Notropis lucifer, Pinion & Kim & Dolan & Portnoy & Voelker & Conway, 2025

Pinion, Amanda K., Kim, Daemin, Dolan, Elizabeth P., Portnoy, David S., Voelker, Gary & Conway, Kevin W., 2025, ? Revision of Notropis stramineus (Cope, 1865), descriptions of three new species and comments on the monophyly of Miniellus Jordan, 1882 (Pisces: Cypriniformes: Leuciscidae), Vertebrate Zoology 75, pp. 699-755 : 699-755

? Notropis lucifer sp. nov.

Figures 10 D , 13 , 14 , 15 , 16E, F , 30 , 31 , 32

Holotype.

TCWC 15689.08 , male, 43.2 mm SL; United States of America, Texas: Val Verde County: Devils River, Devils River Ranch , 29°40'34.32"N, 101°0'5.7594"W; Conway, K. W. and Prestridge, H. L., 18 April, 2012 (Fig. 30 A ) .

Paratypes.

TCWC 15551.09 , 10, 22–40 mm SL; Llano: Sandy Creek @ SH 16 , 30°33'11"N 098°42'05"W; 19 September, 2018 . – TCWC 922.02 , 16, 38–51 mm SL; Llano: Sandy Creek, 19.0 mi SW of Llano ., 30°32'04"N 098°47'52"W; 24 April, 1976 . – TCWC 2069.02 , 12, 23–43 mm SL; Llano: Sandy Creek at Hwy 71 , 30°33'00"N 098°28'00"W; 17 September, 1977 . – JFBM 18636 , 4, 32–43 mm SL; Gillespie: Llano River, at Lang’s Mill (station 32) , 30°28'50"N 099°07'15"W; 5 June, 1956 . – TCWC 4065.01 , 110, 25–37 mm SL; Gillespie: Pedernales River, Trough Springs on headwaters of Wolf Creek ., 30°08'36"N 099°04'42"W; 30 October, 1971 . – TNHC 47383 , 34, 20–49 mm SL; Kimble: Little Devils River at CR 4301 , 30°28'55"N 099°23'07"W; 9 July, 2013 . – JFBM 18522 , 93, 16–40 mm SL; Gillespie: Wolf Creek at Springs, 1 mile Below Texas Highway 16, 4 miles S of junction with Pedernales River , 30°10'28"N 099°00'41"W; 22 November, 1954 . – TCWC 15689.04 , 26, 23–43 mm SL; Val Verde: Devils River , 29°39'51"N 100°57'10"W; 19 September, 2018. – TCWC 7510.06 , 106, 15–37 mm SL; Val Verde: Devils River , 29°53'08"N 100°59'36"W; 8 March, 1993 . – TNHC 30952 , 175, 16–49 mm SL; Val Verde: Devils River , 29°56'21"N 101°05'46"W; 27 July, 2004 . – TCWC 11924.01 , 2, 26–40 mm SL; Val Verde: Devils River , 29°52'23"N 100°59'35"W; October, 1997 . – TNHC 30562 , 559, 15–55 mm SL; Val Verde: Devils River , 29°52'23"N 100°59'35"W; 29 July, 2003 . – TCWC 7509.09 , 14, 23–42 mm SL; Val Verde: Devil’s River; 175 m upstream from Dolan Falls , 29°53'08"N 100°59'36"W; 17 May, 1994 . – JFBM 18795 , 17, 37–42 mm SL; Val Verde: Devil’s Lake, Devil’s River, 8 air miles NW of Del Rio , 29°29'40"N 100°59'19"W; 27 March, 1954 . – TNHC 29785 , 435, 14–49 mm SL; Val Verde: Devils River , 29°52'23"N 100°59'35"W; 30 July, 2002 . – UMMZ 211231 , 23, 24–47 mm SL; Brewster: Terlingua Creek, at Hwy 170, 1.3 mi W of Study Butte (JCT 170 and 118, 29°19'38"N 103°33'12"W; 14 March, 1982 .

Diagnosis.

