The Place of Dali Cranium in Human Evolution
WU Xinzhi
2014, 33(04):
405-426.
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This paper presents a comprehensive comparative study between Dali cranium and other human fossils of Middle Pleistocene and those of later time.The non-metric and metrical features are classified into 9 categories as follows:
1.The features of Dali which are similar to other Middle Pleistocene humans(MPH) and quite different from those of early modern humans(EMH).
The brow ridges are robust and connect each other in glabellar region.
The bregma and vertex coincide in position.
There is angular turn on the occipital portion of mid-sagittal contour.
The cranial wall is thick.
The lacrimal fossa is shallow.
The left infraorbital fissure communicates with the infratemporal fossa through a downward passage instead of a horizontal one.
The infratemporal surface of sphenoid bone locates at a lower level than the orbital floor.
The auricular height of Dali cranium is 102mm. It falls within the variation range (v.r.) of Homo erectus (H.e.) from Zhoukoudian (ZKD) (93.5mm~107mm)[9] and shorter than that of Kabwe (105mm) [9] and Neanderthals (103-122 mm, totally including 7 cases, the sites from which the fossils were enearthed can be checked in the references cited, similarly hereinafter) [9-10]and EMH of China (108~119 mm, including Liujiang, Lijiang, Chuandong and Upper Cave) [11-14].
The length-height index I (ba-b/g-op) of Dali cranium is 57.1. It falls within the v.r. of Plio/Pleistocene humans of Africa (50.4~67.5, including KNM-ER1813; OH 9, KNM-ER 3733, 3883) [5, 15] and that of Dmanisi (55.4~65.4) [16]. The value of Dali is slightly higher than that of the recontructed skull of Yunxian 55.8[16], and lower than that of the reconstructed skull of H.e. from ZKD (59.9)[9] and that of Kabwe (60.2) [15] as well as that of most of MPH of Europe (58.6~69.9, including Petralona, Steinheim, Swanscombe; Ceprano; Atapuerca SH 4,5,6) [15-17] except Ehringsdorf (the index is 55.9)[15]. It is much lower than that of EMH of China (66.7~77.7, including Liujiang and Upper Cave) [11,12] and Europe (65.8~71, including 6 cases) [10].
The length-height index II (po-b ht/g-op) of Dali is 49.6. It falls within the v.r. of that of Dmanisi (46.38~53.59[16]) and Plio/Pleistocene humans of Africa (46.6~53.3, including OH 9, KNM-ER 3733, 3883) [15]. The value of Dali is close to the lower limit of H.e. from ZKD (49.0~53.3) [9]. It is lower than that of H.e. from Hexian (50) [18] and close to the lower limit of the v.r. of MPH of Europe (51.0~65.1, including Petralona, Steinheim; Arago, Ceprano; Atapuerca SH) [15,16,17]. The index of Dali is much lower than that of EMH of China (57.1~72.5, including Liujiang, Upper Cave, Lijiang and Chuandong) [11,12,13,14] and Europe (58~62.3, including 6 cases) [10].
Transverse cranial curvature (au-au/po-b-po) of Dali cranium is 47.2. It is lower than that of H.e.of China (47.4~54.8) [9,18,19] and is between two specimens of Pithecanthropus from Trinil ( I, 52.3 and II, 45.6) [9]. It is lower than that of Kabwe (48.3) [9] and higher than that of Petralona (44.8) [10]. The value of Dali is much higher than that of EMH of China (39.2~42.2, measured by the author on Liujiang and casts of Upper Cave) and Europe (36.5~43.6, including 3 cases)[10].
The ratio of b-ast to occipital breadth (ast-ast) of Dali cranium is 113.9. It is within the v.r. of H.e. of China (103.5~119.8, including those of Hexian and cast of ZKD which are all measured by the author) and MPH of Europe (97.8~117.5, including Atapuerca SH, Petralona and Swanscombe) [17]as well as that of MPH of Africa (107.1~116.3, including Kabwe; Eliye Springs, Omo 2)[17, 20]. It is much lower than those in EMH of China (121.5~132.7, including those of Liujiang, Ziyang and casts of Upper Cave which are all measured by the author).
The angle, l-i-o of Dali is 105°. It is close to the highest value of the v.r. of H.e. of China (98°~106°, those of ZKD are according to Weidenreich 1943[9]; that of Nanjing is measured by the author) and lowest value of that of MPH of Europe (107°~129.1°, including Ehringsdorf, Steinheim ; Atapuerca SH) [10, 17]. It is much lower than that of modern humans (117°~127.3°) [10].
