Etheostoma kimberlae Mayden, 2025

Mayden, Richard L. & Kuhajda, Bernard R., 2025, Two new species of Etheostoma from the Black Warrior River System (Mobile Basin) of Alabama (Teleostei: Percidae), Zootaxa 5618 (3), pp. 351-371 : 352-361

Etheostoma kimberlae Mayden , new species

Locust Fork Darter

Fig. 1

Holotype. UAIC 11031.01 , adult male, 39.6 mm standard length. Locality. Alabama, Blount Co., Mill Creek at gravel road 4.3 km SW Oneonta, T13 S, R1 E, S11, 33.922595 -86.509051, 22 March 1994. Collected by B. R. Kuhajda and R. L. Mayden.

Paratypes. UAIC 11031.03 , same data as holotype, n=4 ; UAIC 10867.06 , n=7, Alabama, Blount County, Mill Creek at Old AL Hwy 75, east of AL Hwy 75, T13 S, R1 E, S1, 33.927813 -86.496049, 2.9 km SSW Oneonta , 16 June 1993, B. R. Kuhajda and R. L. Mayden ; UF 188209 , n=2, same data as holotype ; TU 204140 , n=2, same data as UAIC 10867.06 . UAIC 14799.01 , n=2, same locality data as UAIC 10867.06, 7 October 2005, B. R. Kuhajda ; UAIC 16061.01 , n=3, Alabama, Blount County, Calvert Prong of the Little Warrior River at AL Hwy 75, 5 km NE of Oneonta, T12 S, R2 E, S16, 33.985578 -86.442009, 2 April 2016, B. R. Kuhajda and Freshwater Fishes of Alabama class .

Diagnosis. A member of the Etheostoma chermocki species group as identified by Boschung et al. 1992, Clabaugh et al. (1996) and Boschung and Mayden (2004). Etheostoma chermocki was diagnosed from the Etheostoma bellator complex by Boschung et al. (1992) and Suttkus and Bailey (1993). Etheostoma kimberlae is distinguished from E. michellae new species using the following combination of characters: cream-colored horizontal stripe through lateral band very narrow to absent vs. broad and obvious; snout of breeding males lightly colored with obvious preorbital stripe in breeding males vs. snout dusky and often masking preorbital stripes; suborbital bar diffuse in breeding males, lightly pigmented and short vs. dark, well defined and long.; lateral blotches dark and largely below lateral line vs. diffuse and occurring both above and below lateral line; and crimson line above lateral line formed from spots in continuous to nearly continuous line and not interrupted by lateral blotches vs. crimson line broken up into segments between diffuse upper halves of lateral blotches. Etheostoma kimberlae is further distinguished from E. michellae in the possession of the following alleles at identified protein loci: sAat-A (A vs B)†, mAcon-A (B vs B, C) *, Acp-1 (A vs B)†, Ada-2 (A vs B)†, Est-2 (D vs A)†, Pep-B (B, C vs B) *, Pep-F (D vs B)†, Pnp-A (D vs C, D) *. Etheostoma kimberlae is distinguished from E. bellator using the following characters: cream-colored stripe through lateral band narrow to absent vs broad and obvious; snout lightly colored with obvious preorbital stripes in breeding males vs. snout dusky and often masking preorbital stripes; and suborbital bar diffuse, lightly pigmented and short vs. broad, dark and long. Etheostoma kimberlae is further distinguished from E. bellator in the possession of the following alleles at identified protein loci: sAat-A (A vs B)†, Acp-1 (A vs B)†, Ada-2 (A vs B)†, Pep-B (B, C vs A, B; allele B fixed in comparisons between Five Mile and Gurley creeks but is polymorphic in Murphy Creek) *, Pep-F (D vs B, D) 3, Pnp-A (D vs C, D) *. Etheostoma kimberlae is distinguished from E. chermocki in having a low first dorsal fin with black, red, and blue coloration vs fin tall and mostly red, and ventral orange pigmentation in breeding males in narrow band vs. extending upwards along side to lateral band. Etheostoma kimberlae is further diagnosed from E. chermocki by the possession of the following alleles at identified protein loci: sAat-A (A vs D)†, Acp-1 (A vs B)†, Ada-2 (A vs B)†, Fbp-A (A vs B)†, slcdh-A (B vs A, C)†, Mpi-A (B vs B, C) *, Pep-B (B, C vs A)†, Pep-F (D vs B)†.

