Two erectoid hominids from Sarstedt prompted a detailed examination of the course of the impressions of the Arteria meningea media and their characteristics within the line of hominidae

Alfred Czarnetzki

Dirk von Rautenfeld

Madjid Samii

English

Nature Precedings

 1 
 
 
 
 
 
 
 
The phylogenetic principles of mma; 
A hypothetical biostratigrafic model 
 
 
Czarnetzki, A.1, Berens von Rautenfeld, D.2, Samii, M. 3 
1) formerly: Dep. Paleanthropology and Osteology, Institute of Anthropology and Human Genetics,  
 University of Tuebingen, Wilhelmstr. 27, D-72074 Tuebingen 
2) Medizinische Hochschule Hannover, Inst. f. Funktionelle und  
 Angewandte Anatomie, Carl-Neuberg-Str.1, 30625 Hannover 
3) International Neuroscience Institute (INI), Rudolf-Pichlmayr-Straße 4, D-30625 Hannover 
4) 
 
When filling in our phylogenetic pedigree, no skeletal element can be omitted from 
consideration. The discovery of three relatively small fragments of the human skull[1], i.e. the 
skull fragments of the Neandertal from Neu-Warendorf [2] and the two erectoid hominids from 
Sarstedt [3] , impelled us to initiate a detailed examination of the course of the impressions of 
the Arteria meningea media (median meningeal artery = mma) and their characteristics within 
the hominidae. To this end, we examined 31 human cadavers by trepanning the braincase and 
studying the differences of the indentations between arteries and veins. Unfortunately the 
mma pattern, as found in the last full anatomical description of Dmanisi[4] or in Homo 
ergaster (i.e. KNM-ER3733) from eastern Africa (where it existed at 1.78 million years[5,6]), 
is rarely depicted in the literature. To be sure, different patterns in this artery (mma) have been 
described [7] and a summary[8 – 14]  given of the papers published thus far. Brandt[7] could 
demonstrate that many of the latter [d.h. these papers] described only diffuse types of 
indentation patterns or else differences occurring in these patterns between fossil specimens. 
The literature mentions varying degrees of complexity in the impression patterns left by 
vessels, but omits to say what criteria were used when differentiating between the impressions 
of veins and arteries at the different phylogenetic developmental stages (or, for that matter, in 
their phylogenetic biostratigraphy). The differentiation between arteries and veins was only 
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first insisted [15] on in 2003. Brunner mentioned that a “large amount of self-training and 
experience” is indispensable for differentiating as exactly as possible between veins and  
arteries. An overview was tabled on the extent of variation in the arterial pattern of specimens 
deriving from Eurasia and the northern part of Africa[11, 15].  
 
We are now in a position to differentiate, for the first time, between the main branch of the 
mma that is impressed just after passing the foramen spinosum (including its size) and side 
branches. Based on our data, we could observe throughout the phylogenetic process not only 
the existence of different patterns [9,16,17,]  but also a generally continuous change in the arterial 
pattern found in specimens from Eurasia. Precisely this arterial system and its main branches 
are very conservative on both species and subspecies levels, as is well known and well 
attested. In particular, we could observe changes in (i) the course, (ii) the size and (iii) the 
branching of the meningeal artery – unlike the internal carotid artery, it solely supplies the 
surface of the cerebrum (especially the parietal lobe) with arterial blood.  
Thus, in the earliest phylogenetic representatives of the species Pithecanthropus erectus, we 
find that (i) the rear branch is the one that has the greatest size and features an individually 
variable number of smaller branches. This main branch, (ii) other than in modern humans, 
generally assumes a posterior course soon after passing the Foramen spinosum, as is depicted 
in Fig. 1, Stage 1. This pattern has been described[15] as an “Asian” one. 
These changes can be useful for reconstructing the developmental stages of the parietal lobe. 
Our phylogenetic model is based not only on our own observations, including inter al. on 
fossil specimens like Pithecanthropus I (= Trinil 2) and Pithecanthropus II (= Sangiran 2), 
Smb4, OH9, Zhoukoudien 10, Steinheim and Reilingen, Salé and Arago, Vindija, Mladeč, 
Weimar-Ehringsdorf, 31anatomically modern humans (H.s.s.) and published data [8,9, 10, 11, 13, 
15] . But it is based on a consideration of other schemes [10, 18] as well. A final point: the 
following description of the main branch and its tributaries is based (see fig. on the 
assumption of an anatomically modern brain. 
 