Notropis lucifer sp. nov. is distinguished from all members of the Notropis stramineus species group, except N. chihuahua , by the following characters: absence of cross-hatched (outlined) scales located on the 2–3 scale rows dorsal to the lateral-line canal scale row (vs. present and well developed anteriorly, and present but sometimes poorly developed posteriorly), cleithral streak absent (Devils and Colorado River drainages) or if present greatly reduced ( Colorado River drainage, most obvious in males) (vs. present and well developed, most obvious in males), sparse, large melanophores in the rostral region of the head (vs. many small, diffuse melanophores), few, large melanophores laterally in the fronto-occipital region (vs. many small melanophores scattered across cranial region 8), restriction of few, large melanophores to the anterior half of the internarial region (vs. internarial region completely covered in small, diffuse melanophores), a more pronounced “ wedge ” of melanophores at insertion of dorsal procurrent rays, and extent of ossification of the ethmoid region reduced, with much of the ethmoid region remaining cartilaginous (vs. well-ossified), and a broad (vs. thin and tapering) vomer. Notropis lucifer sp. nov. is distinguished from N. chihuahua by smaller, fewer, and less diffuse macromelanophores generally, except in the internarial region, where melanophores are abundant (vs. sparse in N. chihuahua ). Notropis lucifer sp. nov. is further distinguished from N. chihuahua by a continuous edge of medium to small melanophores outlining scales on nape and dorsal body rows (vs. scales on nape and scale rows dorsal to lateral-line scale row irregularly outlined by macromelanophores). Notropis lucifer sp. nov. is further distinguished from N. topeka and N. procne by the lack of a pronounced lateral stripe, and further distinguished from N. topeka by tubercle size and distribution, being small and distributed across most dorsal and lateral regions of the head (vs. very large and absent from the infraorbital, preopercular, opercular, and subopercular regions). Notropis lucifer sp. nov. is further distinguished from N. stramineus , N. missuriensis , N. multicorniculatus sp. nov. and N. oblitus sp. nov. by the absence (vs. presence) of a bluish sheen along the lateral body side in life, and cephalic pigmentation concentrated in the lacrimal, suborbital, dorsalmost part of opercular, fronto-occipital, and interorbital regions into clusters of melanophores (vs. cephalic pigmentation present in similarly-sized melanophores scattered evenly across most regions of the head, except ventrally), and rostral cap translucent and devoid of pigment with large clusters of melanophores on the lateral surface of the head, particularly in the lacrimal region (vs. rostral cap and lacrimal region covered in evenly distributed field of small melanophores). It is further distinguished from N. missuriensis and N. multicorniculatus sp. nov. by a larger eye (orbit diameter ca. 30 % of HL vs. 25–27 % of HL), and further from N. multicorniculatus sp. nov. by having physically larger pre-dorsal scales (vs. smaller scales, appearing crowded in pre-dorsal scale rows). Notropis lucifer sp. nov. is further distinguished from N. oblitus sp. nov. by the combination of an absent or weakly developed cleithral streak (vs. well-developed), and the absence of a cross-hatched pattern of melanophores in the two to three scale rows dorsal to lateral-line scale row along the length of the body (vs. presence of the cross-hatched pattern, sometimes weakly developed posteriorly).

Description.

Body shape and general appearance as in Figures 30 – 32 . Morphometric and meristic data are listed in Tables 7 . As described for N. stramineus , except for the following. Head deep (ca. 54 % HL), more so in Colorado River drainage. Dorsal profile low, arch from snout to dorsal-fin origin weak. Snout rounded. Tuberculation as described for Notropis oblitus sp. nov. Ethmoid region weakly ossified; vomer broad (Fig. 10 D ).

Coloration.

Coloration in preservative (Figs 30 , 31 ) as in Notropis oblitus sp. nov. except for the following. Melanophores below lateral-line scale row almost entirely absent anteriorly. Head pigmentation in clusters of melanophores, some forming macromelanophores, restricted to lacrimal, suborbital, dorsalmost part of opercular, fronto-occipital, and interorbital regions. Pigmentation reduced in supraorbital region and restricted to anteriormost and posteriormost half of internarial and rostral regions, respectively (Fig. 14 and 13 E, F ).

