Ontogeny and gender dimorphism in pterosaurs – SVP abstract 2016

Unfortunately,
and apparently, this is yet another study (Anderson and O’Keefe 2016) with a priori species assignations prior to a robust phylogenetic analysis and the creation of precise reconstructions. I hope I’m wrong, but no mention of phylogenetic analysis appears in the abstract. Nor do they mention creating reconstructions. Bennett (1993ab, 1995, 1996a, 2001ab, 2006, 2007) failed several times in similar fashion (with statistical analyses) to shed light on the twin issues of pterosaur ontogeny and dimorphism, coming to the wrong conclusions every time, based on results recovered by creating reconstructions and analyses. Further thoughts follow the abstract.

From the Anderson and O’Keefe abstract:
“The relationships of pterosaurs have been previously inferred from observed traits, depositional environments, and phylogenetic associations. A great deal of research has begun to analyze pterosaur ontogeny, mass estimates, wing dynamics, and sexual dimorphism in the last two decades. The latter has received the least attention because of the large data set required for statistical analyses. Analyzing pterosaurs using osteological measurements will reveal different aspects of size and shape variation in Pterosauria (in place of character states) and sexual dimorphism when present. Some of these variations, not easily recognized visually, will be observed using multivariate allometry methods including Principle Component Analysis (PCA) and bivariate regression analysis. Using PCA to variance analysis has better visualized ontogeny and sexual dimorphism among Pterodactylus antiquus, and Aurorazhdarcho micronyx. Each of the 24 (P. antiquus) and 15 (A. micronyx) specimens had 14 length measurements used to assess isometric and allometric growth. Results for P. antiquus analyses show modular isometric growth in the 4th metacarpal, phalanges I–II, and the femur. Bivariate plots of the ln-geometric mean vs ln-lengths correlate with the PCA showing graphically the relationship between P. antiquus and A. micronyx which are argued here to be sexually dimorphic and conspecific. Wing schematic reconstructions of all 39 specimens were done to calculate individual surface areas and scaled to show relative intraspecific wing shape and size. Finally, Pteranodon, previously identified having with sexually dimorphic groups, was compared with ln-4th metacarpal vs ln-femur data, bivariately, revealing similarities between the two groups (P. antiquus and A. micronyx = group 1; Pteranodon = group 2) in terms of a sexual dimorphic presence within the data sets.”

Figure 3. Click to enlarge. The Pterodactylus lineage and mislabeled specimens formerly attributed to this “wastebasket” genus

If these two workers actually had 24 P. antiquus specimens to work with,
then it was only because the labels told them so. Or they came across a cache on a slab of matrix I’m not aware of. Pterodactylus has been a wastebasket taxon for a long time (Fig.1) that, apparently the authors didn’t bother to segregate with analysis. Anderson and O’Keefe do not indicate they arrived at a large clade of P. antiquus specimens after phylogenetic analysis. Having done so, I can tell you that no other tested Pterodactylus is  identical to the holotype and no two adult pterosaurs I’ve tested are alike, even among Rhamphorhynchus, Germanodactylus and Pteranodon. The differences I’ve scored are individual to phylogenetic and they create cladograms that illuminate interrelationships, not sexual dimorphism or ontogeny. There are sequences of smaller species and larger ones. These can appear to be two genders, but that is a false result.

Embryo to juvenile pterosaurs
are isometrically miniaturized versions of their parents as the evidence shows time and again across the pterosaur clade. These facts have been known for over five years and it’s unfortunate that old traditions continue like this unfettered and untested under phylogenetic analysis… or so it seems… I could be wrong having not seen the presentation.

References
Anderson EC and O’Keefe FR 2016. Analyzing pterosaur ontogeny and sexual dimorphism with multivariate allometery. Abstracts from the 2016 meeting of the Society of Vertebrate Paleontology.
Bennett SC 1993a. The ontogeny of Pteranodon and other pterosaurs. Paleobiology 19, 92–106.
Bennett SC 1993b. Year classes of pterosaurs from the Solnhofen limestone of southern Germany. Journal of Vertebrate Paleontology. 13, 26A.
Bennett SC 1995. A statistical study of Rhamphorhynchus from the Solnhofen limestone of Germany: year classes of a single large species. Journal of Paleontology 69, 569–580.
Bennett SC 1996a. Year-classes of pterosaurs from the Solnhofen limestones of Germany: taxonomic and systematic implications. Journal of Vertebrate Paleontology 16:432–444.
Bennett SC 2001a, b. The osteology and functional morphology of the Late Cretaceous pterosaur Pteranodon. Part I. General description of osteology. Palaeontographica, Abteilung A, 260:1–112. Part II. Functional morphology. Palaeontographica, Abteilung A, 260:113–153.
Bennett SC 2006. Juvenile specimens of the pterosaur Germanodactylus cristatus, with a review of the genus. Journal of Vertebrate Paleontology 26:872–878.
Bennett SC 2007. A review of the pterosaur Ctenochasma: taxonomy and ontogeny. Neues Jahrbuch fur Geologie und Paläontologie, Abhandlungen 245:23–31.