Lateral line scale rows of Etheostoma kimberlae usually 47 or 48 (vs. E. michellae with 45–48, usually 45, and 47–51 in E. bellator ). Scale rows below lateral line usually 7 or 8 (vs. usually 8 in E. michellae and 7 or 8 E. bellator ). Transverse scale rows usually 15 (vs. usually 14 and 15 in E. michellae ). Transverse scale rows plus scale rows below lateral line usually 15 (vs. usually 14 or 15 in E. michellae and E. bellator ). Dorsal fin rays usually 11 or 12 (vs. usually 11 or 12 in E. michellae and 11 in E. bellator ). Pectoral fin rays usually 13 (vs. usually 14 in E. michellae and E. bellator ). Caudal fin rays usually 16 or 17 (vs. usually 16 or 17 in E. michellae and 14 in E. bellator ).

Etheostoma kimberlae differs from E. michellae in having usually 7 infraorbital pores (mean = 7.1) (vs. usually 8, mean = 8.1). Etheostoma kimberlae is a small member of the E. chermocki species group, reaching recorded 45.0 mm SL (male) and 41.4 mm SL (female) vs. E. bellator (male = 58 mm SL, Suttkus & Bailey 1993; female = 48.2 herein) and E. chermocki (male = 55 mm SL; female = 51 mm SL; Boschung et al. 1992).

Description. General head and body shape and coloration illustrated in Figure 3 . Morphometric variables for males and females (sexually dimorphic) provided in Table 1 . Distribution of lateral line scales, caudal fin rays, and pectoral fin rays provided in Table 2 .

Lateral line complete and virtually straight, from upper margin of gill opening to base of caudal fin. Scale rows above lateral line 4 (3 spms), 5 (34) or 6 (3) (Mean = 5.0, SD = 0.37). Scale rows below lateral line 6 (3), 7 (7), 8 (26), or 9 (4) (Mean = 7.8, SD = 0.84). Transverse scale rows 14 (6), 15 (27), or 16 (7) (Mean = 14.8, SD = 0.88). Transverse scale rows plus scale rows below lateral line 13 (5), 14 (6), 15 (22), 16 (6) or 17 (1) (Mean = 14.8, SD = 0.98). Caudal peduncle scale rows 10 (3), 11 (20), 12 (15) or 13 (2) (Mean = 11.3, SD = 0.82). Dorsal saddles 8 (40). Dorsal fin spines 6 (3), 7 (20), 8 (13), or 9 (1) (Mean = 7.3, SD = 0.67). Dorsal fin rays 9 (1), 10 (3), 11 (20), 12 (14), or 13 (2) (Mean = 11.3, SD = 0.82). Anal spines = 2 (40). Pelvic spines = 1 (40). Pelvic rays = 5 (40).

Infraorbital pores 6 (2 spms), 7 (5) or 8 (3) (Mean = 7.1, SD = 0.74). Preopercularmandibular pores 7 (1), 8 (5) or 9 (2) (Mean = 8.3, SD = 0.67). Lateral canal pores 2 (10). Supratemporal canal complete, pores 2 (10). Supraorbital pores 3 (10). Coronal pore single (10).

Nape scaled. Belly fully scaled. Breast naked.

Coloration of breeding males. Dorsum of head and body cream colored except for series of black saddles on body, normally numbering 8. Dorsolateral scales lightly pigmented along edges. Lateral blotches black and continuous or nearly so across very narrow light cream line along lateral line. Scale row directly above lateral line deep crimson and continuous or nearly continuous; lateral blotches not oval; above lateral line lateral blotch small, circular, short and not distinct; most of lateral blotch below lateral line and obvious; above lateral line, blotches not clearly separated. Below lateral line single crimson-colored scale row separating dark blotches. Crimson line usually one scale row and dissipating posteriorly directly vertical origin to middle of second dorsal fin. Dark caudal spot present; above and posterior to caudal spot short crimson stripe; stripes overlapping light cream-colored spots dorsally and ventrally at base of fin. Some ventrolateral scales lightly pigmented with melanophores, especially along distal portion of scales. Broad reddish-orange ventrolateral stripe present and involving multiple scale rows, extending length of body and 3–4 scale rows from above base of anal fin and pelvic fin insertion. Ventrolateral color stripe narrow and not connected with crimson scales around complex lateral stripe.

