The larger specimen of Sinopterus atavismus enters the LPT basal to dsungaripterids

Many pterosaur fossils attributed to Sinopterus
have been described. They vary greatly in size and shape.

Presently four Sinopterus specimens have been added
to the large pterosaur tree (LPT, 253 taxa). They are all sister taxa, but as in Archaeopteryx, no two are alike, one is basal to the others, which are, in turn, basal to large clades within the Tapejaridae.

  1. Sinopterus dongi (the holotype) nests basal to the Tupuxuara clade.
  2. Sinopterus liui nests in the Tupuxuara clade.
  3. Sinopterus jii (aka Huaxiapterus jii) nests basal to the Tapejara clade.
  4. Sinopterus atavisms (Figs. 1-4; Zhang et al. 2019; IVPP V 23388) nests basal to the Dsungaripterus (Fig. 4) clade, outside the Tapejaridae.
Figure 1. Sinopterus atavismus in situ.

Figure 1. Sinopterus atavismus in situ. IVPP V 23388

From the Zhang et al. 2019 abstract:
“Here, we report on a new juvenile specimen of Sinopterus atavismus from the Jiufotang Formation of western Liaoning, China, and revise the diagnosis of this species.”

Zhang et al. note that several elements are unfused including a humeral epiphysis. Several pits and grooves in the distal ends of the long bones are also pitted and grooved. Normally these would be good indicators in archosaurs and mammals, but pterosaurs are lepidosaurs and lepidosaurs follow distinctly different ‘rules’ for growth (Maisano 2002). As an example, some pterosaur embryos have fused elements. Some giant pterosaurs have unfused elements. Here the new specimen (IVPP 23388) is considered an ontogenetic adult as its size is similar to other phylogenetic relatives.

“Sinopterus atavismus does not present a square-like crest. Moreover the feature that groove in the ventral part of the second or third phalanx of manual digit IV is not diagnostic of the species.”

Zhang et al. are comparing the new larger IVPP specimen to the smaller, previously described (Lü et al. 2016) XHPM 1009 specimen (then named Huaxiapterus atavismus), which they considered conspecific. The XHPM specimen has wing phalanx grooves while the IVPP specimen does not. The shapes of the skulls do not match (Fig. 3) and we know that pterosaurs grew isometrically. Thus these two specimens are not conspecific.

“In the new material, the skull preserves a pointed process in the middle part of the dorsal marginof the premaxillary crest, which is different from other Chinese tapejarids. Considering the new specimen is known from a large skeleton that differed from the holotype, this difference may be related to ontogeny, as the premaxillary crest of the holotype is short and does not extend as long as that of the new specimen.”

These two specimens are not conspecific, so ontogenetic comparisons should not be made.

Figure 2. Sinopterus atavismus reconstruction.

Figure 2. Sinopterus atavismus reconstruction.

From the Zhang et al. 2019 discussion:
“Except for D 2525 which represents an adult individual of Sinopterus (Lü et al. 2006b), all Chinese tapejarid pterosaurs known so far were immature individuals at the time of death. The new specimen (IVPP 23388) shares some features with the holotype of Sinopterus atavismus. The wingspan of the new material is about twice as long as that of the holotype of S. atavismus.”

As mentioned above, the IVPP V 23388 specimen is here considered an adult with unfused bone elements. It needs both a new generic and specific name. The XHPM 1009 specimen (Fig. 3) requires further study.

Figure 3. Sinopterus atavismus size comparison

Figure 3. Sinopterus atavismus size and shape comparison.

The present confusion about the ontogenetic status of pterosaurs 
could have been largely resolved with the publication of “The first juvenile Rhamphorhynchus recovered by phylogenetic analysis” and other papers suppressed by pterosaur referees. Sorry, readers, we’ll have to forge ahead with the venues we have.

Figure 3. Sinopterus atavismus skull restored (gray areas).

Figure 4. Sinopterus atavismus skull restored (gray areas).

Figure 4. Sinopterus atavisms compared to Dsungaripterus to scale.

Figure 5. Sinopterus atavisms compared to Dsungaripterus to scale.

Sinopterus atavismus (Zhang et al. 2019; Early Cretaceous; IVPP V 23388) was originally considered a juvenile member of the Tapejaridae, but here nests as a small adult basal to Dsungaripteridae. The antorbital fenestra is not taller than the orbit. The carpals are not fused. No notarium is present. The antebrachium is robust. The distant pedal phalanges are longer than the proximal pedal phalanges. An internal egg appears to be present (but half-final-size adults were sexually mature according to Chinsamy et al. 2008,)

Sinopterus dongi IVPP V13363 (Wang and Zhou 2003) wingspan 1.2 m, 17 cm skull length, was linked to Tapejara upon its discovery, but is closer to Tupuxuara.

Sinopterus? liui (Meng 2015; IVPP 14188) is represented by a virtually complete and articulated specimen attributed to Sinopterus, but nests here at the base of Tupuxuara longicristatus.

Sinopterus jii (originally Huaxiapterus jii, Lü and Yuan 2005; GMN-03-11-001; Early Cretaceous) is basal to the Tapejara in the LPT, distinct from the other sinopterids basal to Tupuxuara.

Figure 5. Click to enlarge. The Tapejaridae arise from dsungaripterids and germanodactylids.

Figure 5. Click to enlarge. The Tapejaridae arise from dsungaripterids and germanodactylids.

The present LPT hypothesis of interrelationships
appears to be a novel due to taxon inclusion, reconstruction and phylogenetic analysis. If not novel, please let me know so I can promote the prior citation.

Traditional phylogenies falsely link azhdarchids with tapejarids
in an invalid clade ‘Azhdarchoidea‘. The LPT has never supported this clade (also see Peters 2007), which is based on one character: an antorbital fenestra taller than the orbit (that a few sinopterids lack). Pterosaur workers have been “Pulling a Larry Martin” by counting on this one character and by excluding pertinent taxa that would have shown them this is a convergent trait ever since the first cladograms appeared in Kellner 2003 and Unwin 2003.

Figure 1. Gene studies link swifts to hummingbirds. Trait studies link swifts to owlets. Trait studies link hummingbirds to stilts.

Figure x. Gene studies link swifts to hummingbirds. Trait studies link swifts to owlets. Trait studies link hummingbirds to stilts.

Unrelated update:
The stilt, Himantopus (Fig. x) has moved one node over and now nests closer to the hummingbird, Archilochus. Both arise from the Eocene bird, Eocypselus, which also gives rise to the hovering seagull, Chroicocephalus. The long, mud probing beak of the stilt was adapted to probing flowers in the hummingbird. All these taxa nested close together in the LRT earlier.


