TABLE 1. Meristic data of the type specimens of Paleocharacodon guzmanae gen. and sp. nov. described here. All measurements are in millimeters. The preural centrum 1 and ural 1 (fused to form the caudal complex) are considered into the vertebrae account. Abbreviatures: caudal peduncle height; MBH, maximum body height.
| Males | Females | ? | ||||||||||||||
| IGM 13118 |
IGM 13119 |
IGM 13120 |
IGM 13121 |
IGM 13122 |
IGM 13123 |
Mean or mode, and (range) |
IGM 13117 |
IGM 13124 |
IGM 13125 |
Mean, mode, and (range) |
IGM 13126 |
IGM 13127 |
IGM 13128 |
IGM 13129 |
||
| Total length | 24.42 | 39.84 | 44.75 | 56.88? | - | - | (24.42-56.88?) | 48.27+ | 41.78+ | 29.4? | (41.78-48.27+) | - | - | - | - | |
| Standard length | 21.88 | 34.59 | 40.32 | 55.86? | - | - | (21.88-40.32+) | 37.29 | 36.55 | 25.66 | (25.38-37.29) | - | - | - | 45? | |
| Head length | 5.90 | 9.69 | 10.75 | 16.14? | - | - | 10.73 | 11.88 | 9.27 | - | - | 15.43? | - | |||
| as % of SL | 26.96 | 28.01 | 26.66 | - | - | - | 27.21 (26.66-28.01) |
28.77 | 32.50 | 36.12 | 32.46 (28.77-36.12) |
- | - | - | - | |
| Head height | 5.96 | 8.89 | 8.74 | - | - | - | 10.90 | 13.34 | 7.72 | - | - | 10.73? | - | |||
| as % of SL | 27.23 | 25.7 | 21.67 | - | - | - | 24.86 (21.67-27.23) |
29.23 | 27.39 | 30.08 | 28.9 (27.39-30.08) |
- | - | - | - | |
| Predorsal length | - | 24.02 | 27.96 | 33.14? | - | - | 26.32 | 26.37 | 18.71 | - | - | - | - | |||
| as % of SL | - | 69.44 | 69.34 | - | - | - | 69.39 (69.34-69.44) |
70.58 | 72.14 | 72.91 | 71.87 (70.58-72.91) |
- | - | - | - | |
| Prepelvic length | - | 19.94 | 22.97 | 23.74? | - | - | 23.45 | 26.50 | 15.16 | - | - | - | - | |||
| as % of SL | - | 57.64 | 56.96 | - | - | - | 57.3 (56.96-57.64) |
62.88 | 72.50 | 59.08 | 64.82 (59.08-72.50) |
- | - | - | - | |
| Preanal length | 15.68 | 24.03 | 28.45 | 33.09? | - | - | 29.21 | 26.42 | 17.75 | - | - | - | - | |||
| as % of SL | 71.66 | 69.47 | 70.56 | - | - | - | 70.56 (69.47-71.66) |
78.33 | 72.28 | 69.17 | 73.26 (69.17-78.33) |
- | - | - | - | |
| Dorsal fin length | - | 5.36 | 4.87 | 6.08? | 4.72+ | - | 4.30 | 3.77 | 3.10 | 6.0 | - | 6.78 | 5.37 | |||
| as % of SL | - | 15.49 | 12.07 | - | - | - | 13.78 (12.07-15.49) |
11.53 | 10.31 | 12.08 | 11.3 (10.31-12.08) |
- | - | - | - | |
| Anal fin length | 3.24 | 5.87 | 4.14? | 5.66? | 6.89 | 4.12 | 3.34 | 3.54 | 3.11 | 4.95? | 3.78 | 5.01 | 3.83? | |||
| as % of SL | 14.80 | 16.97 | - | - | - | - | 15.88 (14.80-16.97) |
8.95 | 9.68 | 12.12 | 10.25 (8.95-12.12) |
- | - | - | - | |
| MBH | 4.07 | 5.76 | 9.19 | 15.69? | 22.71 | - | 13.30 | 10.58 | 7.50 | - | 9.33 | - | - | |||
| as % of SL | 18.60 | 16.65 | 22.79 | - | - | - | 19.34 (16.65-22.79) |
35.66 | 28.94 | 29.22 | 31.27 (28.94-35.66) |
- | - | - | - | |
| CPH | 2.41 | 4.11 | 4.50 | 8.88? | 10.01 | - | 6.30 | 6.18 | 5.23 | 6.79 | - | - | 5.20 | |||
| as % of SL | 11.01 | 11.88 | 11.16 | - | - | - | 11.35 (11.01-11.88) |
16.89 | 16.90 | 20.38 | 18.05 (16.89-20.38) |
- | - | - | - | |
| Total vertebrae | 35 | 35 | 34 | - | - | - | 34 (34-35) |
31 | 31 | 32 | 31 (31-31) |