Just for fun: the evolution of decorations and plumage in rock stars

Everything evolves.
Which begs, the question, how did a lineage of rock stars evolve from one another? I’ve chosen the “wild” branch of the rock-n-roll tree here (Fig. 1), a lineage told in pictures.

Did I miss anyone?

Figure 1. Evolution of rock stars beginning with Ike Turner (1951, Rocket 88) and Big Mama Thornton (1952, Hound Dog) and the rest you probably know. From sweet and clean cut rockstars evolved to become androgynous and highly decorated. Not shown: Liberace, who evolved plumage by convergence early on.

If anyone wants to parallel this lineage with the lineage of feathers in birds, crests in pterosaurs or Longisquama (Fig. 3), be my guest. So far, it seems, no reptile (including humans) have invented anything more exotic than plumage, with the possible exception of wings and flight (Fig. 2). When rockstars go “all out” they seem to move toward more exotic plumage. What goes around (in the Triassic), comes around (in the recent).

Figure 2. David Lee Roth of Van Halen demonstrating bipedal leaping and hind wing gliding, as in Longisquama (Fig. 3) and Sharovipteryx.

Have a rockin’ good day!

Figure 3. The lineage of pterosaurs recovered from the large reptile tree. Huehuecuetzpalli. Cosesaurus. Longisquama. MPUM 6009 with increasing plumage and ultimately, fore wings and hind wings.

Another Really Tiny Pterosaur: BMNH 42736

The smallest known pterosaur B St 1967 I 276 (No. 6 of Wellnhofer 1970 ) was discussed earlier. Today we get to meet maybe the second smallest pterosaur, Pterodactylus meyeri BMNH 42736 (Munster 1842, Fig. 1) is the same size as No. 6, but had several distinct traits (Fig. 2). I ran across the BMNH specimen in Unwin’s (2006) The Pterosaurs From Deep Time book on page 151. Dr. Unwin considered the specimen a “flapling” (= newly hatched pterosaur able to fly) with a wingspan of 17 cm, so that is the reconstructed scale (Fig. 3).

The Value of a Reconstruction
It’s a shame that modern workers don’t produce reconstructions of crushed pterosaurs anymore. There is so much to see (Figs. 2, 3), especially when one compares similar specimens. Many traits would go unnoticed if left crushed.

Figure 1. One of the world's smallest pterosaurs, traced from Unwin (2006, p. 151). The feet of the "flapling" were not articulated and a certain amount of guesswork was applied to the idenfication of the pedal elements and their reconstruction. Note how the left radius and ulna are parallel to and beneath the elongated right scapula. The right coracoid is largely beneath the right humerus. The left hand and proximal wing finger are beneath the right hand. Soft tissue stains are highlighted in orange. The wing had a narrow chord at the elbow. Colorizing the bones is a result of employing DGS, digital graphic segregation.

Phylogenetic Nesting
Here the “flapling” nested between No. 6 and No. 12, two other tiny ornithocephalians (and former Pterodactylus) outside of the Pterodactylus lineage, at the base of the Germanodactylus clade. Conveniently (for those looking for transitional taxa) No. 6 was smaller and No. 12 was larger than the BMNH “flapling.”

Distinct from No. 6, the “flapling” had a deeper skull, more and smaller dorsal vertebrae and ribs, a longer scapula (almost touched the pelvis), a deeper and more fully fused pelvis and a larger sternal complex than either of its sisters. Considering the reconstructed differences in quadrate elevation, jugal shape and pelvis dimensions (Fig. 2), you might think the “flapling” would have nested further apart from No. 6 and No. 12. These distinctions suggest that the “flapling” may have been at  the base of its own clade of currently unknown descendants.