Angle l-op-o of Dali cranium is 98°. It equals the lowest value of the v.r. of H.e. of China (98°~108°, including ZKD and Nanjing) [16, 19],much lower than that of Petralona (106°)[16] and much lower than that of modern humans (128°~138°) [16].
The ratio of d-d to fm: a-fm: a of Dali cranium is 22.8. It is between the values of H.e. from ZKD XII (21) and Nanjing (26)(those of ZKD and Nanjing are measured and calculated by the author) and very close to that of Kabwwe (22.6, measured by the author on cast). The value of Dali is much lower than that of Petralona(28.7) [21]and Atapuerca SH 4 and 5 (33.1 and 29.5 respectively) [17] and higher than that in EMH of China (15.5~18.6, measured by the author on fossil from Liujiang and casts of the fossils from Upper Cave).
2. Features similar or close to that in modern humans
The contour in hind view appears as an even curve with the broadest part locating at the temporal squama.
The vertical line passing the most lateral point of mandibular fossa passes through the lateral cranial wall lateral to the joining point between the inner surface of cranial base and lateral cranial wall.
The ratio of calvarium height above g-i chord to g-i chord of Dali cranium is 50.4. It is much higher than those in H.e. from ZKD(34.8~41.2)[9], Pithecanthropus from Trinil(33.3~37.4)[9] that of Kabwe (40.5) [10], Saldanha (45.0) [10] and Jebel Irhoud (43.7) [10], and higher of that of Steinheim (46) [10]. The value of Dali falls into the v.r. of EMH of Europe (49-61, including 5 cases) [10].
The ratio of arc n-i to n-i chord of Dali cranium is 189.9. It is higher than that of Neanderthals (5 cases, 145.1~178.1) [10], between that of male (181.2) and female (203.7) of Obercassel[10]. Dali’s value is close to the average of Middle Age Japanese (200.0) [10].
Angle b-n-op (i) of Dali is 54°. It is much higher than that of H.e. from ZKD (42°~46.5°) [9], Kabwe(48°) [9] and Neanderthals (including 5 cases, 39°~50°) [9]. It is higher than that of Ehringsdorf (52°)[9] and falls into the v.r. of modern humans (45°~59°)[9].
Angle g-i-l of Dali cranium is 82°. It is much higher than that t of H.e. of China (57°~68°)(those of ZKD are from Weidenrech[9]; that of Nanjing is measured by the author), Ehringsdorf (63°) [10], Kabwe (68°) [10], and Neanderthals of Europe (including 7 cases, 59°~69°) [10]. Dali’s value falls within the v.r. of modern humans (80.2°~88.6°) [10].
Upper facial height (fmt-fmt) of Dali cranium is 121mm. It is much longer than those of H.e. from ZKD(III, 109mm; XI, 111mm?)[16] and Nanjing (107mm,measured by the author) and Hexian (113mm). Dali’s value is very close to that of Maba(126mm) and Upper Cave 101(122mm), and much longer than those in Upper Cave 102 and 103 (113mm and 100mm respectively) as well as that in Liujiang(107mm)(All specimens of China except those from ZKD are measured by the author). Dali’s value is shorter than MPH of Europe(125~130mm, including Arago, Atapuerca SH5, Petralona and Ceprano)[16], and much shorter than those from Bodo(136mm) [16]and Kabwe(139mm)[16].
The depth of facial bones (ba-pr) is 105mm. It is much shorter than that of Atapuerca SH 5 (115mm) [16], Petralona (119 mm) [16], Bodo (118 mm) [16] and Kabwe (117.5mm) [16]. Dali’s value falls within the v.r. of EMH of China (100~113.6 mm, including Liujiang and Upper Cave) [11, 12] and is close to the average of that of modern humans (including 60 cases, 97.2 mm) [16].
The ratio n-ba/ba-pr of Dali cranium is 100.5. It is much higher than that in Bodo (88.4) [16]; Kabwe (93.1) [16], Atapuerca SH 5(87.8) [16] and Petralona (94.8) [16]. It falls within the v.r. of EMH of China (99.3~112.0 , including Liujiang and Upper Cave) [11, 12] and is close to the average of modern humans (101.7) [16].