Stage 1  
represents the earliest phylogenetic pattern. Its main criteria are a) anterior convexity in the 
trajectory of the artery after passing the Foramen spinosum and b) branching off within the 
temporal lobe into a main posterior branch and a tiny anterior one. These posterior branches 
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supply the main surface of the parietal lobe, while the anterior sidelines supply only a very 
small part and differ in having a significantly smaller size. (Holotype: Trinil 2). In 
 
Stage 2  
the artery initially turns, after passing the Foramen spinosum, in a posterior direction (anterior 
concavity). The point-by-point branching into tiny anterior and main posterior branch occurs 
within the area of the temporal lobe. The main branch runs to posterior, whereas, after further 
ramifications, the surface of the parietal lobe is almost entirely taken up with tiny branches 
running to anterior. The anterior branch supplies only a small (anterior) part of the surface of 
the parietal lobe and the posterior part of the frontal lobe. Between stages 1 and 2 a 
continuous transition is evident. (Holotype: Sangiran 2). 
Stage 3  
is characterized by the artery curving on the surface of the temporal lobe; the curvature can be 
described as slight, first to anterior, then to posterior. Within this surface area, a tiny branch 
splits off and runs to the anterior surface of the parietal lobe, while, somewhat more 
posteriorly, the main branch heads in a superior and posterior direction. It supplies the main 
surface of the parietal lobe with fine branches running to anterior. (Holotype: Zhoukoudian 
10). In 
Stage 4 
the anterior and posterior branches are nearly the same size and both of them – first the one, 
then shortly thereafter the other – split off from the main trunk of the mma. The posterior 
branch supports the blood supply to the surface of the parietal lobe from the rear, doing so 
from posterior to anterior, while the anterior branch supplies the surface of the precentral 
sulcus. (Holotypes: Reilingen, Salé, Arago). The  
Stage 5  
is distinguished from Stage 4 by the fact that an anterior branch has now become the main 
one. However, unlike in H.s.s., the course of this branch is posteriorly aligned, as too are all 
the other small branches diverging from the main branch. (Holotype: Vindija,). The latest 
developmental stage is reached with 
Stage 6,  
where the mma runs straight to anterior and along the coronal suture. In some specimens, it 
perforates the greater wing of the sphenoid at the sphenoparietal suture. The minor branches 
have split off within the surface of the parietal lobe and run to superior and posterior. 
Sometimes, very tiny branches split off posteriorly, close to the surface of the temporal pole. 
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(Holotype: in more than 1000 H.s.s. from prehistoric and historical times and in 31 modern 
cadavers). 
The excellent state of preservation of bony substance in the two new specimens (Sst IV and 
V)[3] led to us undertaking a precise examination of their meningeal arterial system. The 
developmental stage of mma at Sst IV (a partially preserved left parietal bone) is nearly 
identical to that of Sangiran 2 and corresponds best to our Stage 2. Moreover, the overall 
similarity with Pithecanthropus I (i.e., Trinil 2) is only somewhat undermined by a well-
developed sulcus of the sphenoparietal sinus, which superimposes (in part) the anterior 
arterial branch. In SstV (a completely preserved left temporal bone) the mma curves to 
posterior after passing the Foramen spinosum. This pattern is in exact agreement with Stage 1. 
This places the two fragments (Sst IV and Sst V) biostratigraphically in the earliest stage of 
the human phylogeny (with especial reference to Asia).  
Discussion 
The first point is that this survey of a phylogenetic model is independent from any absolute 
dating of fossil hominids. Second, our results are based on a new method of differentiating 
between the impressions of veins and arteries and between the impressions of main and side 
branches. Some earlier results seem to conflict[17,19]  with those presented here. A high degree 
of correlation[19] has been reported between the geometry of the neurocranium and the 
patterns of the mma in different groups of H.s.s. And it has been suggested that the differences 
among extinct hominids, as opposed to those within H.s.s., are related to the neurocranial 
geometry (“...low vault and posterior endocranial elongation[15,16]. These earlier results are 
only consistent with our observations regarding the pattern of the mma and the geometry of 
fossil hominid skulls. This do not, however, hold up with regard to the phylogenetic stage of 
the skulls – during our research, we were unable to observe any correlation between the 
geometry of the neurocranium and the phylogenetic stage. Our own observations, such as they 
were, led to a hypothetical phylogenetic model of the mma, as proposed in this paper.  
This evolutionary model demonstrates – we believe convincingly and definitively – a clear 
developmental process for mma within the hominid phylogeny [20]. Our model finely 
discriminates differences between specific fossil specimens including in the area of individual 
variability. 
Given that the chief function of the mma is to supply the surface of the parietal lobe with 
blood, as in the subspecies H.s.s., any changes to the position of this artery during the 
phylogeny of the genus Homo might be expected to hint at further developments within the 
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parietal lobe. This observation would signify, for instance, that the parietal lobe would shift to 
anterior during the phylogenetic process of the genus Homo. In tandem with this process, the 
frontal lobe would be shifted to anterior as well, thus reducing the flat frontal bone. In the 
specimen Sst IV, the negative of the central sulcus can be observed in the region where the 
anterior branch of the mma is positioned. This position is consistent with the supposition that 
the anterior branch of the mma will normally depend on that of the central sulcus or, in other 
words, the frontal part of the parietal lobe. 
Therefore, examination of the course of the mma enables precise reconstruction of the 
phylogenetic stage of fossil skulls (and even of fragments from the parietal bone). But there is 
a proviso: careful attention must be paid to a) the differences between arteries and veins and 
b) arteries that develop in the context of meningitis. With a view to the present: finds from 
Africa, say, need to be examined, or re-examined, in light of the phylogenetic development of 
the mma. The data obtained might provide additional indications regarding a possible 
discrepancy or concordance, as the case may be, between morphology and developmental 
stage of the mma.  
 
