TABLE 1. Measurements and proportions of specimens of Armigatus carrenoae sp. nov. from the Tlayúa quarry referred here. All the measurements are given in millimeters, and all percentages are calculated on the base of the standard length (SL).
| IGM 11536 | IGM 11537 | IGM 11538 | Measurements (in mm) and proportions IGM 11539 |
IGM 11540 | IGM 11541 | IGM 1542 | IGM 11543 | Avg. (Range) | |
| Total length | 57.2 | 68.3 | 57 | 81.9? | 35.3 | 57.5 | 56.6? | 68.3? | 55.6 (35.3-68.3) |
| Standard length | 42.7 | 50.1 | 42.4 | 64.2? | 49.7 | 46.0 | 46.9? | 51.8 | 47.1 (18.4-51.8) |
| Body height % SL |
23.0 53.8 |
23.7 47.3 |
22.7 53.5 |
27.3 42.5? |
30.6? 61.5? |
21.7 47.1 |
25.3? 53.4? |
25.3? 53.4? |
24.4 (23-28.2) 51.2 (47.1-54.4) |
| Head length % SL |
17.2 40.2 |
18.2 37.5 |
16.7 39.3 |
22.5 35? |
18.4 37.0 |
16.5 35.8 |
17.8? 37.9? |
18.4? 35.5? |
18.25 (18.4-22.5) 37.9 (35.8-40.2) |
| Predorsal length % SL |
24.2 56.6 |
28.5 56.8 |
24.4 57.5 |
34.9? 54.3? |
28.2 56.7 |
25.8 57.8 |
27.6? 58.8? |
28.4 54.8 |
26.5 (24.2-28.5) 56.7 (54.8-57.8) |
| Dorsal fin length % SL |
7.0 16.3 |
8.2 16.3 |
6.5 15.3 |
9.5? 16.2? |
7.4 14.8 |
8.0 17.3 |
7.6 16.2? |
8.3 17 |
7.4 (6.7-8.3) 16.1 (14.8-17.3) |
| Prepelvic length % SL |
25.2 59 |
30.3 60.4 |
25.2 59.4 |
35? 54.5? |
29.6 60.2 |
26.6 57.8 |
26? 55.4? |
29.4 56.8 |
27.7 (25.2-30.3) 58.9 (56.8-60.4) |
| Preanal length % SL |
33.7 78.9 |
38.5 76.8 |
33.4 78.7 |
52.6? 81.9? |
40.0 80.4 |
35.3 76.7 |
37.5? 79.9? |
42.2 81.4 |
37.1 (33.4-42.2) 78.8 (76.7-81.4) |
| Anal fin length % SL |
8.1 18.9 |
8.3 16.5 |
7.3 17.2 |
10.9 16.9? |
8.5 17.1 |
8.4 18.2 |
7.8 15.3? |
? ? |
8.5 (7.3-10.9 17.5 (16.5-18.9) |
| Meristics | Mode (Range) | ||||||||
| Total vertebrae | 35 | 33 | 34 | 34 | 35 | 35 | 33+ | 29+ | 34 (33-35) |
| Abdominal centra | 17 | 16 | 15 | 16 | 16 | 16 | 14+ | 10+ | 16 (15-17) |
| Caudal centra | 16 | 15 | 17 | 16 | 17 | 17 | 17 | 17 | 17 (15-17) |
| Pair of ribs | 14 | 15 | 15 | 15 | 14 | 14 | 15 | ? | 14-15 |
| Predorsal scutes | 8 | 5+ | 7? | 3+ | 3+ | 4+ | 5+ | 3+ | 8 |
| Abdominal scutes | 22? | 23 | 22 | 18+ | 22? | 21+ | 23 | 5+ | 23 (22-23) |
| Prepelvic scutes | 16? | 17 | 16 | 12+ | 16? | 15+ | 17 | 5+ | 17 (16-17) |
| Postpelvic scutes | 6 | 6 | 6 | 6 | 6 | 6 | 6 | ? | 6 |
| Dorsal fin rays | ii, 11 | i, 11 | ii, 11 | ii,? | ii, 9? | ii,13 | ii, 11 | ii,11 | ii,11 (i-ii,11-13) |
| Dorsal pts. | 11 | 11 | 11 | 11 | 10+ | 12 | 9+ | ? | 11 (11-12) |