Cheek height(WMH) of Dali cranium is 23mm. It is within the v.r. of early modern humans of China(21.7~27.2mm, including 7 sides of 4 cases of Upper Cave and Liujiang, measured by the author). Dali’s value is shorter than that in H.e. from ZKD(XII, 28mm, measured by the author), and those in MPH of Europe(26.7~37.1mm, including 7 sides of 6 cases) [17]. It is close to the H.e. from Nanjing(24.3mm, measured by the author) and Zuttiyeh(24mm)[17], but the dimensions of H.e. from Nanjing are generally shorter than H.e. from ZKD.
The prosthion angle (n-pr-ba) of Dali cranium is 69.5°. It is very close to the average of that of modern humans (71.4°±3.1°) [24] and much higher than that in Kabwe ( 62.1°) [24], Bodo( 59°, calculated by the author based on the data in Rightmire, 1996[23]), Atapuerca SH 5 (60.9°) [17] and Petralona (62.0°) [24].
3. Features far from that in H.e. of China and within the v.r. of that in EMH of China and /or modern humans. These features are also close to that in MPH of Europe and/or Africa
Dali cranium has no supraorbital process.
The maximum cranial length (g-op) is 16.5 mm longer than glabella-inion length (g-i) in Dali cranium. The difference between these two measurements is very short or none in H.e. of China and is between 4 mm and 17 mm in EMH of China (including Liujiang[11] ,Upper Cave[12], Lijiang[13] and Chuandong 2[14]). The difference is larger than 5 mm in Petralona[15], Steinheim[15], Jebel Irhoud [10] and Narmada[15].
Transverse fronto-parietal index (100 x ft-ft/eu-eu) is 69.6 in Dali cranium. It is higher than that of H.e. of China (55.9~64.5, including ZKD, Hexian, and Nanjing) [9, 18, 19, 25] and falls in the v.r. of EMH of China (66.9~77.1, including Liujiang , Upper Cave and Lijiang)[11, 12, 13]. Dali’s value is higher than that of Kabwe(64.3) [15] and Salé (57.5) [15]. It is within the v.r. of MPH of Europe (67.0~77.9, including Ehringsdorf, Steinheim; Arago, Ceprano, Petralona; Atapuerca SH) [15,16,17].
The total cranial arc (n-o arc) is 379 mm in Dali cranium. It is much longer than those in H.e. from ZKD (321 mm~337 mm)[9] and H.e. from Hexian (340 mm?) [18]. It falls within the v.r. of EMH of Chna (335 mm~388.5 mm, including Liujiang, Upper Cave, Lijiang, Chuandong and Ziyang) [11-14, 28] and that of MPH of Europe and Africa (340 mm~380 mm , including Ehringsdorf [9]Atapuerca SH, Petralona, and Kabwe [9]) (the values of Atapuerca SH and Petralona are calculated by the author based on the data presented in Arsuaga et al. [17] and Stringer et al. [21],respectively).
The cranial curvature on the n-o chord is 37.7 in Dali cranium. It is lower than that of H.e. from ZKD (43.2-44.9) [9], Nanjing (48.8) (by the author)and Hexian (38.5) [18]. It falls within the v.r. of EMH of China (36.4~40.3, including Liujiang , Upper Cave and Ziyang ) [11, 12, 28] and v.r. of modern humans (35.2~39.9) [9]. Dali’s value is close to the average of that of modern humans (36.6) [9], Kabwe (37.1) [9]and slightly lower than that of Ehringsdorf (40.1) [9].
The ratio of maximum frontal breadth to occipital breadth (co-co/ast-ast) is 103.5 in Dali cranium. It is much higher than those of H.e. of China (including ZKD, Nanjing and Hexian: 83.9~99.1?)(ZKD is from Weidenreich, 1943[9], Nanjing and Hexian are by the author) and that of Kabwe(90.5, calculated by the author based on Weidenreich, 1943[9]). Dali’s value falls within the v.r. of MPH of Europe (93.6~108.8, including Arago, Atapuerca SH 4 and 5, Petralona, Steinheim and Swanscombe) [29]. Dali’s value falls within the v.r. of EMH of China (including Upper Cave, Liujiang and Ziyang: 100~114, by the author) and is close to the average of Mesolithic humans of Europe (male, 102.9; female, 103.3) and Sepúlveda population (male, 103.0; female, 105.6)[29].