References
Chinsamy A, Codorniú L and Chiappe LM 2008. Developmental growth patterns of the filter-feeder pterosaur, Pterodaustro guinazui. Biology Letters, 4: 282-285.
Kellner AWA 2003. 
Pterosaur phylogeny and comments on the evolutionary history of the group. Geological Society Special Publications 217: 105-137.
Lü J and Yuan C 2005. 
New tapejarid pterosaur from Western Liaoning, China. Acta Geologica Sinica. 79 (4): 453–458.
Maisano JA 2002. The potential utility of postnatal skeletal developmental patterns in squamate phylogenetics. Journal of Vertebrate Paleontology 22:82A.
Maisano JA 2002.
Terminal fusions of skeletal elements as indicators of maturity in squamates. Journal of Vertebrae Paleontology 22: 268–275.
Peters D 2007. The origin and radiation of the Pterosauria. In D. Hone ed. Flugsaurier. The Wellnhofer pterosaur meeting, 2007, Munich, Germany. p. 27.
Unwin DM 2003. On the phylogeny and evolutionary history of pterosaurs. Pp. 139-190. in Buffetaut, E. & Mazin, J.-M., (eds.) (2003). Evolution and Palaeobiology of Pterosaurs. Geological Society of London, Special Publications 217, London, 1-347.
Wang X and Zhou Z 2003. A new pterosaur (Pterodactyloidea, Tapejaridae) from the Early Cretaceous Jiufotang Formation of western Liaoning, China and its implications for biostratigraphy. Chinese Science Bulletin 48:16-23.
Zhang X, Jiang S, Cheng X and Wang X 2019. New material of Sinopterus (Pterosauria, Tapejaridae) from the Early Cretaceous Jehol Biota of China. Anais da Academia Brasileira de Ciencias 91(2):e20180756. DOI 10.1590/0001-3765201920180756.

wiki/Sinopterus

Growth pattern of a new large Romualdo pterosaur

Bantim et al. 2020 document
a new “pteranodontoid pterosaur with anhanguerid affinities (MPSC R 1935) from the Romualdo Formation (Lower Cretaceous, Aptian-Albian), is described here and provides one of the few cases where the ontogenetic stage is established by comparison of skeletal fusion and detailed osteohistological analyses.”

Figure 1. Excellent wing finger carpophalangeal joint from the Bantim et al. 2020 paper. Note the unfused sesamoid (extensor tendon process), a phylogenetic trait of lepidosaurs, not an ontogenetic trait of archosaurs, as phylogenetic analysis documents.

Figure 1. Excellent wing finger carpophalangeal joint from the Bantim et al. 2020 paper. Note the unfused sesamoid (extensor tendon process), a phylogenetic trait of lepidosaurs, not an ontogenetic trait of archosaurs, as phylogenetic analysis documents.

Continuing from the abstract
“The specimen … consists of a left forelimb, comprising an incomplete humerus, metacarpal IV, pteroid and digits I, II, III, IV, including unguals. This specimen has an estimated maximized wingspan of 7.6 meters, and despite its large dimensions, is considered as an ontogenetically immature individual. Where observable, all bone elements are unfused, such as the extensor tendon process of the first phalanx and the carpal series. The absence of some microstructures such as bone resorption cavities, endosteal lamellae, an external fundamental system (EFS), and growth marks support this interpretation. Potentially, this individual could have reached a gigantic wingspan, contributing to the hypothesis that such large flying reptiles might have been abundant during Aptian-Albian of what is now the northeastern portion of Brazil.”

Anhanguera

Figure 2. Anhanguera.

By comparison,
coeval Anhanguera has a 4.6m (15 ft) wingspan. The largest complete ornithocheirid, SMNK PAL 1136 has a 6.6m wingspan.

Bone elements fuse and lack fusion
in phylogenetic patterns (rather than ontogenetic patterns) in the clade Pterosauria, as documented earlier here in 2012. That is why you can’t keep pretending pterosaurs are archosaurs and not expect problems like this to accumulate. Your professors are taking your time and money and giving you invalidated information.

Figure 5. Largest Pteranodon to scale with largest ornithocheirid, SMNS PAL 1136.

Figure 5. Largest Pteranodon to scale with largest ornithocheirid, SMNS PAL 1136.

It is a continuing black mark on the paleo community
that pterosaurs continue to be considered archosaurs by paid professionals when phylogenetic analysis (and Peters 2007 and the LRT) nests pterosaurs with lepidosaurs. That is why pterosaurs have lepidosaur phylogenetic fusion patterns (Maison 2002, 2002) distinct from archosaur ontogenetic fusion patterns. Just add taxa colleagues. The pterosaur puzzle piece does not fit into the archosaur slot… everyone admits that. The pterosaur puzzle piece continues to fit perfectly and wonderfully in the fenestrasaur tritosaur lepidosaur slot.


References
Bantim RAM et al. (5 co-authors) 2020. Osteohistology and growth pattern of a large pterosaur from the lower Cretaceous Romualdo formation of the Araripe basin, northeastern Brazil. Science Direct https://doi.org/10.1016/j.cretres.2020.104667
Maisano JA 2002. The potential utility of postnatal skeletal developmental patterns in squamate phylogenetics. Journal of Vertebrate Paleontology 22:82A.
Maisano JA 2002.
Terminal fusions of skeletal elements as indicators of maturity in squamates. Journal of Vertebrae Paleontology 22: 268–275.
Peters D 2007. 
The origin and radiation of the Pterosauria. In D. Hone ed. Flugsaurier. The Wellnhofer pterosaur meeting, 2007, Munich, Germany. p. 27.

https://pterosaurheresies.wordpress.com/2013/05/14/phylogenetic-fusion-patterns-in-pterosaurs/

Hone et al. 2020 vs. Rhamphorhynchus

Long one today.
Summary, for those in a hurry:
Hone et al. 2020 bring us their views
on Rhamphorhynchus ontogeny (= growth from hatchling to adult). Unfortunately, this study is based on several invalid assumptions. Lacking a phylogenetic context, Hone et al. made the mistake of comparing small adults to large adults. No juveniles were tested. Subsequent ontogeny comparisons to birds and bats were thus rendered moot.

Figure 2. Rhamphorhynchus specimens to scale. The Lauer Collection specimen would precede the Limhoff specimen on the second row.

Figure 2. Rhamphorhynchus specimens to scale based on results from the LPT. No two are alike — except the juvenile Vienna specimen and the adult n81.

Before we get started, you might remember:

  1. A competing paper has been online for 2 years: ‘First Rhamphorhynchus juvenile recovered by phylogenetic analysis’ in which only one juvenile/adult pairing was found among all 31 specimens shown in figure 1. Among the rest, no two are alike. The small ones in the top row are not juveniles, but phylogenetically miniaturized adult basal Rhamphorhynchus species. (Perhaps someday, someone will re-name them all appropriately.)
  2. All pterosaurs (so far tested) develop isometrically (with the exception of tapejarid crests) because that’s what lepidosaurs do.
  3. Hatchling pterosaurs are typically 1/8 as tall as adults.
  4. Only hatchlings of a certain minimum size can fly. Hatchlings below this hypothetical size risk desiccation due to a high surface-to-volume ratio. That’s when quadrupedal locomotion enters pterosaur clades. Extradermal soft tissue that limits desiccation first appears on tiny, flapping pre-pterosaurs like Cosesaurus.
  5. New pterosaur clades often begin with a series of phylogenetically miniaturized transitional taxa (as in Fig. 1). This only appears in phylogenetic analyses when small and large taxa are analyzed together. That has not happened yet in published analyses because other workers make the same mistake as they consider small adult taxa to be mismatched juveniles (thereby destroying the Hone et al. isometry hypothesis).

The Hone et al. 2020 paper was announced today on
Dr. Hone’s email list. After a short comparison to Pteranodon, Hone continues:
“However, if we turn to Rhamphorhynchus we have only a fraction of the number of specimens but pretty much all the other issues are absent. They also cover a near order of magnitude in size with everything for animals of c 30 cm wingspan up to nearly 2 metres and include everything from putative hatchling-sized animals to a couple of genuine outliers that are much bigger than other known individuals.”