| Anal fin rays | ii, 17 | ii, 15 | ii, 17 | ii, 17 | I, 14? | ii, 17 | ?, 5+ | ii, 17? | ii, 17 (ii,15-17) |
| Anal pts. | 17 | 15 | 16 | 15 | 14+ | 17 | 14+ | 12+ | 17 (16-18) |
| Pectoral fin rays | 15 | 15 | 14+ | 13+ | 14 | 5+ | 14 | 15 | 15 (14-15) |
| Pelvic fin rays | 6 | 5 | 5 | 5 | 5? | 5 | 7 | 6+ | 5(5-7) |
| Predorsal bones | 6 | 6 | 6 | 6 | 6 | 6 | 6 | 5? | 6 |
TABLE 2. Nominal species currently included in the order Ellimmichthyiformes.
| Species | Age and Distribution |
| Armigatus alticorpus Forey, Yi, Patterson, and Davies, 2003. | Cenomanian. Namoura, Lebanon. |
| Armigatus brevissimus (Blainville, 1818). | Cenomanian. Heckel and Hajula, Lebanon. |
| Armigatus carrenoae sp. nov. (present manuscript). | Albian. Tlayúa quarry, Mexico. |
| Armigatus dalmaticus Murray, Vernygora, Japundžić, Radovčić, Wilson, Bardack, and Grande, 2016. | Campanian? Dalmatia, Croatia. |
| Armigatus namourensis Forey, Yi, Patterson, and Davies, 2003. | Cenomanian. Namoura, Lebanon. |
| Armigatus oligodentatus Vernygora and Murray, 2016. | Cenomanian/Turonian. Agoult, Morocco. |
| Codoichthys carnavalii Silva Santos, 1994. | Aptian. São Luís-Grajaú Basin, Brazil. |
| Diplomystus altisomus Yabumoto, 1994 | Early Cretaceous. Kyushu, Japan. |
| Diplomystus birdi Woodward, 1895. | Cenomanian. Heckel and Hajula, Lebanon. |
| Diplomystus dentatus Cope, 1877. | Eocene. Green River Fm., USA. |
| Diplomystus dubertreti Signeux, 1951. | Santonian. Sahel Alma, Lebanon. |
| Diplomystus kokuraensis Uyeno, 1979. | Neocomian. Kyushu, Japan. |
| Diplomystus primotinus Uyeno, 1979. | Neocomian. Kyushu, Japan. |
| Diplomystus solignaci Gaudant and Gaudant, 1971. | Senonian. Gabès, Tunisia. |
| Diplomystys shengliensis Zhang, Zhou, and Qing, 1985. | Eocene. Shandong, China. |
| “Diplomystus” trebicianensis Bannikov and Sorbini, 2000. | Paleocene, Trebiano, Italy. |
| Ellimma branneri (Jordan, 1910). | Hauterivian-Aptian. Muribeca Fm., Brazil. |
| Ellimma cruzi Silva Santos 1990. | Aptian. Cabo Fm., Brazil. |
| Ellimma longipectoralis Polck, Gallo, Figueiredo, Viana, Santos, Quiroz-Neto, and Jahnert, 2020. | Aptian. Velha Fm., Santos Basin, Brazil. |
| Ellimmichthys longicostatus (Cope, 1886). | Hauterivian-Barremian. Recôncavo Fm., Brazil. |
| Ellimmichthys goodi Eastman, 1912. | Aptian-Albian. Cocobeach Series, Equatorial Guinea. |
| Ellimmichthys maceioensis Malabarba, Do Carmo, Gómez-Pérez, and Queiroz-Neto, 2004. | Aptian-Albian. Maceio Fm., Brazil. |