The minimum frontal breadth of Dali cranium is 104 mm. It is much longer than those of H.e.of China (80.0 mm~93 mm including ZKD, Hexian and Nanjing) [9, 18, 19] and falls within the v.r. of EMH of China (83 mm~110 mm , including Upper Cave, Liujiang, Ziyang, Longlin and Maludong) [1, 12, 28, 30]. It falls also in the v.r. of EMH of Europe (91 mm~111 mm) [30] and West Asia (96 mm~110 mm) [30]. The mean values of last two groups are 105±5mm and 103±5 mm respectively). Dali’s value falls also in the v.r. of MPH of Europe (102 mm~117 mm , including Arago, Steinheim; Ceprano, Petralona ; Atapuerca SH) [15, 16, 17]and is close to that of Bodo (103 mm) [16] , but is longer than that of Kabwe (96) [16].
The ratio of minimum frontal breadth to maximum frontal breadth (ft-ft/co-co) is 87.4. It is higher than those of H.e. of China (77.8~84.3, including ZKD [9], Nanjing [19] and Hexian which is measured and calculated by the author)and falls within the v.r. of MPH of Europe (86.1~100, including Arago, Petralona, Steinheim; Ceprano , Atapuerca SH,) [15, 16, 17] and that of African MPH ( 78.3~89.6, including Bodo, Kabwe and Salé) [16, 31]. It falls also within the v.r. of EMH of China (76.0~90.5, including Upper Cave, Liujiang, Ziyang, and Maludong which is from Curnoe et al., 2012[30]; U.C., Liujiang and Ziyang are measured and calculated by the author).
The bistaphanic breadth (st-st) of Dali cranium is 108 mm. It is longer than those of H.e. of China ( 78mm~103mm , including ZKD [9], Nanjing and Hexian, latter two are measured by the author), and falls within the v.r. of EMH of Chian(105~119.5, including Upper Cave 101, 103, Liujiang and Ziyang, all of these are measured by the author). The mean value of modern humans (110.42mm) [16] is close to that of Dali’s value which falls also within the v.r. of MPH of Europe (102mm~130mm, including Arago, Petralona, Ceprano; Atapuerca SH) [16, 17].
Length-height index of temporal squama is 64.6 in Dali cranium. It is higher than those in H.e.of Chinna (45.2~60, including ZKD and Hexian) [9, 18], and slightly lower than those in Atapuerca SH (including 5 cases, 69.3~79.7) [32]. Dali’s value is close to the average of that of modern man (65.2) and well within his v.r. (49.4~87.5). [9]
The frontal profile (angle m-g-i) of Dali cranium is 74°. It is much larger than those of H.e. of China (56°~63°,including ZKD, Hexian, and Nanjing ) [9, 18, 19], Pithecanthropus from Trinil (47°and 55°) [9] and MPH of Africa (including Jebel Irhoud, 67°[10]; Saldanha, 61°[10]; Kabwe, 60°[9]). Dali’s value is very close to that of Ehringsdorf (73.5°) [9] and falls in the v.r. of mordern man (70°~96°)[9].
4. Features intermediate between MPH and EMH
Dali cranium has a weak bregmatic eminence, a fusiform median sagittal ridge at the middle part of frontal bone, and a tympanic plate the thickness of which is intermediate between that of H.e.from ZKD and modern man.
An angular torus presents at the outer surface of parietal bone.
Tranverse cranial curvature (au-au / arc po-b-po) of Dali cranium is 47.2. The comparsions between Dali cramium and other specimens have been persented among the features of Group 1.
5. Features intermediate between H.e. and EMH of China and close to those in MPH of Europe and/or Africa
The cranial curvature above chord n-op of Dali cranium is 51.8. It is much lower than those in H.e. from ZKD (55.7~57.3) [9], slightly lower than that in H.e. from Hexian (52.9, measured by the author). It is slightly higher than the upper limit of the v.r. of EMH of China (including Upper Cave, Liujiang, and Ziyang, 47.2~51.0, by the author) and is well within the v.r. of MPH of Europe (49.5~54.4, including Ehringsdorf[9], Atapuerca SH[17] and Petralona[21]). That of Kabwe (54.2) [9] is also within the v.r. of MPH of Europe.