- | - | - | - | |
| Abdominal | 15 | 15 | 15 | 10+ | 11+ | ? | 15 (15) |
13 | 14 | 14 | 14 (13-14) |
2+ | 7+ | ? | ? | |
| Caudal | 20 | 20 | 19 | 19 | 20 | 18 | 20 (18-20) |
18 | 17 | 18 | 18 (17-18) |
17 | 11+ | 14+ | 15+ | |
| Ribs (pairs) | 15 | 15 | 14 | 11? | 9+ | 4+ | 15 (14-15) |
12 | 13 | 12 | 12 (12-13) |
? | 11? | 12? | 12 | |
| Dorsal fin rays | - | 15 | 15 | 14 | 8+ | ? | 15 (14-15) |
12 | 12 | 11 | 12 (11-12) |
11 | ? | ? | 14? | |
| Anal fin rays | 16 | 18 | 16 | 17 | 16 | 15+ | 16 (16-18) |
16 | 15 | 16 | 16 (15-16) |
11+ | ? | 6+ | ? | |
| Andropodium | 6 | 6 | ? | - | 6? | 5 | 6 (5-6) |
- | - | - | - | - | - | - | - | |
| Pectoral rays | - | 6+ | 12 | 4+ | - | 11? | 12 | 13 | 14 | 12? | 13-14 | ? | ? | ? | 4+ | |
| Pelvic rays | - | 4+ | 6 | ? | 4+ | 6 | 6 | 6 | 6 | 6 | 6 (6) | 4? | 3+ | 6? | 6 | |
TABLE 2. Otolith measurements of Paleocharacodon guzmanae gen. and sp. nov., from the Paleolake Amajac deposits, Sanctórum, Mexico, based on the left otolith recovered in IGM 13119 (Figure 9). All the lengths and heights are in micrometers.
| measurement | as % of total length | |
| Total length (ML+RL) | 1237.1 | - |
| Medial length (ML) | 994.4 | 80.3% |
| Dorsal length (DL) | 907.3 | 77.3% |
| Ventral length (VL) | 920.4 | 74.4% |
| Rostrum length (RL) | 242.7 | 19.6% |
| Antirostrum length (AL) | 92.0 | 74.3% |
| Maximum height (MH) | 907.3 | 73.3 |
| Rostrum height (RH) | 364.1 | 29.4% |
| Antirostrum height (AH) | 349.7 | 28.2% |
| Exisura angle (ea) | 100° | - |
| Posterior angle (pa) | 72° | - |
| Posteroventral angle (pva) | 128° | - |
TABLE 3. Valid nominal species of the family Goodeidae (the listed numbers are the same present in Figure 16).
| Species | Status and distribition |
| 1. Allodontichthys hubbsi Miller and Uyeno, 1980. | Endangered. Upper Coahuyana River Basin. Jalisco. |
| 2. Allodonthichthys polylepis Rauchenberger, 1988. | Vulnerable. Upper Ameca River Basin. Jalisco |
| 3. Allodontichthys tamazulae Turner, 1946. | Vulnerable. Coahuayana River Basin. Jalisco. |
| 4. Allodontichthys zonistius Hubbs, 1932. | Vulnerable. Armeria River Basin. Colima and Jalisco. |
| 5. Alloophorus robustus Bean, 1892. | Vulnerable. Cotija, Cuitzeo, Pátzcuaro, Zacapu, Zirahuén, Santiago, and Balsas basins. Michoacán and Jalisco. |
| 6. Allotoca catarinae De Buen, 1942. | Critically endangered. Upper Cupatizio River Basin. Michoacán. |
| 7. Allotoca diazi Meek, 1902. | Critically endangered. Lake Pátzcuaro Basin. Michoacán. |
| 8. Allotoca dugesii Bean, 1887. | Endangered. Middle and lower Rio Lerma Basin and upper Rio Santiago Basin. Michoacán, Guanajuato, and Jalisco. |
| 9. Allotoca goslinei Smith and Miller, 1987. | Extinct in wild. Arroyo Potrero Grande in Ameca River basin. Jalisco. |
| 10. Allotoca maculata Smith and Miller, 1980. | Critically endangered. Santa Magdalena Lake Basin. Jalisco. |
| 11. Allotoca meeki Álvarez, 1959. | Critically endangered. Lake Zirahuén Basin. Michoacán. |
| 12. Allotoca zacapuensis Meyer, Radda, and Domínguez, 2001. | Critically endangered. Lake Zacapu Basin. Michoacán. |