 
 
Abstract 
 
This study has a presupposition: while each skeletal element may be of pivotal importance in 
reconstructing human phylogenesis, none is more so than the cranium. Based on the recovery 
of relatively small fragments of Ossa parietalia and temporalia antedating the Elster glaciation 
(MIS 13 – 17), the question arises as to whether certain criteria are met for assigning them to 
different stages in the phylogenetic development of the human line. The impressions of the 
Arteria meningea media were such as to require closer examination of their pathways, the size 
of their diverse branches, and also the specifics of the arterial impressions (as opposed to the 
venous ones). To this end, the casts of a total of 12 crania and/or their endcasts were assessed, 
as were also the schemata developed by a number of different authors [10, 11, 15]. In addition, 31 
corpses of anatomically modern humans were dissected and examined. This led to the 
determination that it is essential to distinguish between arterial and venous impressions. This 
underscores[15] yet again that “a large amount of self-training and experience”, to quote one 
author, is indispensable when it comes to differentiating as exactly as possible between 
arteries and veins. 
 
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In light of the above, we were able to demonstrate (indeed this was our principal result) that 
the main branch of the Arteria meningea media in early human finds progressed continuously 
through time from posterior to anterior. In consequence, the side branches are aligned. Based 
on this progressive displacement, it was possible to distinguish six (6) stages of phylogenetic 
development, although no exact timeline could be assigned for any stage. Nor was this 
displacement from posterior to anterior an isolated occurrence: the Sulcus centralis too 
progresses in an anterior direction, thus displacing the Lobus frontalis in such a way that the 
Os frontale now stands upright. 
 
Acknowledgment 
 We are much indebted to K.W. Frangenberg (for his constructive discussion) and 
Mihnea Nicolescu (for his revision of our figures). 
 
 
 
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The phyogenetic principles of mma: A hypothetical biostratigraphic model

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The phyogenetic principles of mma: A hypothetical biostratigraphic model

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