Opercle darkly pigmented dorsally becoming lighter ventrally; dark oblique line between cheek and preopercle; remainder of opercle lightly pigmented to immaculate; subopercle, branchiostegals, under side of head, preorbital area, breast and tip of snout turquoise. Distinct narrow suborbital bar, dark preorbital bar, middle of upper lip pigmented; remainder of lips yellowish to turquoise. Snout lightly pigmented and in distinct contrast to preorbital bars.

Pectoral-fin rays pigmented and with two stripes, one a basal band covering 1/3 of rays and boarded distally by turquoise colored stripe; distal to this basal band light area, bordered distally by darker stripe, and distal-most 1/3 of fin with lightly pigmented rays; membranes without pigmentation. Base of fin and area of insertion reddish-orange. Spine of pelvic fin immaculate to turquoise; remaining rays turquoise to darkly pigmented. Base of fin and first three branched rays turquoise. Posterior rays immaculate. Spine and first three rays also pigmented with melanophores medially, creating a dark stripe; all membranes turquoise. Spinous dorsal fin with four stripes. Base of fin with narrow dark green stripe, area between this stripe and adjoining more distal stripe narrow and light cream. Distal dark stripe with pigment on rays and membranes. Subdistal stripe beginning with dark red irregular shaped spot between first and second spines and stripe that is less intense in color, continuing posteriorly from being narrow in anterior membranes and significantly increasing in width towards last membranes. Narrow blue distal band beginning at sixth spine and extending to posterior of fin. Distal margin of anterior five rays clear. Distal-most black stripe separated from brick red stripe by narrow clear stripe. Soft (second) dorsal fin with three distinct bands. Distinct broad median brick red stripe below and above darkly pigmented median stripe formed from pigmented rays and membranes. Basal half of first three rays may have alternating black and yellow/gold narrow stripes. Procurrent, principle and two adjacent branched rays of caudal fin turquoise. Two-three dark vertical bands located medially on caudal fin separated by yellow- to cream-colored narrow bars; distal margin lightly pigmented, appearing almost transparent.

Coloration of Preserved breeding males. Adult males with dark brown dorsal saddles extending from occiput to posterior extent of caudal peduncle; occasionally blotches may be connected; blotches formed from dense concentrations of melanophores, and separated by pale areas having only light concentrations of melanophores. Dorsolateral area above lateral line with stripe, usually 1.5 to 2.5 scales high, extending posteriorly to vertical of middle of second dorsal fin to near hypural plate; stripe formed largely from dorsal halves of slightly lighter scale areas relative to dark lateral blotches (hereafter referred to as lighter) that, in live specimens, were dark red; dark lateral blotches weakly developed above lateral line and most strongly developed below lateral line where appearing somewhat oval in shape; blotches separated by lighter scale areas. Posterior to middle of second dorsal fin base dark lateral blotches connecting across and centered on lateral line; lighter intervening scale areas only present above lateral line. Lighter blotches below lateral line forming continuous stripe to hypural plate, becoming darker and centered on lateral line posterior to middle of second dorsal fin. Belly and breast lightly pigmented.

Dorsum of head tan to brown. Two horizontally rectangular dark blotches immediately posterior to head along lateral line poorly developed, usually small, if present, and consisting of broken lines. Snout with melanophores uniformly distributed except for distinct dark line connected to preorbital strip; line directly above upper lip and continuous across snout in adult males; smaller males with interruption in snout band; upper lip pigmented medially. Suborbital bar present; bar most distinct in females. Postorbital stripe usually not continuous but broken into two distinct, dark spots; first spot immediately posterior to orbit and second at junction of dorsal arm of preopercle and anterodorsal area of opercle. Remainder of cheek, opercle, preopercle, and subopercle only lightly pigmented except for distinct irregularly shaped concentrations of melanophores formed on the cheek and upper operculum. Cheek spot located slightly ventral and posterior to postorbital spot immediately behind orbit. Opercular spot located near center or posterodorsal area of opercle below dorsal margin of opercle. Lower cheek, branchiostegals, and gular areas with light scattering of melanophores to immaculate; lower lip immaculate.

