Description
Aneides klamathensis is a large, robust-bodied member of the Aneides flavipunctatus complex with a standard length of 68.2 – 80.6 mm in males and 65.4 – 84.8 mm in females. Total length is inaccurate as most specimens of the species have regenerated tail tips. The flat head is very large with a broad, rounded snout. In the lateral view the snout extends beyond the mandible. The nasolabial protuberances are modest. The distance between the nostrils is less than the distance between the eyes. The nostrils are closer to the snout than the eye. The eyes are prominent and moderately large but do not protrude. The jaw muscles are prominent and bulging. This salamander has a varying number of premaxillary teeth: 4 - 8 in males, and 3 - 7 in females. There is a large, prominent oval-shaped mental gland that is wider than long. The integumentary grooves of the head are distinct. They have a well-defined neck (Reilly and Wake 2019).

The holotype has 15 costal grooves. The short, robust limbs consist of shorter forelimbs than hindlimbs. The limb interval, when adpressed along the body, is about 0.5 – 3 costal grooves in males and 2 – 4 in females. The digits are short, small, and not prominent. The hand phalangeal formula is 1 – 2 – 3 – 2 with the fourth finger being much shorter than the third, about the same length or shorter than the second. The foot phalangeal formula is 1 – 2 – 3 – 3 – 2 with the fifth toe being much shorter than the second and about the same or shorter than the second. The longest digits are slightly expanded at the tips in a scalloped fashion and has well-developed subterminal pads (Reilly and Wake 2019).

The short, robust tail is tapered at the tip and about as wide as it is high (Reilly and Wake 2019).

Previously thought to be a subspecies of Aneides flavipunctatus and extremely morphologically similar to others in the A. flavipunctatus complex, A. klamathensis distinguishes itself in several key ways. First, its standard size is between the smaller A. hardii, A. vagrans, and A. ferreus and the larger A. lugubris. It has a relatively large body, short limbs, and more robust and prehensile tail compared to other Aneides salamanders such as A. aeneus and A. hardii. Unlike A. niger, which is typically solid black in dorsal coloration, A. klamathensis has some small white speckles on its dorsal side. Aneides klamanthesis can be distinguished from A. iecanus by having fewer small dorsal iridophores (pigment cells) and an average of 17, not 16 trunk vertebrae. Lastly, A. klamathensis can be distinguished from A. flavipunctatus by geographical range and DNA sequences, however, they are morphologically difficult to differentiate in their contact zone in Humboldt County, California (Reilly and Wake 2019).

In life, A. klamathensis has a black background color that is densely covered with intense gold to grey frosting on the dorsal surface, intense white spotting or frosting on the lateral surfaces and arms and belly. The blackish ventral coloration has white speckles, more as body size increases. There is a frosted ecomorph, characterized by cream-colored splotches on its limbs. The eyes and pupils appear dark all over. In preservative, the dorsum is black except for some white speckles and splotches on the lateral margin of dorsal surface, lateral surfaces, dorsal surfaces of the hands and feet, the neck, and tail. The ventrum of the body is completely black. The mental gland and nasolabial protuberances and grooves are unpigmented to lightly pigmented. About half of the gular region unpigmented or pale. The undersides of the proximal portions of the limbs appear to lose pigmentation, and the hands and feet are unpigmented. The posterior 15 - 20% of the tail is also unpigmented (Reilly and Wake 2019).

Juveniles have greenish-grey frosting over their black background coloration. This frosting is especially intense on the flanks. There is a sharp boundary between the flank and the black ventrum. There are also small to moderately large whitish to cream or yellow spots on the dorsal surfaces of the limbs and widely scattered across the other dorsal surfaces (Reilly and Wake 2019).

There is slight sexual dimorphism with males having larger jaw muscles, relatively longer limbs, and more premaxillary teeth. There is also a frosted eco-morph with more cream-colored splotches on the limbs (Reilly and Wake 2019).

Distribution and Habitat

Country distribution from AmphibiaWeb's database: United States

U.S. state distribution from AmphibiaWeb's database: California, Oregon

View distribution map in BerkeleyMapper.

Aneides klamathensis occupies the northernmost regions amongst members in the A. flavipunctatus complex, residing in northern California and southern Oregon of the United States of America. Specifically, they can be found in Jackson County, Oregon and Del Norte, Siskiyou, and Humboldt Counties in California. In Jackson County, A. klamethensis ranges from the Applegate river drainage, southwards, and in Del Norte County, leading into the southern bank of the Smith River. Its range then extends south along the boundaries of the Van Duzen River, including the Klamath and Trinity rivers. Their primary elevation range for this species tends to be at 500 m or below, though it has been known to exist near Hilt, Siskiyou County, at roughly 1000 m in elevation (Reilly and Wake 2019).

