Assessed as Endangered A2c; B2ab(i,ii,iii,iv,v), ver 3.1; date assessed: August 30, 2020

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Common name(s): Mid-Atlantic Comb-over, Beach Broccoli

Taxonomic Notes: This species was included within Cladonia mitis until Evans (1943) separated it on the basis that C. submitis occupies a different habitat, produces unique secondary compounds, and has shorter, more robust podetia. Four subspecies of C. submitis were also recognized by Evans (1943), and while they have not been subsequently investigated in detail, they are not presently recognized as taxonomically distinct (Hoffman et al. 2020). The reported occurrences of this species in east Asia are very likely ascribable to a different taxon. While the individuals do share some characteristics (e.g. production of usnic acid and pseudonorrangiformic acid), there are notable morphological differences. The podetia of North American individuals are thicker and coarser than Asian individuals (Ahti 1961). Furthermore, the North American populations occur in coastal sandy habitats while the Asian populations occur on rocky outcrops at higher elevations (Hoffman et al. 2020). Thus, the Asian occurrences are treated as a distinct taxon, and C. submitis isconsidered endemic to eastern North America.

ASSESSMENT JUSTIFICATION [criteria: A2c; B2ab(i,ii,iii,iv,v)]

Cladonia submitis is an endemic macrolichen native to the Mid-Atlantic coast of North America, inhabiting primarily pine barren and sand dune habitat. It has a restricted area of occupancy (AOO = 196 km²), and is considered to be severely fragmented. The species shows evidence of decline due to habitat loss resulting from land use changes and development. Based on recent large-scale surveys, there hasbeen a 59.3% decline in AOO and a 39.3% decline in extent of occurrence (EOO) in the past century (Hoffman et al. 2020). A similar range of decline in the population size is inferred based on the reduction in EOO and AOO. Overall, therefore, the species is listed as Endangered under criteria A2c; B2ab(i,ii,ii,iv).

Assessor/s: Hoffman, J., Ohmura, Y. & Lendemer, J.; Reviewer/s: Yahr, R.; Facilitator(s) & Compiler(s): Chandler, A. & Allen, J.

Bibliography:

Ahti, T. (1961) Taxonomic studies on reindeer lichens (Cladonia, subgenus Cladina). Annales Botanici Societatis Zoologicae-Botanicae Fennicae Vanamo 32(1): 1-160.

Ahti, T., & Oksanen, J. (1990) Epigeic lichen communities of taiga and tundra regions. Vegetation 86: 39-70.

Bachman, S., Moat, J., Hill, A.W., de la Torre, J. & Scott, B. (2011) Supporting Red List threat assessments with GeoCAT: geospatial conservation assessment tool. In: V. Smith & L. Penev (eds) e-Infrastructures for data publishing in biodiversity science. Zookeys 150: 117–126.

Brodo, I.M., S.D. Sharnoff, & S. Sharnoff (2001) Lichens of North America. Yale University Press, New Haven and London.

Evans, A.W. (1943) Microchemical studies on the genus Cladonia, subgenus Cladina. Rhodora 45(539):417-438.

Hoffman, J.R., Ohmura, Y. & Lendemer, J.C. (2020) Combing for Beach Broccoli: surveys of the endemic macrolichen Cladonia submitis determines endangered status under IUCN guidelines. Biodiversity and Conservation 29: 2439–2456.

IUCN (2021) The IUCN Red List of Threatened Species. Version 2021-1. Available at: www.iucnredlist.org. (Accessed: 25 March 2021).

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Keeley, J. & Syphard, A. (2016) Climate change and future fire regimes: examples from California. Geosciences 6(3): 37.

Maikawa, E. & Kershaw, K.A. (1976) Studies on lichen-dominated systems. XIX. The postfire recovery sequence of Black Spruce–lichen woodland in the Abitau Lake region, NWT. Canadian Journal of Botany 54(23): 2679-2687.

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Ray, D.G., Calahan, G.D. & Lendemer, J.C. (2020) Factors influencing the persistance of reindeer lichens (Cladonia subgenus Cladina) within frequent-fire environments of the Mid-Atlantic Coastal Plain, USA. Fire Ecology 16(1): 1-18.

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MB# 365718

TYPE. UNITED STATES. MASSACHUSETTS, Plymouth County, Wareham, on rich humus and decayed wood, 1924, C.A. Robbins s.n. (FH, lectotype; US, isolectotype)

Description. Lichenized fungus.

Thallus dimorphic to fruticose. Primary thallus rarely seen, crustose, disappearing, visible only when growing on wood. Podetia yellow-green, highly branched with thick, twisted stems, 0.7-3 mm diam., forming prostrate colonies with erect branches in the center. Podetial surface ecorticate, strongly wrinkled in older stems, often with short, isidia-like side branches; medulla thick, compact.  Photobiont Asterchloris glomerata alga. Ascomata biatorine apothecia, brown. Pycnidia tiny, brown, often on branch tips.

Chemistry. UV-, K-, KC+ yellow, C-, PD-; usnic and pseudonorrangiformic acids.

Substrate and Habitat. On wood or humus over sand or sandy soil.

Distribution. Eastern North America, Japan(?); in North Carolina throughout.

Notes. Cladonia submitis is considered endemic to the Mid-Atlantic coast of eastern North America; records from Japan may be misidentifications (Hoffman et al. 2020).

Literature

Brodo, I.M., S. Duran Sharnoff & S. Sharnoff. (2001) Lichens of North America. Yale University Press, New Haven & London. 795 pp.

Evans, A.W. (1943) Microchemical studies on the genus Cladonia, subgenus Cladina. Rhodora 45: 417-438 (original description).

Hoffman, J.R., Y. Ohmura & J.C. Lendemer (2020). Combing for beach broccoli: surveys of the endemic macrolichen Cladonia submitis determines endangered status under IUCN guidelines. Biodiversity and Conservation 29: 2439–2456.

Lendemer, J. & N. Noell. (2018) Delmarva Lichens: an illustrated manual. Memoirs of the Torrey Botanical Society 28: 1-350.

Pino-Bodas, R. & S. Stenroos. (2021) Global Biodiversity Patterns of the Photobionts Associated with the Genus Cladonia (Lecanorales, Ascomycota). Microbial Ecology 82: 173–187.