A good sample of Rhamphorhynchus taxa are shown above (Fig. 1) in phylogenetic order. Note this genus has its genesis as a phylogenetically miniaturized series following Campylognatoides in the large pterosaur tree (LPT, 250 taxa). The sole juvenile shown above is the Vienna specimen, nesting with one of the ‘genuine outliers that are much bigger.’ This adult and juvenile pairing nest together with virtually identical scores, despite the great difference in size.The LPT was able to lump and split all tested Rhamphorhynchus taxa. So it can be done. Hone et al. omitted this all important step and ruined their paper.

Hone continues:
“The numbers of course are not tiny, well over 100 good specimens, and that alone would make them an exceptional sample of most terrestrial Mesozoic archosaurs.”

Our first red flag! Hone et al. do not realize that when taxa are added, pterosaurs move over to lepidosaurs. On another note: relative to ‘100 good specimens’, 31 are shown above (Fig. 1).

Hone explains
that Wellnhofer (1975) featured 108 specimens. Hone’s group looked at 129, but, as Hone confesses, “The ‘real’ total is actually a little lower.” Oddly, in the text of the paper, Hone et al. report testing 135 specimens of R. muensteri.

Hone continues:
“This post inevitably marks the publication of an analysis of growth in Rhamphorhyunchus. In a lot of ways, this mirrors Chris Bennett’s fantastic 1995 paper on this genus where he convincingly demonstrated that all specimens belonged to a single species and not multiple ones as previously thought, and part of his arguments for doing this looked at the relationships between various elements based on Wellhofer’s dataset.”

Our second red flag! Bennett’s 1995 paper likewise did not include a phylogenetic analysis. When several specimens of Pteranodon were added to the LPT, no two nested together as conspecific taxa (Fig. 2). Small specimens were closer to the genesis of Pteranodon following Germanodactylus. Large specimens split into several clades.

Figure 2. The Tanking-Davis specimen compared to other forms. Specimen w and specimen z appear to be the closest to the Tanking-David specimen. Specimen 'w' = Pteranodon sternbergi? USNM 12167 (undescribed). Specimen 'z' = Pteranodon longiceps? Dawndraco? UALVP 24238. Click to enlarge.

Figure 2. The Tanking-Davis specimen compared to other forms. Specimen w and specimen z appear to be the closest to the Tanking-David specimen. Specimen ‘w’ = Pteranodon sternbergi? USNM 12167 (undescribed). Specimen ‘z’ = Pteranodon longiceps? Dawndraco? UALVP 24238. Click to enlarge.

Hone is delighted to announce
“Chris’ point [in Bennett 1993, 1995] was that while there were some discreet clusters of specimens (which he attributed to year classes) most of the alleged differences between the putative species vanished when you put them on a graph and the rest were classic ontogenetic traits like the fusion of the pelvis in large individuals of big eyes in small ones. So while he didn’t really deal with growth as such, he was already showing similar patterns to what I and my coauthors confirm now – Rhamphorhynchus was weirdly isometric in growth.”

Our third red flag! Dr. Hone does not appear to realize that ALL pterosaurs  develop isometrically during ontogeny. They do this because pterosaurs are lepidosaurs. By contrast, archosaurs develop allometrically. I’m also going to throw in the objection that a graph or two (as in Bennett 1993, 1995, Hone et al. 2020] is no substitute for a thorough phylogenetic analysis.

Hone continues:
“In other words, in the case of the vast majority of their anatomy, young animals are basically just scaled down adults.”

This is an odd statement to make considering the fact that Hone et al. are looking at phylogenetically miniaturized adults (Fig. 1) and regarding them as juveniles. That Hone considers the little specimens, “basically carbon copies of the adults” makes one question the precision of their observations. They did cherry-pick two similar taxa (Figs. 3, 4), avoiding the wider variation of other specimens. A competing online analysis (subset Fig. 5) was able to split and lump all Rhamphorhynchus specimens.

For comparison, Hone et al. also looked at ontogeny in bats,
noting hand/wing development accelerated close to sexual maturity (= shortly after weaning). He notes, “This is the pattern we would expect.”

Our fourth red flag! Since Hone et al. have blinded themselves to the possibility that pterosaurs are lepidosaurs (Peters 2007) they don’t look at lepidosaurs for comparison. Here’s why they should: pterosaurs hatch with adult proportions from leathery eggs held within the mother’s body longer than in any archosaur.

Hone continues:
“Birds are functionally poor analogues of pterosaurs but are much closer phylogenetically and are the only other powered flying tetrapod so we also looked at some existing datasets for them too.”

More traditional myth perpetuating here. I find this all so disheartening. Colleagues, just add taxa. If I can do it as an outsider, you can do it as a PhD. Do not be afraid to do the work of constructing a cladogram.

Hone continues:
“If you grow isometrically you wings will get longer and wider but your weight will increase much faster since you as a whole will get longer and wider and deeper. Birds increase penumaticity as they grow and there’s evidence this is the case in other pneumatic clades too and if so for pterosaurs, then the mass increase in adults would also be offset somewhat by a proportionally lower mass in adults for a given volume than juveniles.”

Very good point. But I’m ot sure of any pneumaticity studies comparing hatchling and adult pterosaurs.

Hone continues:
“Precociousness has been suggested in pterosaurs before based on the evidence for them flying while young, but it has also been challenged. It suggested that to be flying at that size would require a huge amount of effort and this would leave little energy for growth.”

Wait a minute! Didn’t he just say the weight would increase by the cube in adults? That means juveniles were that much lighter.

Hone continues:
“That’s largely true, but overlooks that there could be post hatching parental parental care. That is normal for archosaurs (including dinosaurs) and we would expect it for pterosaurs.”

If only pterosaurs were archosaurs, but at this point they still nest with lepidosaurs. Most lepidosaurs fend for themselves after hatching, and if pterosaur hatchlings could fly, then they would be able to fly off on their own shortly after hatching. Best not to ‘expect’ anything without a valid phylogenetic context, evidently lacking in Hone et al. 2020.

Hone continues:
“So in short, Rhamphorhynchus is perhaps the best pterosaur for large studies about populations and growth and this genius at least grew isometrically, and this may or may not be the same for other pterosaurs.”

But for the present, every pterosaur known from embryo, juvenile and adult shows strict isometric growth (except for tapejarid crests).

“But it does imply that young pterosaur could fly, and fly well.”

Sadly, Hone et al. seem to be looking at small adults (Fig. 1) and calling them ‘young’. Of course these adult pterosaurs can fly well!

Apparently Hone et al. are comparing linear measurements and graphing them. That method produced false positives for Bennett 1995. There is no substitute for phylogenetic analysis.

In this topsy-turvy world of pterosaurs,
myths are popularized by PhDs while comprehensive phylogenetic analyses compiled by amateurs are ignored and suppressed. Not sure why this problem is not more widely recognized. For other missteps made by Dr. Hone with regard to pterosaurs, click here or use the keyword ‘Hone’ for that long list.

Moments ago the paper itself arrived.
In my morning email was a message from Dr. Hone: “Attached” along with a PDF of their Rhamphorhynchus paper. Two sets of graphs are present, but only a single figure combining bat allometry and Rhamphorhynchus isometry (isolated in Fig. 3).

Figure 3. Image from the only non-graph figure in Hone et al. 2020. Identification and permission note from that caption. Compare these taxa to those in figures 1 and 4.

Figure 3. Image from the only non-graph figure in Hone et al. 2020. Identification and permission note from that caption. Compare these taxa to those in figures 1 and 4.