Coloration in life silvery, pale, much of body side without yellow pigment, bluish sheen absent (Fig. 32 ). Chromatophores on dorsal midline creating thick golden streak, continuing onto head and covering much of dorsal surface (Fig. 32 D ). Scale rows in dorsal midline posterior to dorsal fin with thick margin of melanophores on posterior margin of scale. Cleithral streak absent or weakly developed and short. Pectoral fins largely hyaline; in males, pigment on pectoral fins restricted to few scattered melanophores on leading edge.

Distribution.

Notropis lucifer sp. nov. is distributed in the upper portions of the Colorado River basin of Texas, and in the Rio Grande basin (Fig. 29 ), where it is distributed in the Devils River (Fig. S 17). Notropis lucifer sp. nov. is presumably also present in Pinto Creek, a tributary of the Rio Grande south of the Devils River, though recent attempts to collect the species there have failed. In Mexico, Notropis lucifer sp. nov. is presumed to be distributed in tributaries to the Rio Grande, including the Rio Salado and Rio San Juan, though genetic material is not available for these populations, making assignment of these populations to this species tentative. The report of N. stramineus sensu lato from the Río Pilón near Montemorelos by Miller et al. (2005: fig. 6.123, image of specimen from UMMZ 210715 , not included in distribution map) appears to be based on a misidentification. Collections from the lower Pecos River, including Independence Creek (not evaluated herein), may also belong to this species and deserve further study.

Etymology.

Lucifer in Latin means light bearer or morning star. This species is so named for the comparatively bright appearance to that of the remaining members of the Notropis stramineus species group. This name also alludes to the Devils River, the type locality of this species, in reference to the connotation of Lucifer as Satan, the Devil, in Jewish and Christian literature. Additionally, the authors hope the recognition of this new species will cast a light upon the conservation risks of this geographic region, experienced by much of the arid regions of southwestern U. S. and adjacent Mexico. A noun in apposition.

Remarks.

Populations of Notropis lucifer sp. nov. from the Colorado River basin of Texas exhibit some morphological characteristics that are intermediate between those seen in the Devils River populations and those of individuals of N. oblitus sp. nov. (see, e. g., Figs 13 – 16 E, F vs. 13–16 C). These include a deeper body, the occasional presence of a subtle cleithral streak (most pronounced in males), and increased pigmentation posteriorly dorsal to the lateral line scale row (though typically not forming complete cross-hatching pattern). These intermediate conditions could be the result of past hybridization events between N. lucifer sp. nov. in the Colorado River basin and N. oblitus sp. nov., a scenario that likely explains the close relationship and low genetic distance in analyses of mitochondrial DNA.

Figures

Figure 10. Cleared and stained ethmoid region of select members of Notropis stramineus species group. a, N. stramineus sensu stricto (Clade 1), TCWC 21295.01, Michigan, Huron River; b, N. multicorniculatus sp. nov. (Clade 2), TCWC 15789.07; c, N. missuriensis (Clade 4), TCWC 21299.01, Missouri, Contrary Creek, Missouri River drainage; d, N. lucifer sp. nov. (Clade 5, 6), TCWC 15689.04, 40.0 mm SL. V, vomer; P, parasphenoid; LE, lateral ethmoid. White outline demarcates border of vomer; note size and extend of cartilage in D (N. lucifer sp. nov.) relative to the other three.

Figure 13. Lateral view of heads of males of each member of Notropis stramineus species group; A Notropis stramineus sensu stricto (Clade 1), ROM 11967, 46.6 mm SL; B N. multicorniculatus sp. nov. (Clade 2), TCWC 15789.07, 43.2 mm SL; C N. oblitus sp. nov. (Clade 3), TCWC 16802.06, 41.2 mm SL; D N. missuriensis (Clade 4), KU 4186, 48.4 mm SL; E N. lucifer sp. nov., Devils River (Clade 5), TCWC 15689.04, 43.2 mm SL; F N. lucifer sp. nov., Col. drainage (Clade 6), TCWC 19721.01, 42.3 mm SL; G Notropis topeka, KU 40725, 46.1 mm SL; H Notropis procne, NCSM 12688, 51.1 mm SL; I Notropis chihuahua, UMMZ 211231, 41.8 mm SL. Scale bars = 2 mm.