Figure 2. The tiniest pterosaurs. On the left, Unwin's "flapling" Pterodactylus meyeri BMNH 42736. On the right, B St 1967 I 276, No. 6, the former sole owner of the title.

Juvie or Adult?
If the BMNH tiny pterosaur was indeed a juvenile of a larger more established taxon, which one did it match up to? And if so, why did it nest with other tiny pterosaurs? No. The BMNH specimen was an adult. The many autapomorphies (= differences) in the “flapling” also follow a larger trend seen in other tiny pterosaurs: morphological innovation.

Figure 3. Full scale image of ginkgo leaf and the two smallest pterosaurs to scale on a 72 dpi screen. Yes, these are tiny, but just look at the size of a hatchling on the far right, no bigger than a small fly.

Special Premaxillary Teeth
In the BMNH “flapling” we see more substantial anteriorly-directed medial teeth forming the tip of the premaxilla. Those two teeth evolve to become one in the rostral tip of Germanodactylus. That tooth is the only one retained in so-called “toothless” pterosaurs like Pteranodon and Nyctosaurus that have sharply tipped jaws.

Bigger Eggs?
A deeper pubis and pelvis in the BMNH specimen could have produced a larger egg. A stronger sternal complex and longer scapula could have made the “flapling” a more powerful flyer.

Soft Tissue Preservation
Despite a flipped mandible and poorly preserved feet, the “flapling” is otherwise well preserved and largely articulated. A soft tissue stain can be seen (overprinted in Fig. 1) that demonstrates a narrow chord at the elbow wing membrane construction.

As always, I encourage readers to see specimens, make observations and come to your own conclusions. Test. Test. And test again.

Evidence and support in the form of nexus, pdf and jpeg files will be sent to all who request additional data.

References
Meyer H von 1842. Notes on labyrinthodonts and fossil reptiles, including a description of Belodon plieningeri, new gen. and sp. Neues Jahrbuch fur Mineralogie, Geologie und Palaontologie 1842, pp. 301-304.
Unwin D M 2006. The Pterosaurs From Deep Time. 347 pp. New York, Pi Press.
Wellnhofer P 1970. Die Pterodactyloidea (Pterosauria) der Oberjura-Plattenkalke Süddeutschlands. Abhandlungen der Bayerischen Akademie der Wissenschaften, N.F., Munich 141: 1-133.

Ontogeny: Pteranodon vs. Diomedea

The Albatross (Diomedea exulans) and Pteranodon ingens (Fig. 1) were two large soaring reptiles, the former a bird, the latter a pterosaur. Insight into the ontogeny, maturity and lifespan of the extant Diomedea may shed some light on the extinct Pteranodon.

Figure 1. Left: Pteranodon. Right: Diomedea (albatross).

Widest Wingspan Among Birds
The albatross has a wingspan that reaches an extreme of 12 feet and averages 9 feet. The pterosaur Pteranodon had a similar wing plan with an enlarged wingspan up to 20 feet. Thus, and for little other reason, the albatross is the closest living analog to Pteranodon.

Albatross: Latest Maturation Among Birds
Most birds reach maturity in less than a year and many (crow, ostrich) do so in two years. By contrast, albatross males begin breeding at age 7, females at age 10. Some wait until 13. The life expectancy of an albatross is 30 years. According to Couzens (2008) it takes years for the albatross to become proficient at finding enough food for itself and more to take on the extra task of feeding a chick. The albatross also takes a long time to establish a pair bond.

Lizards vs Birds: Traditional Views
Most workers follow the paradigm that cold-blooded lizards mature more slowly than warm-blooded birds and mammals. Often, but not always, this is the case. However, more than mere physiology, size is typically an overriding factor. As everyone knows, mice mature faster than dogs, which mature faster than elephants and humans. The blue whale, which matures at 5 (females) or 8+ (males), does not follow this pattern. Among cold-blooded reptiles, iguanas are sexually mature at 50% of maximum size before the end of the second year (Kaplan 2007). Varanus hatchlings triple in size to sexual maturity and reach maximum size by the end of the first year (Pianka 1971). However some may continue growing thereafter, reaching up to 50% longer.