Figure 4. Lateral view of Hone et al. 2020 Rhamphorhynchus taxa taken from ReptileEvolution.com (Fig. 1).

Figure 4. Lateral view of Hone et al. 2020 Rhamphorhynchus taxa taken from ReptileEvolution.com (Fig. 1). Hone et al. cherry-picked these two somewhat similar by convergence taxa assuming the smaller one was a juvenile of the other other. Phylogenetic analysis separates these two (see Fig. 1). Note the differences in pedal element proportions.

From the paper:
“We test whether pterosaurs show a similar pattern of rapid forelimb growth during post‐hatching/ontogeny to that of bats and birds, and thus infer when in ontogeny R. muensteri would have become volant.”

Sounds laudable. Let’s see how they do it.

From the paper:
“All Rhamphorhynchus specimens from Bavaria are now considered a single species (Bennett 1995).”

No. That’s why figure 1 was created and a phylogenetic analysis of pterosaurs was run (subset Fig. 5), to see how specimens could be lumped and separated. Like Hone et al., Bennett likewise eschewed the use of phylogenetic analysis. Sadly, Hone et al. adopted without further consideration Bennett’s invalid assumption, rather than testing Rhamphorhynchus with a phylogenetic analysis.

Figure 4. Subset of the LRT focusing on Rhamphorhynchus.

Figure 5. Subset of the LRT focusing on Rhamphorhynchus.

From the paper:
“Four lines of evidence suggest that the smallest R. muensteri specimens were very young animals and potentially hatchlings.

  1. Histology reveals incomplete ossification of long bones in the smallest specimens tested (Prondvai et al. 2012),
  2. A disproportionate number of known specimens are small, consistent with high juvenile mortality (Bennett 1995; Hone & Henderson 2014)
  3. Late‐stage embryos of pterosaurs had well‐developed, ossified wings (Wang & Zhou 2004; Codorniú et al. 2018)
  4. and finally while few fossilized pterosaur embryos are known, the ratio by which adults are larger than embryos (Lü et al. 2011; Wang et al. 2017) is similar to the size ratio between the largest R. muensteri specimens and the smallest.”

Incomplete ossification: the smallest specimen studied by Prondvai et al. (2012) was BSPG 1960 I 470a = n9 (Figs. 1, 5) is also the second most primitive tested specimen (next to n28) in a phylogenetic miniaturization series that began with Campylognathoides. Among the neotonous / juvenile traits retained was incomplete ossification of the long bones. Lacking a phylogenetic context, neither Prondvai et al. nor Hone et al. were aware of the miniaturized adult status of n9.

Figure 5. the B St 1960 I 4709A specimen of Rhamphorhynchus is the first and one of the smallest phylogenetically miniaturized specimens.

Figure 5. the B St 1960 I 470a specimen of Rhamphorhynchus (at right)  is the second most primitive and one of the smallest phylogenetically miniaturized specimens attributed to Rhamphorhynchus. One of the neotonous traits was incomplete ossification. Hatchlings were 1/8x the size of adults, similar to house flies in size.

Disproportionate number of specimens are small: lacking a phylogenetic context, Hone et al. were not aware of the phylogenetic miniaturization that preceded the evolution of larger Rhamphorhynchus specimens. In the LPT only one Rhamphorhynchus specimen is a valid juvenile nesting with larger adults.

Late‐stage embryos of other pterosaurs had well‐developed, ossified wings: So did miniaturized adults.

Size ratio of largest R. muensteri specimens to smallest similar to embryo vs adult sizes in other pterosaurs: lacking a phylogenetic context, Hone et al. were not aware of the phylogenetic miniaturization that preceded the evolution of larger Rhamphorhynchus specimens. Hone et al. made the mistake of labeling small adults as juveniles. Notably, Hone et al. did not try to match their purported juveniles with adults phylogenetically. Other tiny Rhamphorhynchus specimens have juvenilized proportions (smaller rostrum, larger orbit), but these were ignored by Hone et al., who cherry-picked two comparative taxa out of 135.

From the paper:
“We tested for isometric versus allometric growth across 135 specimens of R. muensteri using bone length and composite measures (e.g. total wing length and total leg length) relative to: (1) total body length, from rostrum tip to the end of the tail; (2) skull length; and (3) humerus length.”

Lacking a phylogenetic context (available online for several years), Hone et al. made the mistake of comparing adults to adults. No juveniles were tested. Subsequent comparisons to birds and bats were thus rendered moot.

From the paper:
“Our results suggest that even the smallest Rhamphorhynchus had adult skeletal proportions and thus wings sufficient for flight.” This confirms the conclusions of Peters (2018) using a phylogenetically validated juvenile Rhamphorhynchus, rather than a dataset full of large and small adults.

From the paper:
“Wang et al. (2017) noted that in embryos of the pterodactyloid Hamipterus, although there was greater ossification of the limbs and vertebrae than the head, including of the shafts of longbones, there was limited ossification of some other parts of the skeleton that may have related to flight. They hypothesize in this case that hatchlings may have been able to walk before they could fly, though still imply relatively early flight for these animals.”

These were not hatchlings, but embryos still developing within the egg, within the mother in the tradition of lepidosaurs.  Eruptive gases killed flocks en masse. Details here.

From the paper:
“Pterosaurs, like almost all other archosaurs, probably provided parental care (Witton 2013), and precocial flight need not preclude this possibility.” 

This is myth. We’ve known since Peters 2007 that adding taxa moves pterosaurs to nest within Lepidosauria.

From the paper:
“Thus, while Rhamphorhynchus apparently flew at a young age, such volant offspring may have plausibly received parental care, including provisioned food, as they became independent foragers.” 

There is no evidence for this bit of speculation. But it cannot be ruled out. According to Gans 1996, “Many aspects of reptilian reproductive patterns prove to be vagile among the vertebrates. Reversals complicate, and may even invalidate, the characterization of broad trends. Furthermore, the 7000 species of reptiles show dozens of modes that seem to enhance the fitness of their offspring, thereby providing a vast opportunity of testing the reality of these adaptations.”  (‘vagile’ = able or tending to move from place to place or disperse)

In summary:
Hone et al. assumed that phylogenetically miniaturized adults at the genesis of Rhamphorhynchus were juveniles. While testing small adults against large adults (Figs. 1–5) the authors determined that Rhamphorhynchus ontogeny proceeded isometrically.

Ironically this confirms earlier findings by Peters (2018 and elsewhere in this blog) using the only known phylogenetically validated juvenile and a matching adult Rhamphorhynchus. As longtime readers know, all other pterosaurs develop isometrically because they are lepidosaurs arising from taxa close to late-surviving Huehuecuetzpalli, known from matching juvenile and adult specimens.

Dr. Hone needs to show more leadership. He needs to create reconstructions of the specimens under study so visual comparisons can be made by his team and readers. Roadkill specimens are too difficult to compare otherwise. He also needs to run a phylogenetic analysis to determine interrelationships between pterosaur taxa and within all amniotes to see where pterosaurs nest. At present he’s perpetuating old myths and traditions that were invalidated twenty years. He’s that far behind the times.

I’ll never forget the day several decades ago
when Dr. Kevin Padian and Dr. Chris Bennett told me, “nothing can be known about a taxon until it is put into a phylogenetic context.” I took that advice to heart. That is why the LRT and LPT now include more than 2000 taxa.