| 13. Ameca splendens Miller and Fitzsimons, 1971. | Critically endangered. Upper Ameca River basin, Teuchitlán springs. Jalisco. |
| 14. Ataenobius toweri Meek, 1904. | Endangered. Pánuco Basin, Verde River and La Media Luna Lagoon. San Luis Potosi. |
| 15. Chapalichthys encaustus Jordan and Synder, 1899. | Vulnerable. Chapala Lake and adjacent portions of Lower Lerma basins. Jalisco and Michoacán. |
| 16. Chapalichthys pardalis Álvarez, 1963. | Critically endangered. Springs near Tocumbo, San Juanico, Michoacán. |
| 17. Chapalichthys paraticus Álvarez, 1963. | Endangered. San Juanico reservoir. Cotija, Michoacán. |
| 18. Characodon audax Smith and Miller, 1986. | Endangered. Ojo de agua de las Mujeres. Durango. |
| 19. Characodon garmani Jordan and Evermann, 1898. | Extinct. Parras Valley, Nazas-Aguanaval River basin, Mayrán Lagoon. Coahuila. |
| 20. Characodon lateralis Günter, 1866. | Critically endangered. Mezquital River Basin. Durango. |
| 21. Crenichthys baileyi (Gilbert, 1893). | Endangered. White River system, springs of Moapa River. Southeastern Nevada. |
| 22. Crenichthys nevadae Hubbs, 1932. | Vulnerated. Railroad Valley. Nye County, Nevada. |
| 23. † Empetrichthys erdisi Uyeno and Miller, 1962. | Fossil (Pliocene). Santa Clara River Valley, in the Piru Mountains, Ventura County, California. |
| 24. Empetrichthys latos Miller, 1948. | Critically Endangered (only introduced). Pahrump Valley, Nye County, Nevada. |
| 25. Empetrichthys merriami Gilbert, 1893. | Extinct. Ash Meadows, Amargosa Desert, Nye County, Nevada. |
| 26. Girardinichthys multiradiatus Meek, 1904. | Endangered. Basins of Balsas, upper Lerma, and Tuxpan rivers; Zempoala lake. Estado de México, Michoacán, and Morelos. |
| 27. Girardinichthys viviparus, Bustamante, 1837. | Endangered. Valley of Mexico Basin, Río Tula. Estado de México, Mexico City, and Hidalgo. |
| 28. Goodea atripinnis Jordan, 1880. | Least Concern. Widely distributed in basins of Lerma, Santiago, Balsas, Armeria, Ameca and River, as well as Tierra Quemada, Zirahuén, Zacapu, Chapala, and Pátzcuaro lakes. Hidalgo, Jalisco, San Luís Potosí, and Michoacán. Recently, it was discovered in the Río Metztitlán, Pánuco Basin. Hidalgo (Miranda et al., 2010). |
| 29. Hubbsina turneri De Buen, 1940. | Critically endangered. Cuitzeo basin, Yuriria reservoir and Zacapu Lake in Lerma River Basin. Guanajuato and Michoacán. |
| 30. Ilyodon cortesae Paulo-Maya and Trujillo-Jiménez, 2000. | Threatened. San Juan Tacámbaro River, Michoacán. |
| 31. Ilyodon furcidens Jordan and Gilbert, 1882. | Least Concern. Marabasco, Armeria, and Ameca River basins. Colima and Jalisco. |
| 32. Ilyodon lennoni Meyer and Förster,1983. | Vulnerable. Chacambaro stream, Lerma River Basin. Guerrero. |
| 33. Ilyodon whitei Meek, 1904. | Vulnerable. Balsas River basin. Jalisco, Michoacán, Morelos, Puebla, and Guerrero. |
| 34. Neoophorus regalis Álvarez, 1959. | Critically endangered. Balsas River basin in the Los Reyes. Michoacán. |
| 35. † Paleocharacodon guzmanae. Here. | Fossil (Pliocene). Sanctórum site. Hidalgo. |