Color pattern of dorsal fins as described for live specimens except colors muted.Caudal fin with light pigmentation on rays only, forming two to four bands. Anal and pelvic fins with few melanophores on membranes and little to no pigment on rays. Pectoral fin with bars formed from melanophores on rays separated by depigmented portions of rays; membranes clear. Spot at base of caudal fin only darkly pigmented like lateral blotches and surrounded by pale oval areas.

Coloration of live breeding females. Without bright coloration. Dorsum of body with distinct dark dorsal saddles separated by lighter scales cream in color. Dorsolateral scales with cream bases and darker posterior edges; some scales darker than others and in short continuous lines, creating mottled appearance; scattered scales partially brick red to orange. Scale row directly above lateral line brick red to orange and continuous or nearly continuous. Lateral blotches mostly below lateral line where dark pigment covers 1–2 rows of scales; blotches, if present, above lateral line may be half scale row. Anterior five blotches with oblique line of pigmented scales (best developed anteriorly) directed anteroventrally; posterior blotches with short (1–2 scales) pigmented along ventral margins. Scattered scales one scale row below lateral-line and between lateral blotches may be orange. Some ventrolateral scales may have some orange in center of scale.

Dorsum of head dark brown; preorbital and postorbital stripes and suborbital bar brown and well developed. Preorbital stripes connecting along pigmented upper lip. Cheek with distinct dark blotch posteriorly; opercle mottled. Venter of head and body cream. Pectoral, second dorsal, and caudal with pigment on rays only, creating lines on fins; membranes transparent. First dorsal fin with dark pigment on rays and membranes, small red to orange blotch in first membrane. Anal fin immaculate except for pigment on membranes between spines and posterior spine and first ray. Spot at base of caudal fin darkly pigmented and surrounded by pale cream oval areas.

Coloration of preserved females and juveniles. Dorsum of body with dark saddles; areas between saddles and ventrally to lateral line somewhat lighter but less obvious than in males; most scales with darkened centers, creating a mosaic pattern of dark and light irregularly shaped spotting pattern. Below lateral line distinct dark and squarish blotches, 3–4 scales deep, separated by areas of near equivalent width with light or depigmented scales; most larger females with distinct narrow anteroventral extension of dark lateral blotch, usually connected to blotch and sometimes extending anteriorly to connect with or nearly connect with immediate blotch anteriorly; anteroventral extensions of dark blotches creating pattern of pale circles or ovals between dark blotches below lateral line. Lighter pigmented line, as described in males, not as well developed and occurring below lateral line posteriorly to usually origin of second dorsal fin. Narrow light line below lateral line only anteriorly, extending posteriorly to near terminus of first dorsal fin. Ventrolateral areas, belly, breast and lower caudal peduncle immaculate.

Head coloration as in males except for the following; preorbital stripe present, suborbital bar usually well developed, and concentrations of melanophores on check and posterodorsal area of opercle darker and may include more than two spots.

Membranes of dorsal fins clear; rays with concentrations of melanophores separated by areas of same size with no melanophores, creating a pattern of bands in the fin; no indication of a red blotch on first membrane of first dorsal fin. Like dorsal fins, caudal and pectoral fins with melanophores on rays and melanophore distributions form two or more bands. Anal and pelvic immaculate. Spot at base of caudal fin distinct, similar in intensity as dark lateral blotches, and surrounded by pale ovals.

Distribution and Habitat. This species is only known from the upper Locust Fork of the Black Warrior River drainage ( Figure 5 ). It is presently known from Calvert Prong at AL Hwy 75 just north of Oneonta and a tributary to Calvert Prong, Mill Creek, in and around Oneonta, Blount County, Alabama. It is known historically from another Calvert Prong tributary, Sand Valley Creek, Blount County, Alabama (UMMZ 158288, 1939) and Little Cove Creek, Etowah County, Alabama (UAIC 3307.11, 1969) ( Suttkus & Bailey 1993, Kuhajda 2004a). No systematic status survey has been done for this species, and it is predicted to occur at other localities in the Calvert Prong system and different streams of geologies dominated by the Chapultepec and Copper Ridge Dolomites undifferentiated formation in Blount and Etowah counties. The species is found in small to moderate-sized upland creeks associated with moderate to little current over a sand/gravel to cobble substrate, typically in the glide above riffles and in the transition to pool habitat below riffles, but can occur in riffles proper ( Kuhajda 2004a).