References
Bennett SC 1993. The ontogeny of Pteranodon and other pterosaurs. Paleobiolgy 19(1):92-106.
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.
Gans C 1996. An overview of parental care among the Reptilia. Advances in the Study of Behaviour 25:145–157.
Hone DWE, Ratcliffe JM, Riskin DK, Hemanson JW and Reisz RR 2020. Unique near isometric ontogeny in the pterosaur Rhamphorhynchus suggests hatchlings could fly. Lethaia. Paywall access here.
Hone 2020. Email post. How to grow your dragon – pterosaur onotgeny [sp]
Peters D 2007. The origin and radiation of the Pterosauria. In D. Hone ed. Flugsaurier. The Wellnhofer pterosaur meeting, 2007, Munich, Germany. p. 27.
Peters D 2018. First juvenile Rhamphorhynchus recovered by phylogenetic analysis. PDF here.
Prondvai E, Stein K, Ösi A, Sander MP 2012. Life History of Rhamphorhynchus Inferred from Bone Histology and the Diversity of Pterosaurian Growth Strategies. PlosOne. online pdf
Wellnhofer P 1975a-c. Teil I. Die Rhamphorhynchoidea (Pterosauria) der Oberjura-Plattenkalke Süddeutschlands. Allgemeine Skelettmorphologie. Paleontographica A 148: 1-33. Teil II. Systematische Beschreibung. Paleontographica A 148: 132-186. Teil III. Paläokolgie und Stammesgeschichte. Palaeontographica 149:1-30.

wiki/Rhamphorhynchus

https://pterosaurheresies.wordpress.com/2012/03/23/not-another-rhamphorhynchus-growth-series-without-a-phylogenetic-analysis/

 

Flugsaurier 2018: ‘Young istiodactylid’ nests with tall pterodactylids in the LPT

Flugsaurier 2018 opens today, August 10,
and the abstract booklet is out. So it’s time to take a look at some of the news coming out of that Los Angeles pterosaur symposium. Since the purpose of the symposium is increase understanding of pterosaurs, I hope this small contribution helps.

Figure 1. The Erlianhaote specimen attributed by Hone and Xu 2018 to istiodactylidae nests in the LPT with the large derived pterodactylids.

Figure 1. The Erlianhaote specimen attributed by Hone and Xu 2018 to the clade Istiodactylidae (within Ornithocheiridae) nests in the LPT with the large derived pterodactylids. Note the un-warped deltopectoral crest and lack of a deep cristospine, along with the long legs and short wings.

Hone and Xu at Flugsaurier 2018
describe, “An unusual and nearly complete young istiodactylid from the Yixian Formation, China (Fig. 1). The specimen shows the characteristic istiodactylid cranial features of tooth shape and enlarged nasoantorbital fenestra. However, it has proportionally large hindlimbs and wing proportions that are similar to those of azhdarchids. This has led to suggestion that the specimen may be a composite and that only the cranial material is istiodactylid. Preparation work around some key parts revealed no inconsistencies in the matrix or evidence of glue. The specimen is held in the Erlianhaote Dinosaur Museum, Erlianhote, China.”

Figure 2. The Erlianhaote specimen nests with these pterodactylids in the LPT, not with Istiodactylus (Fig. 3). Compare to valid istiodactylids in figures 4–6/

Figure 2. The Erlianhaote specimen nests with these pterodactylids in the LPT, not with Istiodactylus (Fig. 3). Compare to valid istiodactylids in figures 4–6/

Reconstructed as is
(Fig. 2) and added to the large pterosaur tree (LPT, 233 taxa, not yet updated due to no museum number nor genus name) the young ‘istiodactylid’ nests as a large derived pterodactylid. 13 steps separate this taxon from the Istiodactylus clade.

Ornithocheirids,
like Istiodactylus (Figs. 3, 4) and the SMNL PAL 1136 specimen (Fig. 5), share a very large wing finger, a short metacarpus, a warped deltopectoral crest, small free fingers and deeply keeled sternal complex not found in the Erlianhote specimen.

Figure 3. Istiodactylus has a shorter neck, longer wing finger and deep cristospine, among other traits not found in the new Erlianhaote specimen.

Figure 3. Istiodactylus has a shorter neck, longer wing finger and deep cristospine, among other traits not found in the new Erlianhaote specimen.

Figure 4. Istiodactylus sinensis is an istiodactylid from China sharing few traits with the new Erlianhaote specimen. Note the warped deltopectoral crest not warped in the new specimen.

Figure 4. Istiodactylus sinensis is an istiodactylid from China sharing few traits with the new Erlianhaote specimen. Note the warped deltopectoral crest not warped in the new specimen. Manual 4.1 is shorter than in other well-known istiodactylids.

The largest ornithocheirid

Figure 5. The unnamed largest ornithocheirid, SMNK PAL 1136, nests with Istiodactylus.

Figure 6. The Erlianhaote pterodactylid reconstructed in several views.

Figure 6. The Erlianhaote pterodactylid reconstructed in several views. The imagined (gray) areas of the skull here were imagined as an istiodactylid, but the better restoration is shown in figure 2.

It’s better not to eyeball certain specimens.
Sometimes you have to run them through a phylogenetic analysis to find out what they are. That’s what the LPT is for. It minimizes taxon exclusion and handles convergence.

Pterosaurs are still lepidosaurs.
So they follow lepidosaur fusion patterns, which follow phylogeny. Hone and Xu made the mistake of imagining pterosaurs might have archosaur fusion patterns that follow ontogeny.

Why am I not at Flugsaurier 2018?
In addition to about a dozen reasons that I can list later, or your can guess now, I can be more helpful and timely here.

References
Andres B and Ji Q 2006. A new species of Istiodactylus (Pterosauria, Pterodactyloidea) from the Lower Cretaceous of Liaoning, China. Journal of Vertebrate Paleontology, 26: 70-78.
Bowerbank JS 1846. On a new species of pterodactyl found in the Upper Chalk of Kent P. giganteus). Quarterly Journal of the Geological Society 2: 7–9.
Bowerbank JS 1851. On the pterodactyles of the Chalk Formation. Proceedings of the Zoological Society, London, pp. 14–20 and Annals of the Magazine of Natural History (2) 10: 372–378.
Bowerbank JS 1852. On the pterodactyles of the Chalk Formation. Reports from the British Association for the Advancement of Science (1851): 55.
Hone DWE and Xu 2018. An unusual and nearly complete young istiodactylid from the Yixian Formation, China. Flugsaurier 2018: the 6th International Symposium on Pterosaurs. Los Angeles, USA. Abstracts: 53–56.
Hooley RW 1913. On the skeleton of Ornithodesmus latidens. An ornithosaur from the Wealden shales of Atherfield (Isle of Wight)”, Quarterly Journal of the Geological Society, 69: 372-421
Howse SCB, Milner AR and Martill DM 2001. Pterosaurs. Pp. 324-335 in: Martill, D. M. and Naish, D., eds. Dinosaurs of the Isle of Wight, The Palaeontological Association
Wang X, Rodrigues T, Jiang S, Cheng X and Kellner AWA 2014. An Early Cretaceous pterosaur with an unusual mandibular crest from China and a potential novel feeding strategy. Scientific Reports 4 : 6329, pp. 1-9. | DOI: 10.1038/srep06329
Witton MP 2012. New Insights into the Skull of Istiodactylus latidens (Ornithocheiroidea, Pterodactyloidea). PLoS ONE 7(3): e33170. doi:10.1371/journal.pone.0033170

wiki/Istiodactylus

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.”