The ratio of maximum frontal breadth to maximum cranial breadth (co-co/eu-eu) is 79.3 in Dali cranium. It is intermediate between those in H.e. of China (68.5~76.9, including ZKD [9], Nanjing and Hexian which are measured and calculated by the author) and those in EMH of China (79.7~93.1, including Upper Cave, Liujiang and Ziyang, measured by the author). It falls in the v.r. of MPH of Europe (Atapuerca SH: 75.0~87.9) [17] and v.r. of Neanderthals of Europe (including 11 cases, 75.5~83.4) [29]. It is much lower than the averages of early Upper Paleolithic man of Europe (male: 87.9; female: 85.2) [29]and within the v.r. of Sepúlveda population (male, 41 cases: 72.9~90.1; female, 57 cases: 77.7~90.8) [29].
6. Features close to those in fossil humans of East Asia and distant to those of MPH of Europe and / or Africa
The median sagittal ridge of frontal bone is higher and has narrower base than those in MPH of Europe and Africa.
The ratio of nasion subtense fraction of n-b chord to n-b chord is 43.5 in Dali cranium. It is slightly lower than those in H.e. from ZKD(45.3~48.7) [16] and slightly higher than that of H.e from Nanjing(42.4, by the author). It falls in the v.r. of EMH of China ( 43.0~46.0, including Upper Cave and Liujiang, all are measured and calculated by the author). On the contrary, Dali’s value is much lower than those in MPH of Europe (including Petralona, 50.0[16]; Arago, 49.6, based on the data presented by de Lumley[16]) and is lower than those in MPH of Africa (44.9~54.1, including Laetoli OH18, Eliye Springs , Omo 1, Jebel Ithoud 1 and 2 and Singa) [3].
The occipital angle of Dali cranium is 96°. It is within the v.r. of H.e. of China (including ZKD and Nanjing, 95°~108°, measured on the Figures in Weidenreich[9] and Wu et al. [19] respectively) and is much lower than those of Atapuerca SH (106.5°~126.1°) [17]. Dali’s value is also much lower than those of Plio/Pleistocene huamns of Africa( 101°~114°, including KNM-ER 3883, 3733, 1813?) [34], Dmanisi (115.6° and 108°) [34] and Sangiran (105° and 100°) [34].
The anterior interorbital breadth (mf-mf) of Dali cranium is 21.5 mm. It is close to that in EMH of China (including Upper Cave and Liujiang: 19.1mm~21.2mm, by the author), H.e. from ZKD(No.XII, 22.5mm), Nanjing(19mm) and slightly shorter than that of EMH of Europe (including 7 cases: 23.4±2.9 mm) [22]. Dali’s value is much shorter than that in MPH of Europe, Africa and West Asia (including 5 cases, 29.5±2.2 mm) [22].
The upper facial index (n-pr/zy-zy) of Dali cranium is 53.2. It is close to that of Jinniushan(50.1)[26], H.e. of Nanjing (49.9) [19] and ZKD XII (54.5, measured by the author on cast). Dali’s value is within the v.r. of EMH of China(48.5~53.8) [11, 12] , but is lower than those in MPH of Europe (56.0~59.0, including Petralona, Steinheim; Atapuerca SH) [10, 17] and Africa (54.2~64.7, including Jebel Irhoud, Kabwe; Bodo) [10, 35].
The distance between infraorbital foramen and inferior border of orbit is 8.3 in Dali cranium. It is close to that in H.e. of Nanjing (7.5 mm, measured by the author) and much shorter than those in Atapuerca SH (14.1 mm~17.7 mm) [17] and Petralona (16.4 mm) [17].
The ratio d-d to fm:a of Dali cranium is 22.8 mm. The comparisons between Dali cranium and other specimens have been presented among the features of Group 1.
7. Features quite different from those in H.e. of China and are close to those in MPH of Europe and/or Africa
In Dali cranium supraorbital process is absent, the middle part of the supraorbital torus is much thicker than the medial and lateral part, and there is a bulge between the orbit and pyriform aperture.
The l-ast chord of Dali cranium is 94 mm. It is within the v.r. of MPH of Europe (74.5 mm~95.6 mm , including 10 cases, 17 sides of Atapuerca SH and Petralona) [17, 21] and much longer than those in H.e. of China (77 mm~87 mm, including ZKD[9] and Hexian which is measured by the author).
The bifrontal breadth (fm: a-fm: a) of Dali cranium is 114 mm. It is much longer than the MPH of China (96 mm~104 mm, inluding H.e. from ZKD, Naanjing and Hexian, Maba , all of these are measured by the author) and close to those in MPH of Europe, Africa and West Asia ( mean value of 6 cases: 114.7±8.5) [22].