| Eoellimmichthys superstes Marramà, Bannikov, Kriwet, and Carnevale, 2019. | Early Eocene. Pesciara, Italy. |
| Ezkutuberezi carmenae (Poyato-Ariza, López-Horgue, and García-Garmilla, 2000). | Valanginian-Barremian. Villaro Fm., Spain. |
| Foreyclupea loonensis Vernygora, Murray and Wilson, 2016. | Albian. Loon River Fm., Canada. |
| Gateroclupea branisai Signeux, 1964 (see Marramà and Carnevale, 2017). | Late Cretaceous-Paleocene. Cayara, Bolivia. |
| Horseshoeichthys armaserratus Newbrey, Murray, Brinkman, Wilson, and Neuman, 2010. | Maastrichtian. Horseshoe Canyon Fm., Canada. |
| Kwangoclupea dartevellei (Casier, 1965). | Cenomanian. Kwango, Zaire. |
| Ornategulum sardinoides (Pictet, 1850). | Cenomanian. Heckel and Hajula, Lebanon. |
| Paraclupea chetungensis Sun, 1956. | Early Cretaceous. Chawan Fm., China. |
| Paraclupea seilacheri Alvarado-Ortega and Melgarejo-Damián, 2017. | Albian. Tlayúa, Mexico. |
| Ranulfoichthys dorsonudum Alvarado-Ortega, 2014. | Albian. Tlayúa, Mexico. |
| Rhombichthys intoccabilis Khalloufi, Zaragüeta-Bagils, and Lelièvre, 2010. | Cenomanian. Ein Yabrud, Palestine. |
| Scutatoclupea bacchiai Bannikov, 2015. | Cenomanian. Namoura, Lebanon. |
| Scutatuspinosus itapagipensis Santos and Corrêa, 1985. | Hauterivian?. Marfim Fm., Brazil. |
| Sorbinichthys elusive Bannikov and Bacchia, 2000. | Cenomanian. Namoura, Lebanon. |
| Sorbinichthys africanus Murray and Wilson, 2011. | Cenomanian/Turonian. Agoult, Morocco. |
| Thorectichthys rhadinus Murray and Wilson, 2013. | Cenomanian/Turonian. Agoult, Morocco. |
| Thorectichthys marocensis Murray and Wilson, 2013. | Cenomanian/Turonian. Agoult, Morocco. |
| Triplomystus applegatei Alvarado-Ortega and Ovalles-Damián, 2008. | Cenomanian. El Espinal, Mexico. |
| Triplomystus noorae Forey, Yi, Patterson, and Davies, 2003. | Cenomanian. Namoura, Lebanon. |
| Triplomystus oligocostatus Forey, Yi, Patterson, and Davies, 2003. | Cenomanian. Namoura, Lebanon. |
| Tycheroichthys dunveganensis Hay, Cumbaa, Murray, and Plint, 2007. | Cenomanian. Dunvegan Fm., Canada. |
| Tunisiaclupea speratus Boukhalfa, Wu, Ben Ali, and Fang, 2019. | Late Barremian. Chotts Basin, Tunisia. |
TABLE 3. Comparison of the nominal species of the genus Armigatus. Maximum standard length (SL) expressed in millimeters, (based on data from table 3; Forey et al., 2003; Murray et al., 2016, tables 2, 3; Vernygora and Murray, 2016, table 1; Vernygora and Murray, 2021).