The Pterodactylus lineage and mislabeled specimens formerly attributed to this "wastebasket" genus

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 RhamphorhynchusGermanodactylus 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.

One more look at Rhamphorhynchus growth

Usually I avoid histological (bone microstructure) studies.
But here’s one that merits one more extended report based on its many incorrect assumptions and overlooked comparisons.

Summary of key facts in this long blog:

  1. both phylogenetically miniaturized adult pteros and mammals had juvenile-like “woven” bone texture
  2. Pterosaur embryos develop in utero and had adult proportions, so they could fly upon hatching
  3. Pterosaurs develop isometrically, thus immature pteros can only be identified in phylogenetic analysis (= when larger identical adults are known).

Prondvai et al. 2012 tested growth strategies in Rhamphorhynchus. As noted earlier, Prondvai et al. confused small adults with juveniles and hatchlings, not following the clear data that pterosaurs grow isometrically, not allometrically. Thus the morphological difference shown here (Fig. 1) are phylogenetic, not ontogenetic. Phylogenetic analysis supports this hypothesis.

Figure 1. Bennett 1975 determined that all these Rhamphorhynchus specimens were conspecific and that all differences could be attributed to ontogeny, That is clearly false as shown here and by phylogenetic analysis. Only the juvenile between the two largest specimens is a non-adult. Click to enlarge.

Figure 1. Bennett 1975 determined that all these Rhamphorhynchus specimens were conspecific and that all differences could be attributed to ontogeny, That is clearly false as shown here and by phylogenetic analysis. Only the juvenile between the two largest specimens is a non-adult. Click to enlarge.

 

Age of first flight
Prondvai et al. 2012 report,
The initial rapid growth phase early in Rhamphorhynchus ontogeny supports the non-volant nature of its hatchlings, and refutes the widely accepted ‘superprecocial hatchling’ hypothesis. We suggest the onset of powered flight, and not of reproduction as the cause of the transition from the fast growth phase to a prolonged slower growth phase. Rapidly growing early juveniles may have been attended by their parents, or could have been independent precocial, but non-volant arboreal creatures until attaining a certain somatic maturity to get airborne.” Prondvai et all did not realize they were examining small adult pterosaur specimens, not juveniles. So rapid growth was part of their growth strategy. More refutations relevant to the above statements follow.

Powered flight is one of the most energy-consuming locomotion types in tetrapods, therefore high growth rates and a superprecocial onset of the flying lifestyle in a highly developed hatchling are mutually exclusive developmental parameters. The validity of this simple trade-off model is supported by the fact that the only extant superprecocial fliers, the megapod birds have very low if not the lowest growth rates among extant birds.” Prondvai et al. ignore the fact that megapodes have their rapid growth phase inside the egg shell. Hatchling megapodes are relatively “very large with a wingspan up to half that of the adult).”  By contrast, pterosaurs hatch at 1/8 the height of the adult and 1/8 the wingspan.

In support of supreprecocial flight…
pterosaur hatchlings had adult proportions. Tiny adults, the size of sparrows and hummingbirds, had larger pterosaur proportions. The smallest pterosaur that Prondvai et al. tested had wing tips that extended way over their heads when folded and quadrupedal (Figs. 1, 2). We’ve seen the short wings of flightless pterosaurs. Hatchlings of volant taxa don’t have short wings. Tiny adult pterosaurs may have ‘rapidly growing” bone microstructure because they matured quickly, reproduced as often as possible then died early, like tiny mammals do. More on this below:

Sexual maturity vs. size:
Prondvai et al. report, “According to the hypothesis presented here, the onset of powered-flight in Pterodaustro occurred after attaining 53% of adult size. Here we prefer the hypothesis that bone growth is slowed down by the initiation of a new, and much more energy consuming locomotory activity, namely powered flight.” Not by coincidence, this is the size that Chinsamy et al. (2008) determined that sexual maturity was attained. After observing the morphology of the embryo Pterodaustro, which matches the morphology of the adult, there is no supporting evidence for the Prondvai et al. hypothesis.

Archosaur vs. lepidosaur
Prondvai et al. do not consider the growth strategies and histology of lepidosaurs, only archosaurs. So they are making comparisons to the wrong clade. Pterosaurs nest within the Lepidosauria. Growth patterns in lepidosaurs are distinct and do not follow archosaur growth patterns (Masisano 2002). But this may not be the key factor in observed differences.

Chinsamy and Hurum 2006
looked at the basal lepidosaur, Gephyrosaurus. “The [bone] compacta consists of essentially parallel−fibred bone tissue interrupted by several lines of arrested growth (LAGs). The first LAG visible from the medullary cavity appears to be a hatchling line with its more haphazardly oriented, globular-shaped, osteocyte lacunae.”  This was not a phylogenetically miniaturized taxon even though it was a basal lepidosaur.

More to the point
Chinsamy and Hurum 2006 also looked at the basal and phylogenetically miniaturized mammal, Morganucodon. They report on, “distinct woven bone tissue with large, irregularly oriented osteocyte lacunae and several primary osteons. No secondary osteons were visible, though several enlarged erosion cavities are evident in the compacta. In the same section, it appears that substantial endosteal resorption had occurred, and parallel−fibred bone tissue is evident only in a localized area peripherally. This area includes several rest lines, which indicate pauses in the rate of bone formation, and hence, pauses in growth.” Perhaps these pauses indicate a lifespan of “several” years. Note the “woven bone” texture description.

Figure 1. Several tiny Rhmphorhynchus adults, among them is the n7 specimen tested by Prondvai et al. and considered a juvenile by them.

Figure 2. Several tiny Rhmphorhynchus adult sister taxa, among them is the n7 specimen tested by Prondvai et al. and considered a juvenile by them shown here about 7/10 of in vivo size. As you can see, these pterosaurs do not appear to have any impediments to flapping and flying. However their tiny hatchlings would probably not have flown based on their high surface/volume ratio. The adults had juvenile traits due to phylogenetic miniaturization.

The smallest sampled Rhamph bone microstructure
Prondvai et al. report about the tiny Rhamph, BSPG 1960 I 470a, (n7 in the Wellnhofer 1970 catalog, Fig. 2): “A thin layer of lamellar bone of endosteal origin rims the medullar [central] cavity. There seem to be only a few longitudinally oriented vascular canals, but these have rather large diameter in relation to the overall thickness of the cortex. The bone matrix is typically woven with some poorly defined, immature primary osteons, hence the majority of the cortex does not show the mature fibrolamellar pattern yet. The osteocyte lacunae are large and plump throughout the cortex, and possess an extremely well-developed system of dense, radially oriented canaliculi implying extensive communication and nutrient-exchange between the osteocytes. No LAGs or any other growth marks can be observed.”  Maybe LAGs were never present in this taxon if it lived for just a short time. Remember, we’re talking about phylogenetic miniaturization here.  If the small precocial Rhamphorhynchus specimens were maturing quickly and laying eggs early, they likely followed the life patterns of other tiny tetrapods, like Morganucodon (above) and died early, perhaps living only one or two years, not five or more as in mid-sized pterosaurs.

Note: Like Morganucodon (above) the phylogenetically miniaturized mammal, 
the bone structure in the smallest tested Rhamphorhynchus is described as “woven”.

