply didn't have them preserved. It further adds to my distrust of these trees. Call me old as I was a grad student in the 1990s, before many of you were born, and had to do cladistics by hand... but something just feels... off. 

 Conclusions

It seems like every time I sit down to track down a simple answer on non- sauropods it turns into a deep dive that ends up uncovering all sorts of muck. Maybe non-sauropods are inherently dirty? :-)

Appalachiosaurus does not have long arms. The pubis is a pubis. Yes, the short arms are based on negative evidence, but it is strong negative evidence, considering its closest tyrannosauroid cousins all have reduced arm lengths. Yes, they won't be the ultra-derived tiny arms of T. rex or Tarbosaurus, but they aren't the long arms of dromaeosaurids, spinosaurids, or allosaurids, either.  

Coda 1 - Oh, AL...

Would someone describe the "Big AL" family? I can't find good measurements or descriptions anywhere. The pathologies were the stars then people moved on. It is frustrating. But then I reflect upon myself and the skeletons I haven't described yet. Please know I am working diligently on them when I'm not George R. R. Martin-ing on long write-ups like this ;-).

Coda 1.5 - Happens to all of us

Every paper has errors. In a recent Haplocanthosaurus paper I was a co-author on we reversed two digits within a specimen number in one of its nine appearances. We didn't catch it. We also provided a wrong measurement for a bone. There were five sauropod specialist authors on it, and we all missed that one to. Granted, there were dozens of measurements, but still... Ray Wilhite and I left our names off of our 2024 SVP poster!  Yep, that was mortifying. We were so focused on our work that we forgot to take credit for it! So mistakes happen, and I've made them myself. However, theropod experts letting a non-existent forelimb get scored, or scoring tyrannosauroids as edentulous? I don't want to judge so I'll say it just goes to show even the best have room for error, and improvement. 

Coda 2

Links to online content are wonderful... when they work. The Carr et al. 2005 paper provides hyperlinks to Tables in this statement:

Tables 14-16, on-line at http://www.vertpaleo.org/jvp/

Yet clicking on that link results in:

As this blog is on my family-friendly business account, the site automatically holds all comments. I'll strive to keep an eye on them so I can share them. I'd "turn people loose" but I've had some unpleasant experiences, thus the "restrictor plate."

BC 8/11/2024 started (7 hours), competed 12/4/2024 (5 hrs more)