Life History, Abundance, Activity, and Special Behaviors
Aneides klamanthensis is a terrestrial species found in moss-covered rockslides (rock talus microhabitats) and under surface covers in conifer forests (Hammerson 2004, Reilly and Wake 2019).

The species reproduces via direct development and exhibits the highest “paedomorphism index” amongst all other populations in the A. flavipunctatus complex based on the retention of juvenile cryptic coloration. Adults have more deeply embedded iridophores, resulting in a more copper rather than greenish coloration (Lynch 1981).

Aneides klamanthensis shares a contact zone with A. flavipuncatus where the two are morphologically indistinguishable. The zone is an approximately 3 km region between Dobbyn Creek and the town of Blocksburg (Reilly and Wake 2019).

Trends and Threats
As of 2020, A. klamathensis has not been assessed by the IUCN Red List and A. flavipunctatus was last assessed in 2004, essentially making the assessment for the whole complex. Based on IUCN’s 2004 assessment, the species is “Near Threatened” as a result of observed population decline caused, in part, by habitat destruction. However, some declines are occurring in areas where habitat is protected, and research is needed to better understand why these populations are declining (Hammerson 2004).

Possible reasons for amphibian decline

General habitat alteration and loss

Comments

The species authority is: Reilly, S. B., Wake, D. B. (2019). “Taxonomic revision of black salamanders of the Aneides flavipunctatus complex (Caudata: Plethodontidae).” PeerJ, 7(e:7370):1–36.

Aneides klamathensis is closely related to other species in the Aneides flavipunctatus complex, especially A. iecanus, A. niger, and A. flavipunctatus. Trees generated by mtDNA suggest that A. klamathensis and A. iecanus are sister taxa, whereas *BEAST analysis at 13 nuclear DNA loci indicates that A. klamathensis and A. flavipunctatus are sister taxa. This is supported by the combined mtDNA and nuclear DNA data that generate a tree indicating A. klamathensis and A. flavipunctatus are sister taxa (Reilly and Wake 2019).

Aneides klamathensis shares an approximately 3 km contact zone with A. flavipunctatus in Humboldt County, California, United States of America. This zone ranges from Dobbyn Creek to the town of Blocksburg, and results in infrequent migration and hybridization based on genetic analysis (Reilly and Wake 2019).

Population genetics of A. klamathensis indicate that the species was restricted to the Klamath River watershed during the Pleistocene and later colonized the Smith and Rogue River water sheds (Reilly and Wake 2019).

The species epithet, “klamathensis”, is in reference to the Klamath Mountains in northern California. Its river, the Klamath River (and tributaries), runs through much of the range of this salamander. The Klamath Mountains are also a unique geomorphic region, whose orientation is east-west rather than north-south like many other mount ranges in the region. The Klamath Mountains are known for their high level of endemism and species diversity (Reilly and Wake 2019).

Aneides klamathensis has been referred to as Aneides “sequoiensis'', however, this is an invalid name as the description was never formally published and the type locality was in the geographic range of A. flavipunctatus (Reilly and Wake 2019).

References

Hammerson, G. (2004). “Aneides flavipunctatus.” The IUCN Red List of Threatened Species 2004: e.T59116A11884308. https://dx.doi.org/10.2305/IUCN.UK.2004.RLTS.T59116A11884308.en. Downloaded on 10 August 2020.

Lynch, J. F. (1981). ''Patterns of ontogenetic and geographic variation in the Black Salamander, Aneides flavipunctatus.'' Smithsonian Contributions to Zoology, (324), i-iv, 1-53.

Reilly, S.B., Wake D.B. (2019). ''Taxonomic revision of black salamanders of the Aneides flavipunctatus complex (Caudata: Plethodontidae).'' PeerJ, 7, e7370 . [link]

Species Account Citation: AmphibiaWeb 2020 Aneides klamathensis: Klamath Black Salamander <https://amphibiaweb.org/species/9046> University of California, Berkeley, CA, USA. Accessed Apr 15, 2024.

Citation: AmphibiaWeb. 2024. <https://amphibiaweb.org> University of California, Berkeley, CA, USA. Accessed 15 Apr 2024.

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