Age vs size:
Prondvai et al. report, “The ontogenetic validity of the smallest size category of Bennett is clearly supported by the overall microstructure found in the bones of the three small specimens.” Unfortunately, without a phylogenetic analysis, Prondvai et al. did not realize that the smallest specimens were small due to phylogenetic miniaturization. Their ancestors were larger. Thus small Rhamphs retained juvenile and embryonic traits into adulthood, including the typical short rostrum and smaller wings. These traits also included juvenile “woven” bone tissue. Essentially these tiny pterosaurs were precocious sexually active adults in the former juvenile phase of development.

Precocial hatchling?
Prondvai et al. report, “Superprecocial embryos require substantial amount of nutrients stored in their eggs to reach an advanced level of somatic maturity state by the time the embryo hatches. If the egg volume of Darwinopterus was relatively as low as that of squamates, then how could it have contained so much yolk as to cover the energy requirements of an extremely well-developed, volant hatchling?” Prondvai are assuming that pterosaur eggs developed outside the uterus. As lepidosaurs, pterosaur embryos developed inside the uterus and the super thin eggshell was deposited last. Thus they could “cover the energy requirements.”

Apparently Prondvai et al. are not looking
at verified pterosaur hatchlings (in eggs), which are identical in morphology to adults. In some cases large embryos can be larger than small adult sister taxa! The Prondvai team know that the tiny Rhamps don’t have the same morphology as the medium or big rhamphs. Unfortunately, and this is a continuing problem… they don’t realize those changes are phylogenetic, not ontogneric.

With similar proportions of bone and muscle,
but at 1/8 as tall and therefore (8 cube rooted or) 1/512 as massive, juvenile pterosaur bone tissue would have been strong enough for sustained flight in such lightweight specimens. But that overlooks reality, where the specimens Prondvai are looking at are in fact tiny adults with juvenile bone structure, as in Morganucodon. We don’t know where small, medium and large Rhamphorhynchus laid its eggs, which were likely ready to hatch shortly after deposition. We don’t have any hatchling Rhamphorhynchus fossils. Hatchlings of the small and tiny adults would have been in danger of desiccation (high surface area/volume ratio), so we can presume they grew up in moist environs. Unfortunately Prondvai et al. did not test the one verified juvenile among in the Rhamphorhynchus clade, NHMW 1998z0077/0001 (Fig. 3), the Vienna specimen. No one thinks this juvenile could not fly based on its age/relative size.

Figure 1. Two specimens attributed to Rhamphorhynchus longiceps along with a third specimen that nested with the larger of the two with identical scores, thus identifying it as a juvenile R. longiceps.

Figure 3. Two specimens attributed to Rhamphorhynchus longiceps along with a third specimen, NHMW 1998z0077/0001, that nested with the larger of the two with identical scores, thus identifying it as a juvenile R. longiceps. No one thinks this Rhamph could not fly, despite its young age.

To their credit, Pronvai et al. suggest (following a hypothesis first presented here): “Alternatively, Rhamphorhynchus hatchlings could have been precocial to the effect that they could have left their nests immediately after hatching, but they must have been exclusively terrestrial or rather arboreal. They could have clambered around quadrupedally on the branches of trees feeding themselves with smaller invertebrates or vertebrates without any parental contribution.”

No universal growth strategy in pterosaurs
Prondvai et al. report, “In the light of the histological results it becomes evident that there is no universal pattern in the growth strategy of pterosaurs.” I am concerned that this conclusion was made without the the benefit of a phylogenetic analysis and without knowledge of phylogenetic miniaturization in the clade.

To their credit
Prondvai et al. report, “In contrast to Bennett’s  suggestion, the second size category of Rhamphorhynchus does not only include subadult but also adult specimens, hence it cannot be used as an indicator of real ontogenetic stage.”

References
Chinsamy A, Codorniu ́ L, Chiappe L 2008. Developmental growth patterns of the filter-feeder pterosaur, Pterodaustro guinazui. Biol Lett 4: 282–285.
Chinsamy A and Hurum JH 2006. Bone microstructure and growth patterns of early mammals. Acta Palaeontologica Polonica 51 (2): 325–338.
Maisano JA 2002. Terminal fusions of skeletal elements as indicators of maturity in squamates. Journal of Vertebrae Paleontology 22: 268–275.
Prondvai E, Stein K, O0 si A, Sander MP 2012. Life History of Rhamphorhynchus Inferred from Bone Histology and the Diversity of Pterosaurian Growth Strategies. PLoS ONE 7(2): e31392. doi:10.1371/journal.pone.0031392
Sekercioglu C 1999. Megapodes: A fascinating incubation strategy. Online article. 

Allometry during ontogeny in the basal tritosaur, Huehuecuetzpalli

Huehuecuetzpalli (Reynoso 1998) is a basal tritosaur according to the large reptile tree, a lepidosaur nesting outside of the Squamata, and ancestral to Tanystropheus, Macrocnemus, drepanosaurids, fenestrasaurs and ultimately, pterosaurs. The lineage of pterosaurs is shown here.

Huehuecuetzpalli specimens are only known from the Early Cretaceous, with ghost lineage origins going back to the Late Permian. Long species survival is not uncommon among lepidosaurs, as in the extant Sphenodon with relatives in the Triassic.

Figure 1. Two specimens of Huehuecuetzpalli were found, one adult and one juvenile. Here they are shown together to scale along with manus and pes comparisons scale to a common length for metacarpal 4 and metatarsal 4.

Figure 1. Two specimens of Huehuecuetzpalli were found, one adult and one juvenile. Here they are shown together to scale along with manus and pes comparisons scale to a common length for metacarpal 4 and metatarsal 4.

Reynoso 1998 reported,
“The relative length of the snout, and the proportions of the skull and limbs relative to the presacral vertebral column, do not show signifcant differences between the juvenile and adult specimens, although these features usually change in ontogeny. This suggests that adult proportions were already acquired at the ontogenetic stage of the younger specimen in spite of its relatively smaller size.”

I have been repeating this observation
with regard to pterosaurs, which likewise do not show any significant differences (apart from the enlargement of any skull crests) in their morphological proportions. For examples click here, here and here and other references therein.

But is it true for Huehuecuetzpalli?
That’s why side-by-side comparisons are so useful. Sadly, I have not done so until just yesterday (Figs. 1, 2).

Figure 2. Huehuecuetzpalli, adult and juvenile skulls to scale. note the relatively shorter rostrum in the juvenile, which also had smaller teeth and a shorter set of parietals (with a smaller braincase and smaller jaw adductor chamber). In the juvenile the ascending process of the premaxilla was more robust.

Figure 2. Huehuecuetzpalli, adult and juvenile skulls to scale. note the relatively shorter rostrum in the juvenile, which also had smaller teeth and a shorter set of parietals (with a smaller braincase and smaller jaw adductor chamber). In the juvenile the ascending process of the premaxilla was more robust and the tooth-bearing portion was shorter with fewer teeth.

Reynoso 1998 reported,
“The complete fusion of the cranial elements suggests that the larger specimen is of post-juvenile age, and probably an adult condition was already acquired. The olecranon process of the ulna, however, is not completely ossified and attached to the ulna, and only a ball of hard tissue (calcified cartilage or bone) is preserved. It was impossible to find information in the literature about the time when the precursor of the olecranon process become fused to the ulna.