The EQ of Dali cranium, average of H.e. from ZKD and MPH of Europe and Africa are 5.30, 4.6 and 5.3±1.29 respectively, as estimated by Rightmire[36].
8. Feature close to those in MPH of Africa and distant from other MPH of China and Europe
The ratio of glabella subtense fraction of g-b chord to g-b chord is 43.4 in Dali cranium. It is lower than those of H.e. from ZKD (47~50.9, measured and calculated by the author based on the figures in Black[37] and Weidenreich[9]), Nanjing (49.7, by the author ) and Maba (45.1, by the author). But it is within the v.r. of EMH of China (41~48.5, including Huanglong[40], Upper Cave, Liujiang and Ziyang, that of the latter three are made by the author), Dali’s value is within the v.r. of MPH of Africa (42.6~58.5, including Florisbad, Jebel Irhoud, Kabwe, Laetoli, Omo 1 and Saldanha) [3]. But it is much lower than those in Arago (51, calculated by the author based on the cast and the data presented in Spitery [38]), and Ceprano (60.8, measured and calculated by the author based on the figure presented in Ascenzi et al [39]).
9. Features rarely seen in other MPH or uniquely seen in Dali
A quadrangular shaped process of the size of 10×7 mm, extending from the antero-superior part of temporal squama and connects with frontal bone. This makes the sutures in pteryon region obliquely posited ∏ shape.
The crista galli is thin and low with a large transverse diameter. The angle g-i-o of Dali is 21°. It is much smaller than those in H. e. from ZKD (37°~44°,measured by the author on the figures presented in Weidenreich [9]) and Nanjing (49°, measured by the author on cast) , Neanderthals of Europe and Asia (31°~54°, including 7 cases) [10] and modern man (31°~40°) [10].
Discussion and conclusion
The features including in Group 1 confirm the position of Dali cranium in Middle Pleistocene. Features of Group 2 show that this cranium is one of the specimens closer to EMH than H.e. if not also MPH of Europe and Africa. Part of the features of Group 2, 3, 4 and 5 suggest that thepopulation represented by Dali cranium provides more contribution to the formation of modern man than H.e. of China, if not also the MPH of west part of Old World. The features of Groups 1 and 2 and some features of other Groups indicate that Dali cranium represents a mosaic with primitive and progressive characters. Features of Group 6 and part features in Group 1, 2, 3 and 5 suggest close relation of Dali cranium to populations of East Asia. Many features presented in Groups 2, 3, 4, 5, 7, 8 and 9 indicate that Dali cranium probably belong to a population different from H.e. Thhe features of Group 3, 5, 7, 8 and part of Group 4 suggest that Dali may have close relation with the populations in the western part of Old World. So Dali cranium is a mosaic joining some features of H.e. of China, MPH of Europe and Africa as well as some modern features. This cranium belongs to neither H.e., nor H.heidelbergensis. The population represented by Dali cranium have made more contribution in the formation of EMH of China than H.e. of China and MPH of Africa. Considering the geographical factor and the association with the Paleoliths of Mode I of Dali cranium, more reasonable inference may be that the root of the population represented by Dali cranium is in East Asia and the antecessors of this population originally had higher affinity with the populations of Europe or had absorbed the gene flow from the West before evolving to Dali population.
With regards to the relation between the humans of Middle and Late Pleistocene Rightmire (1995)[41] has proposed four hypotheses, the evidence exhibited in the present paper are more favourable to the fourth hypothesis of him, namely the recent humans are most closely related with Archaic Asians.
In spite of the limitation of the data for comparison the present author would like to say that the complexity of the morphology shown in Dali cranium suggeats that the human evolution in East Asia is not as simple as we thought until present. The Middle Pleistocene humans may be classified into several morphs: Dali morph, erectus morph, Narmada morph, Zuttiye morph, Rhodesia morph (for Africa), Heidelberg morph (for Europe) etc. Hexian specimens may represent a submorph of erectus morph, Jinniushan, Maba and Quyuan River Mouth may represent separate submorphs of Dali morph or separate morphs for themselves. Atapuerca SH and Arago may represent separate submorphs. The model of human evolution in Middle Pleistocene is like a river network.
The above mentioned comparisons are based on limited information available. The author looks forward to the accumulation of new data to renew the preliminary conclusion based on this study.