| 36. Skiffia bilineata Bean, 1887. | Endangered. Cuitzeo Lake and Lerma River basins. Michoacán and Guanajuato. |
| 37. Skiffia francesae Kingston, 1978. | Extinct in wild. Teuchtitlán, Ameca River basin. Jalisco |
| 38. Skiffia lermae Meek, 1902. | Endangered. Pátzcuaro, Zirahuén, Cuitzeo, Zacapu lake basins; Lerma River. Michoacán and Guanajuato. |
| 39. Skiffia multipunctata Pellegrin, 1901. | Endangered. Lower Lerma River Basin, Lake Chapala, and the upper part of the Santiago River basin. Jalisco and Michoacán. |
| 40. † Tapatia occidentalis Álvarez and Arreola-Longoria, 1972. | Fossil (Pliocene). Amatitán. Jalisco. |
| 41. Xenoophorus captivus Hubbs, 1924. | Endangered. Upper Pánuco River basin, springs in the Venados, Moctezuma, and Agua de Enmedio; Santa María del Río and Tierra Quemada rivers; Jesús María reservoir. San Luis Potosi. |
| 42. Xenotaenia resolanae Turner, 1946. | Vulnerable. Marabasco and Purificación River basins. Pacific slope of Jalisco. |
| 43. Xenotoca doadrioi Domínguez-Domínguez, Bernal-Zuñiga, and Piller, 2016. | Critically Endangered. Hacienda San Sebastián pond, El Moloya spring, Estancia de Ayoles reservoir, Oconahua Dam. Etzatlán region, in Jalisco. |
| 44. Xenotoca eiseni Rutter, 1896. | Endangered. Santiago River basin and Compostela River, Magdalena Lake basin, and upper parts of Ameca and Coahuayana River basins. Nayarit and Jalisco. |
| 45. Xenotoca lyonsi Domínguez-Domínguez, Bernal-Zuñiga, and Piller, 2016. | Critically Endangered. Middle and upper part of the Coahuayana River Basin; Tuxpan and Tamazula rivers. Jalisco. |
| 46. Xenotoca melanosoma Fitzsimons, 1972. | Endangered. Magdalena Lake and the Ameca, Armería, Santiago, and Coahuayana River basins. Jalisco. |
| 47. Xenotoca variata Bean, 1887. | Least concern. Pánuco River Basin, middle Lerma River basin, and upper Santiago River basin; and Cuitzeo and Zacapu Lake basins. Aguascalientes, Jalisco, Guanajuato, Querétaro, and Michoacán. |
| 48. Zoogoneticus purhepechus Domínguez-Domínguez, Pérez-Rodríguez, and Doadrio, 2008. | Vulnerable. Lower Lerma River, including Duero, Grande, and Santiago rivers; Upper Ameca River; Chapala Lake; and Magdalena, Sayula, San Marcos, Zacoalco, and Atotonilco lagoons. Michoacán and Jalisco. |
| 49. Zoogoneticus quitzeoensis Bean, 1898. | Endangered. Middle and lower Lerma River Basin, upper part of basins of Santiago, Armeria, Ameca rivers, Chapala Lake. Michoacán and Jalisco. |
| 50. Zoogoneticus tequila Webb and Miller, 1998. | Endangered. Teuchitlán springs, upper Ameca River Basin, Jalisco. |
TABLE 4. Comparison of Paleocharacodon guzmanae gen. and sp. nov. and other goodeids (characters based on Parenti, 1981; Webb, 1998; data of supraorbital canal also based on Fitzsimons (1981); see the main text of Discussion). Isolated numbers between parentheses correspond to those characters shown in Figure 15. Abbreviations between parentheses: +, a character present in the taxa noted ahead;-, a character absent in the taxa noted ahead; v, a character variable in the taxa noted ahead).