Relationships. Clabaugh et al. (1996) discussed possible relationships of this species relative to others of the E. chermocki group based on allozyme variation. Near et al. (2011), using sequence variation, identified E. kimberlae (therein referred to as E. cf. bellator (Locust Fork Darter) as the sister species of a clade composed of E. bellator and E. chermocki . Interestingly, in all of the supplemental dendrograms by Near et al. (2011) (cytb, S7 intron 1, RAG1) presented for the group, there are no data for the Locust Fork Darter. Yet the species appears in the concatenated dendrogram in the body of the paper (two specimens each of Locust Fork and Sipsey Fork darters). In the definition of the clade Adonia, Near et al. (2011) mentions the Locust Fork Darter but not the Sipsey Darter. Kim et al. (2023) provide a short dialog about the genetic variation in this group relative to the geological formations in the area. Like Clabaugh et al. (1996), using only a relatively small number of allozyme loci, Kim et al. (2023), using 25,393 ddRAD loci revealed the exact similarities between E. bellator (sensu lato), E. michellae , and E. kimberlae in both the maximum likelihood concatenated dendrogram and the unrooted PoMo dendrogram. Etheostoma kimberlae forms the basal sister group to all other members of the E. chermocki clade. In their analysis, the authors confidently discuss the existence of not only E. bellator in the Mulberry Fork of the Black Warrior River but also two undescribed forms in Valley and Gurley creeks (no data were provided to substantiate any of the forms). Hence, this is why we refer to E. bellator as sensu lato. Neither lineage was identified in our analyses (genetic or morphological), the coloration of breeding and nonbreeding males and females based on descriptions, personal observations, and photographs of breeding and nonbreeding coloration, meristic, morphometrics, and morphological observations analyses.

Etymology. This species is named after my daughter Kimberly Linnae Mayden. The common name Locust Fork Darter refers to the river system where Etheostoma kimberlae is endemic.

Conservation Status.

Species is currently only recorded from Calvert Prong north of Oneonta and approximately 5 km of Mill Creek in and surrounding Oneonta, Blount County, Alabama ( Kuhajda 2004a). Given the very small distribution of E. kimberlae and localized industrial activities, urbanization, and agriculture, this species is endangered ( Jelks et al. 2008) and needs to be considered for State and Federal protection. It is currently listed as a species of High Conservation Concern (P2) by the State of Alabama ( Kuhajda 2004a, ADCNR 2015) and S1 by the Alabama Natural Heritage Program but is not on the State list of protected species ( ALNHP 2015). A complete systematic status survey is needed ( Kuhajda 2004a), followed by long-term monitoring of extant populations.

Figures

FIGURE 1. Male (Top) and Female (Midde) Vermilion Darter, Etheostoma chermocki Boschung, Mayden, & Tomelleri. Images from Boschung, Mayden & Tomelleri (1992) and Boschung & Mayden (2004; plate 80B and A) and copyrighted by Joseph R. Tomelleri (used with permission). (Bottom) General habitat of E. chermocki, Turkey Creek, Jefferson Co., Alabama, 12 July 2014. Photography copyrighted by B. R. Kuhajda.

FIGURE 3. (Top) Male, Locust Fork Darter, Etheostoma kimberlae new species Mayden. Images from Boschung & Mayden (2004; plate 79D). Image copyrighted by Joseph R. Tomelleri (used with permission). (Bottom) General habitat of E. kimberlae, Calvert Prong Creek, Jefferson Co., Alabama, 18 February 2017. Photography copyrighted by B. R. Kuhajda.

FIGURE 5. Distributions of Etheostoma bellator (circles), E. kimberlae (black squares), and E. michellae (gray squares) in the river systems of Alabama. The inset to the right represents the distributions of these species relative to the major river systems of the Southeastern United States. Images of each species are provided for reference. Maps modified from Boschung & Mayden (2004) and used with permission. Images of darters copyrighted by Joseph R. Tomelleri (used with permission.)