Pterodaustro Ontogeny
The only pterosaur for which a complete record of growth is known is the filter-feeder Pterodaustro. Chinsamy et al. (2008) reported: “…upon hatching, Pterodaustro juveniles grew rapidly for approximately 2 years until they reached approximately 53% of their mature body size, whereupon they attained sexual maturity. Thereafter, growth continued for at least another 3–4 years at comparatively slower rates until larger adult body sizes were attained.” So, this pterosaur’s growth rate, despite an apparent warm-blooded metabolism and active lifestyle, was not dissimilar to that of other lizards, reaching sexual maturity at 50% of the ultimate size. However, Pterodaustro took twice as long as Varanus, a cold-blooded lizard. Apparently, growth was not so rapid in pterosaurs — more along the lines of Iguana.

Pteranodon
If we add in the factor of increased size to what we know of Pterodaustro, we can imagine that Pteranodon might have had a maturation rate similar to that of the albatross (sexually mature at 7 to 10) along with a similar lifespan (30 years). However, if Pteranodon was more like Pterodaustro we get 2 years until half-grown, 5 to 6 years until fully grown.

Where Are the Juvenile Pterosaurs?
A long maturation brings up a problem. Where are the juveniles and immature forms (ages 0 to 6)? The Pterodaustro bone beds (nesting sites) provide the only evidence. There we find all sizes of Pterodaustro.

Ptweety the Only Juvenile Pteranodon
We know of only one juvenile Pteranodon. All others are adults that fit neatly into a phylogenetic framework of increasing size and crest size originating with a specimen of Germanodactylus (SMNK-PAL 6592) as an outgroup. This falsifies the current paradigm presented by Bennett (1991, 2001) and followed by others (Hone et al. 2011) of gender and maturation variation in most known specimens of Pteranodon. Here there is evidence of speciation leading to the largest crested forms (that in one clade only preceded a continuing clade of smaller crested, smaller forms.)

Bone Histology
The age of sexual maturity in Pteranodon has not yet been determined. Neither has the lifespan. Bone histology in Pteranodon has not provided the data needed due to crushing and resorption of the inner walls of the extremely thin long bones. At present we can only guess using extant analogs, like the albatross, and extinct analogs, like Pterodaustro.

Then There’s the Tiny Pterosaur Hypothesis
Tiny pterosaurs giving birth to fly-sized hatchlings were likely terrestrial until reaching adult-size due to desiccation problems, as discussed earlier. Larger pterosaur hatchlings, like Pteranodon (and all known, apparently flight ready pterosaur embryos), did not have a problem with desiccation — but they may have retained some sort of non-flying lifestyle living in environments not conducive to fossilization. This may explain the lack of immature pterosaurs in the fossil record (contra all traditional studies that considered tiny adults to be juveniles and embryos to be flight ready).

Not ready to jump on the flightless hatchling hypotheses quite yet, but it’s something to consider when faced with current and future evidence.

Just a Reminder
Maisano (2002) provides guidance on lizard ontogeny that can be applied to pterosaurs. That is: fusion can precede maturation and ultimate size or fusion may never take place in the oldest individuals, depending on their phylogeny. Recent work by Lü et al. (2012) show that the archosaur model continues to be wrongly applied to pterosaur studies.

As always, I encourage readers to see specimens, make observations and come to your own conclusions. Test. Test. And test again.

Evidence and support in the form of nexus, pdf and jpeg files will be sent to all who request additional data.

References
Chinsamy A, Codorniú L and Chiappe LM 2008. Developmental growth patterns of the filter-feeder pterosaur, Pterodaustro guinazui. Biology Letters, 4: 282-285.
Couzens D 2008. Extreme Birds: The world’s most extreme and bizarre birds. Firefly Books.
Hone DWE Naish D and Cuthill IC 2011. Does mutual sexual selection explain the evolution of head crests in pterosaurs and dinosaurs? Lethaia, DOI: 10.1111/j.1502-3931.2011.00300.x
Kaplan M 2007. Iguana Age and Expected Size. iguana/agesize online
Lü J, Unwin DM, Zhao B, Gao C and Shen C 2012. A new rhamphorhynchid (Pterosauria: Rhamphorhynchidae) from the Middle/Upper Jurassic of Qinglong, Hebei Province, China. Zootaxa 3158:1-19. online first page
Maisano JA 2002. Terminal fusions of skeletal elements as indicators of maturity in squamates. Journal of Vertebrae Paleontology 22: 268–275.
Pianka E 1971. Notes on the Biology of Varanus tristis. West. Aust, Natur, 11(8):80-183.