| Species→ Number of individuals considered→ |
Armigatus carrenoae sp. nov. (8) |
Armigatus brevissimus (10) |
Armigatus namouraensis (11) |
Armigatus alticorpus (6) |
Armigatus oligodentatus (8) |
Armigatus dalmaticus (6) |
|
| Averages of proportions expressed as % of SL (ranges) |
Maximum TL | ≈ 81.9 | 105 | 139 | 47 | 37 | 48.6 |
| Body height | 51.2 (47.1-54.4) |
37.3 (33-42) |
37.7 (31-50) |
48.9 (46.1-52.7) |
32.1 (28-36) |
40.8 (34.6-44-8) |
|
| Head length | 37.9 (35.8-40.2) |
33.6 (30-38) |
31.9 (28-35) |
34.2 (33.3-36.1) |
32.5 (31-35) |
31.3 (28-33) |
|
| Predorsal length | 56.7 (54.8-57.8) |
46.5 (44-49) |
47.3 (43-52) |
49.2 (47.9-51) |
48.6 (46-52) |
46.3 (43.3-50.2) |
|
| Preanal length | 78.8 (76.7-81.4) |
67.1 (64-72) |
69.1 (66-77) |
69.9 (66.6-73.7) |
77.8 (75-82) |
75.4 (66.5-81.9) |
|
| Prepelvic length | 58.9 (56.8-60.4) |
50 (47-56) |
51.5 (49-56) |
53.6 (55.5-50) |
55.2 (50-58) |
56.4 (55.4-57.8) |
|
| Modes of accounts (ranges) |
Anal fin rays | ii, 17 (ii,15-17) |
28 - |
25 (24-27) |
25+ (?-25+) |
? (12?-14?) |
16 - |
| Dorsal fin rays | ii,11 (i-ii,11-13) |
19 - |
19 (18-19) |
16 (15-17) |
11 (10?-13?) |
13, 14 (13-14) |
|
| Pectoral fin rays | 15 (14-15) |
20 (18-20) |
12 (12-15) |
12 or 13? (12-13) |
8 (8-13) |
? (12-13) |
|
| Pelvic fin rays | 5 (5-7) |
7 - |
8 (8-9) |
? - |
7 (6-9) |
7? - |
|
| Predorsal bones | 6 (6) |
6 (6) |
6 (5-6) |
6 (5-6) |
8 (7?-8) |
7, 8 (7-8) |
|
| Total vertebrae Abdominal Caudal (including urals) |
34 (33-35) 16 (15-17) 19 (17-19) |
31 (31-32) 17 (16-18) 15 (13-15) |
38 (36-38) 18 (17-19) 20 (17-21) |
36 or 37 (35-37) 16 (16-18) 20 (19-21) |
38 (37-39) 24? (23-25) 14, 15 (14-15) |
34 (32-37) ¿ (14-19) 18 (13?-18) |
|
| Predorsal scutes | 8 (8) |
9 (9) |
17 (17-19) |
11? (7+-11?) |
? (5?-9?) |
11 (10-11) |
|
| Prepelvic scutes | 17 (16-17) |
16 - |
20 (19-21) |
17 - |
13 (10-13) |
13 (13) |
|
| Postpelvic scutes | (6) |
7 (6?-7) |
8 (8-9) |
7 (7) |
10, 11 (8-11) |
9-10 (9-10) |
|
| Other features |
Skull roof bones | Stout ridges ornament the frontal, parietal, and pterotic. | Smooth | ||||
| Parasphenoid | with the osteoglossid-like tooth patch | ? | toothless | ||||
| Opercle ornaments | Basal short and straight grooves | ? | Smooth | ||||
| Epineurals and epipleurals | close to caudal peduncle | far from the caudal peduncle | close to caudal peduncle | far from the caudal peduncle | |||
| Predorsal bones below predorsal scutes | Posterior 4-3 (= predorsal series inccomplete | All (= predorsal series complete) | Posterior 4 | ? | All (= predorsal series complete) | ||
| Posterior predorsal scutes |
Posterior ones subtriangular, others are ovoid, all with spine | Posterior ones subtriangular with a small spine, others ahead become heart-shaped | All are heart-shaped | All are wide and rectangular
With no posterior spine |
All are heart-shaped | All subrectangular with anterior and posterior spines | |