Tables

TABLE 1. Body and head measurements as proportions of standard or head length for E. bellator, E. michellae, and Etheostoma kimberlae. D1 = First dorsal Fin, D2 = second

Males	Etheostoma bellator (N = 19)				Etheostoma michellae (N = 20)				Etheostoma kimberlae (N = 20)
	Min	Max	Mean	SD	Min	Max	Mean	SD	Min	Max	Mean	SD
SL	43.75	54.08	48.81	3.09	29.87	44.94	38.40	4.46	36.05	47.17	41.35	2.87
Percent standard length
HL	0.21	0.27	0.24	0.01	0.23	0.25	0.24	0.01	0.23	0.26	0.25	0.01
BW	0.13	0.15	0.14	0.01	0.10	0.14	0.12	0.01	0.11	0.14	0.12	0.01
HW	0.13	0.15	0.14	0.01	0.12	0.15	0.14	0.01	0.12	0.15	0.14	0.01
Tip of snout to PC base lower	0.23	0.26	0.24	0.01	0.22	0.25	0.24	0.01	0.23	0.25	0.24	0.00
Posterior orbit to origin of D1	0.20	0.22	0.21	0.01	0.20	0.23	0.21	0.01	0.19	0.22	0.21	0.01
Origin D1 to origin to D2	0.27	0.31	0.30	0.01	0.28	0.32	0.30	0.01	0.27	0.31	0.28	0.01
Origin D1 to origin AF	0.31	0.36	0.34	0.01	0.30	0.35	0.33	0.01	0.31	0.34	0.33	0.01
Origin of D2 to lower PC base	0.37	0.41	0.39	0.01	0.37	0.39	0.38	0.01	0.35	0.38	0.36	0.01
D2 base length	0.15	0.21	0.18	0.01	0.14	0.19	0.17	0.01	0.16	0.21	0.19	0.01
Origin D2 to origin AF	0.16	0.20	0.18	0.01	0.15	0.18	0.17	0.01	0.15	0.18	0.17	0.01
Posterior of D2 base to posterior of AF base	0.13	0.16	0.14	0.01	0.11	0.16	0.13	0.01	0.11	0.15	0.13	0.01
Caudal peduncle depth	0.09	0.11	0.10	0.01	0.10	0.11	0.10	0.00	0.09	0.11	0.10	0.01
Posterior of D2 base to hypural plate	0.21	0.25	0.22	0.01	0.21	0.27	0.24	0.01	0.21	0.26	0.23	0.01
Caudal peduncle length	0.28	0.32	0.30	0.01	0.29	0.32	0.30	0.01	0.28	0.32	0.30	0.01
Base of AF	0.11	0.14	0.13	0.01	0.09	0.14	0.12	0.01	0.10	0.15	0.13	0.01
Origin AF to PC base lower	0.31	0.36	0.34	0.01	0.32	0.36	0.34	0.01	0.31	0.36	0.34	0.01
PC base	0.05	0.07	0.06	0.00	0.05	0.06	0.06	0.00	0.05	0.07	0.06	0.00
Percent head length
Interorbital width	0.16	0.22	0.18	0.02	0.14	0.21	0.18	0.02	0.16	0.21	0.19	0.01
Preorbital	0.24	0.33	0.27	0.02	0.21	0.29	0.24	0.02	0.22	0.28	0.26	0.02
Snout to end of maxilla	0.21	0.35	0.25	0.03	0.22	0.28	0.25	0.02	0.18	0.29	0.24	0.03
End of maxilla to PC base lower	0.65	0.86	0.72	0.05	0.68	0.82	0.73	0.03	0.66	0.76	0.71	0.02
End of maxilla to PC base upper	0.70	0.90	0.76	0.05	0.72	0.93	0.78	0.05	0.72	0.85	0.77	0.04
Orbit width	0.23	0.31	0.26	0.02	0.24	0.30	0.27	0.02	0.22	0.28	0.26	0.01
Posterior orbit to PC base lower	0.51	0.66	0.55	0.03	0.51	0.61	0.56	0.03	0.49	0.62	0.54	0.03

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