|
|
Empetrichthyinae | Goodeinae | ||||||
| † Tapatia occidentalis |
† P. guzmanae | Goodeiini | Ilyodontini | Girardichthyini | Chapalichthyini | Characodontini | ||
| Viviparity | No | ? | Yes | |||||
| Anal fin ray 1 | Normal | Rudiementary (8) | ||||||
| Andropodium | Absent | Present (9) | ||||||
| Pelvic fin | Absent (10) | Present | ||||||
| Epibranchial | Y-shaped (11) (v, Crenichthys) |
? | Unforked (With no accessory strut near the base) |
|||||
| Surface of neurocranial bones | Rugouse (12) | Smooth | ||||||
| Anterior edge of the vomer | Abruptly concave (13) | ? | Slightly concave to Straight | |||||
| Supraorbital canal ossification | Partially ossified (14) | With out any ossification. | ||||||
| Pores of supraorbital canal | With no break 4a-4b (15) | ? | With the break 4a-4b (v, Allodontichthys zonistius, Allotoca diazi, A.meeki, Characodon lateralis, Chapalichthys encaustus, Ilyodon whitei, Xenoophorus captivus, Xenotoca variata) (-, Allotoca dugesii, A. maculata, Allodontichthys except for A. zonistius, Allotoca catarinae) (Pit organs only in Girardinichthys, Hubbsina, and Skiffia) |
|||||
| Pharyngobranchials | Higher than their teeth (16) | ? | Teeth higher than pharyngobranchials (-, Xenotoca melanosoma) | |||||
| Third postcleithrum | Present (17) | Absent | ||||||
| Origin of dorsal fin | Posterior, it rises at or behind 67% of SL | Anterior, it rises anterior to 67% of SL (18) | Posterior, it rises at or behind 67% of SL | Anterior, it rises anterior 67% of SL (18) (+, Skiffia, Zoogoneticus) (-, Xenotaenia, Allotoca diazi) (v, Allotoca dugesii, A. goslinei) | Posterior, it rises at or behind 67% of SL | |||
| Neurocranium height in its posterior view | Low, the heigh/width ratio is 0.45 to 0.52 | ? | Low | High, the heigh/width ratio is 0.58 to 0.66 (19) (-,Alloophorus) (+, Skiffia, Zoogoneticus) |
||||
| Presence of rostral cartilages | Yes (20) | ? | Yes (20) | No | ||||
| Anterior parasphenoid end. | No inged | ? | No winged | Winged (21) | No winged | |||
| Length of Pelvic fin | Long | Short (22) | Long | Short (22) | ||||
| Epioccipital posterior process | Short | ? | Long (23) (+, Skiffia) |
Long (23) | ||||
| Gill rackers in the branchial arch 1 | 9-19 in Empetrichthys, 24-59 (24) in Crenichthys | ? | 24-59 (24) (-, Allodontichthys polylepis, A. tamazulae, A. zonistius) (+, Skiffia) |
9-19 (-, Ameca, Xenoophorus) |
||||
| Shape of principal teeth in jaws | Unicuspid in Empetrichthys. ricuspid in Crenichthys | Bicuspid and straight (25) | Bicuspid and sharp | Bicuspid and straight (25) | Bicuspid and sharp (-, unicuspid in Alloophorus, Allotoca, Girardinichthys viviparus, Zoogoneticus) (-, tricuspid in Allodontichthys) (v, uni and bicuspid in Xenoophorus) | |||
| Attachement of principal teeth bases in jaws | Co-ossified | Loose (26) | Co-ossified | Loose (26) | Co-ossified (26) (-, loose in Ilyodon, Skiffia francesae, S. lermae) (v, S. multidenatata, Xenoophorus) |
|||
| Anguloarticular angle | ≈ 90° | ≈ 180° (27) | ≈ 90° (+, Zoogoneticus) (-, ≈180° in Skiffia and Ilyodon) |
|||||
| Posteroventral margin of the coracoid | With a straight angle | Convex, vertical, and notched (28) | With a straight angle (-, convex, vertical, and notched in Ameca and Xenotoca variata (28) (-,broad convex in Skiffia) (v, straight and broad convex in Ilyodon) |
|||||
| Palatoquadrate arch | Inclined slightly forward | Vertical 29) (+, Skiffia) |
Inclined slightly forward (+, Zoogoneticus) |
|||||
| Retroarticular in the facet for the quadrate | Excluded | ? | Included (30). (It is in the center in P. guzmanae. It is lingually in Goodiini, Ameca, Ilyodon, and Skiffia) |
Excluded (-, Ameca, Skiffia, Ilyodon) | Included (30). It is in the center | |||
| Facet for the quadrate Donut-like (31) | No | ? | Yes | No, (+, Skiffia, Zoogoneticus) | Yes | |||
| Anterolateral projection in the palatine | Present | ? | Present | Absent (32) (+, Skiffia francesae, Zoogoneticus) |
Present (-, Allophorus) |
Absent (32) | ||
| Ventral process of postemporal | Absent (33) | ? | Very small | Large (+, Skiffia) | Absent (33) (+, Zoogoneticus) |
Large | Absent (33) | |