Figure 14. Dorsal view of heads of males of each member of Notropis stramineus species group; A Notropis stramineus sensu stricto (Clade 1), ROM 11967, 46.6 mm SL; B N. multicorniculatus sp. nov. (Clade 2), TCWC 15789.07, 43.2 mm SL; C N. oblitus sp. nov. (Clade 3), TCWC 16802.06, 41.2 mm SL; D N. missuriensis (Clade 4), KU 4186, 48.4 mm SL; E N. lucifer sp. nov., Devils River (Clade 5), TCWC 15689.04, 43.2 mm SL; F N. lucifer sp. nov., Col. drainage (Clade 6), TCWC 19721.01, 42.3 mm SL; G Notropis topeka, KU 40725, 46.1 mm SL; H Notropis procne, NCSM 12688, 51.1 mm SL; I Notropis chihuahua, UMMZ 211231, 41.8 mm SL. Scale bars = 2 mm.

Figure 15. Ventral view of heads of males of each member of Notropis stramineus species group; A Notropis stramineus sensu stricto (Clade 1), ROM 11967, 46.6 mm SL; B N. multicorniculatus sp. nov. (Clade 2), TCWC 15789.07, 43.2 mm SL; C N. oblitus sp. nov. (Clade 3), TCWC 16802.06, 41.2 mm SL; D N. missuriensis (Clade 4), KU 4186, 48.4 mm SL; E N. lucifer sp. nov., Devils River (Clade 5), TCWC 15689.04, 43.2 mm SL; F N. lucifer, Col. drainage (Clade 6), TCWC 19721.01, 42.3 mm SL; G Notropis topeka, KU 40725, 46.1 mm SL; H Notropis procne, NCSM 12688, 51.1 mm SL; I Notropis chihuahua, UMMZ 211231, 41.8 mm SL. Scale bars = 2 mm.

Figure 16. Caudal peduncles of males of each member of Notropis stramineus species group; A Notropis stramineus sensu stricto (Clade 1), ROM 11967, 46.6 mm SL; B N. multicorniculatus sp. nov. (Clade 2), TCWC 15789.07, 43.2 mm SL; C N. oblitus sp. nov. (Clade 3), TCWC 16802.06, 41.2 mm SL; D N. missuriensis (Clade 4), KU 4186, 48.4 mm SL; E N. lucifer sp. nov., Devils River (Clade 5), TCWC 15689.04, 43.2 mm SL; F N. lucifer sp. nov., Col. drainage (Clade 6), TCWC 19721.01, 42.3 mm SL; G Notropis topeka, KU 40725, 46.1 mm SL; H Notropis procne, NCSM 12688, 51.1 mm SL; I Notropis chihuahua, UMMZ 211231, 41.8 mm SL. Scale bars = 2 mm.

Figure 29. Map of southern Texas and Northern Mexico, depicting distribution of samples of Notropis oblitus sp. nov. and N. lucifer sp. nov. used in 3 - gene dataset. Type localities of each new species indicated by a star.

Figure 30. Notropis lucifer sp. nov., Texas, Val Verde County, Devils River; A TCWC 15689.08 holotype, male, 43.2 mm SL; B TCWC 15689.04, paratype, female, 38.6 mm SL. Scale bars = 5 mm.

Figure 31. TCWC 19721.01, Notropis lucifer sp. nov., paratypes, Texas, Kimble County, Little Devils River; A male, 42.3 mm SL; B female, 42.2 mm SL. Scale bars = 5 mm.

Figure 32. Notropis lucifer sp. nov. photographed at Devils River Ranch, Devils River, Texas, April 2022 (A, B, D), and at the San Saba River (C); A male, B female, C male, D, sex undetermined, demonstrating dorsal midline pigmentation. Scale bars = 5 mm.