“The age of the smaller specimen is more difficult to establish. The complete ossification of the fourth distal tarsal and the still separated astragalus and calcaneum undoubtedly suggest a post-hatchling stage. The complete fusion of the frontal, however, shows that it is older than Rieppel’s specimen number 18 and the hatchling of Cyrtodactylus pubisulcus (Gekkonidae) illustrated by Rieppel (1992a: ¢g. 1). The high degree of ossification indicates that it is close to the latest stages of development preceding complete ossification. Juvenile skull characters are the presence of a broader parietal table with short lateral processes. Compared with the adult skull, the juvenile parietal table is more than 15% broader on the narrower section excluding the ventrolateral flanges for the dorsal attachment of the jaw adductor musculature.”

We looked at olecranon ossification in tritosaurs earlier here.

As a rule, lepidosaurs don’t change much during ontogeny
as we’ve seen earlier here with Shinisaurus. But they do change… a little.

References
Reynoso V-H 1998. Huehuecuetzpalli mixtecus gen. et sp. nov: a basal squamate (Reptilia) from the Early Cretaceous of Tepexi de Rodríguez, Central México. Philosophical Transactions of the Royal Society, London B 353:477-500.

 

Allometry and Isometry in Shinisaurus Ontogeny

There are those who insist that pterosaur juveniles and hatchlings had a short rostrum and large orbit (Bennett 1995, 1996), citing similar allometric changes during ontogeny in mammals and archosaurs. The fact that pterosaurs are not mammals or archosaurs does not appear to matter. The large reptile tree nests pterosaurs firmly within the Fenestrasauria, within the Tritosauria, within the Lepidosauria (outside the Squamates) and within the Lepidosauriformes.

Earlier we looked at isometry (relative lack of change) during ontogeny (maturation) in several pterosaurs for which we have juveniles associated with adults. These observations don’t seem to matter much to pterosaur experts who want to believe that hatchling pterosaurs had cute features. Isometry during ontogeny is generally found trait among lepidosaurs and especially so among tritosaur lepidosaurs, as evidenced by Reynoso (1989) who noticed little to no change between a juvenile and an adult Huehuecuetzpalli.

Today we’ll take a look at allometry AND isometry during ontogeny in a rare living lizard (Squamata, Autarchoglossa, Anguimorpha), Shinisaurus crocodilurus (Figs. 1, 2, the Chinese crocodile lizard).

Figure 1. Lateral views of Shinisaurus adult and juvenile, to scale and to the same skull length. While the skull proportions are roughly the same (isometry) changes that can be noted are noted (allometry).

Figure 1. Lateral views of Shinisaurus adult and juvenile, to scale and to the same skull length. While the skull proportions are roughly the same (isometry) changes that can be noted are noted (allometry). Note there is no rostral elongation during maturation in this taxon. Images from Digimorph.org

The skulls of the juvenile and adult
show very little rostral elongation during maturity. The orbit is only slightly reduced in the adult. Larger changes are noted on the figures. Surprisingly, the teeth are relatively smaller in the adult. The expanded braincase in the juvenile is reduced in the adult, but an expanded (inflated) occiput is retained and further expanded in several burrowing lizards, retained in a process called neotony.

Figure 2. Dorsal views of Shinisaurus juvenile and adult with notes on isometric and allometric changes.

Figure 2. Dorsal views of Shinisaurus juvenile and adult with notes on isometric and allometric changes. Left image from Digimorph.org.

Wikipedia reports: Shinisaurus, the Chinese crocodile lizard, was once also regarded as a member of Xenosauridae, but most recent studies of the evolutionary relationships of anguimorphs consider Shinisaurus to be more closely related to monitor lizardsand helodermatids than to Xenosaurus. It is now placed in its own family Shinisauridae. The large reptile tree agrees with this nesting.

References
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 1996. Year-classes of pterosaurs from the Solnhofen limestones of Germany: taxonomic and systematic implications. Journal of Vertebrate Paleontology 16:432–444.

http://digimorph.org/specimens/Shinisaurus_crocodilurus/juvenile/
wiki/Chinese_crocodile_lizard

NOT a new Zhenyuanopterus: XHPM1088

Very, very close, but no cigar.

And not a juvenile either.
A new paper by Teng et al. (2014) reports on a small partial Zhenyuanopterus (XHPM1088, Fig. 1) that does quite fit the morphology of the holotype. No worries. They said it was a juvenile with some odd sorts of allometry going on.

I hate to say it, but we can blame Chris Bennett for this bit of wishful thinking as his 1995 and 1996 papers on Solnhofen pterosaurs opened the doors to letting almost any small specimen become the juvenile of any somewhat similar, but much larger specimen based on the false notion of allometry during ontogeny. Several specimens falsify that little fantasy, including all the embryos now known.

Phylogenetic analysis would have put a stop to such nonsense, but no analysis was undertaken, either in 1995, 1996 or 2014.

Figure 1. XHPM1088 in situ. Only the posterior half is preserved here.

Figure 1. XHPM1088 (mistakenly referred to Zhenyuanopterus) in situ. Only the posterior half is preserved here.

Here’s the problem
The new specimen has a relatively long and robust tail (15 caudals) and a more robust forelimb than hindlimb, plus a Yixian Formation (Early Cretaceous) locality. These facts identified this pterosaur as Zhenyuanopterus to its authors. With identical length ratios between the humerus and femur, Teng et al. thought growth was isometric in these bones, but not others. The scapula has an odd sort of shape otherwise found only in Zhenyuanopterus. However the coracoid was not the same shape or size ratio (Fig. 1). They thought the length of the coracoid would slow dramatically during growth compared to other bones, not realizing that taxa just outside of Zhenyuanopterus (i.e. Boreopoterus, Arthurdactylus, Fig. 2) had a similar long, straight coracoid. They also blamed the coracoid length problem on the holotype of Zhenyuanopterus, saying it was not well-preserved and giving it a longer redicted length based on XHPM1008. That’s not good Science, especially when the coracoids are well preserved and articulated in the holotype.

Unfortunately
Teng et al. thought one of the unique characters of Zhenyuanopterus was its small feet, but the reality is ALL ornithocheirids (more derived than the JZMP embryo) had tiny feet.

Figure 2. The partial pterosaur XHPM1088 to scale with Boreopterus and Zhenyuanopterus and also scaled up to a similar humerus length with Zhenyuanopterus.  Note the coracoids don't match. This is one of the few pterosaurs in which the tibia is shorter than the femur. Boreopterus is similar in this regard.

Figure 2. Click to enlarge. The partial pterosaur XHPM1088 to scale with Boreopterus and Zhenyuanopterus and also scaled up to a similar humerus length with Zhenyuanopterus. Note the coracoids don’t match. This is one of the few pterosaurs in which the tibia is shorter than the femur. Boreopterus is similar in this regard.

A beautiful illustration of Zhenyuanopterus is included in the paper (Fig. 3) sadly flawed by bat-like, deep chord wing membranes and an odd sort of hanging posture for a pterosaur, especially one with such small feet. Some traditions are very hard to kill.

Zhenyuanopterus-illustration

Figure 3. Zhenyuanopterus illustration by Zhao Chuang, a very talented artist. Sadly the wing membranes are wrong and the hanging posture is unlikely based on the tiny feet.

I encourage pterosaur workers
to start putting bones together in reconstructions, then adding new taxa to good phylogenetic analyses before assigning a juvenile status to a small pterosaur that doesn’t match a large one. Here’s a new genus that Teng et al. could have named, but didn’t.

Reference
Teng F-F, Lü J-C, Wei X-F, Hsiao Y-F and Pittman, M 2014. New Material of Zhenyuanopterus (Pterosauria) from the Early Cretaceous Yixian Formation of Western Liaoning. Acta Geologica Sinica (English) 88(1):1-5.