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Hildenbrandia
The darker red alga encrusting this rock fragment is H. crouaniorum
SEM of a H. rivularis gemma. Scale bar: 50 μm
Scientific classification
Clade:	Archaeplastida
Division:	Rhodophyta
Class:	Florideophyceae
Order:	Hildenbrandiales
Family:	Hildenbrandiaceae
Genus:	Hildenbrandia Nardo, 1834
Synonyms
Hildenbrandtia

Hildenbrandia is a genus of thalloid red alga comprising about 26 species. The slow-growing, non-mineralized thalli take a crustose form. Hildenbrandia reproduces by means of conceptacles and produces tetraspores.

Morphology

Hildenbrandia cells are around 3–5 μm in diameter and the filaments are around 50–75 μm in height.

The thallus comprises two layers: the hypothallus, which attaches to the rock, and the perithallus, a pseudoparenchymous layer comprising vertical filaments, which unlike coralline red algae is not further differentiated.

Growth

Hildenbrandia comprises orderly layers of vertical oblong cells with thick vegetative cell walls, occasionally connected by secondary pit connections with pit plugs in the septal pores. It grows at its margins, away from the centre, and is able to quickly repair any gaps arising by regenerating from a basal layer of cells. As plants become more mature, they become multi-layered and strongly pigmented near their centres, whilst their single-layered margins begin to grow more slowly. Multi-layered areas may develop in the margins; these will detach and float away as gemmae to form new colonies, leaving a single layer of cells beneath them once they separate from the host plant.

Newly settled gemmae form rhizoids.

Conceptacles develop in a haphazard manner; cells in conceptacle regions deform one another and become less regularly shaped as they grow larger.

In a similar fashion to the coralline algae, the outer layer of the thallus is shed seasonally, presumably to avoid colonization by epiphytes.

Taxonomy

The genus name of Hildenbrandia is in honour of Franz Xaver von Hildenbrand (1789-1849), who was an Austrian physician and botanist.

The genus was circumscribed by Giovanni Domenico Nardo in Isis (Oken) vol.27 on page 675 in 1834.

Habitat

The freshwater species H. rivularis and H. angularis seems to form a clade, and require an alkaline pH and hard water, preferring clean water. Unlike most other freshwater red algae (which prefer running water), H. rivularis prefers still water, particularly shady lakes or ponds. H. rubra and other marine species are found in brackish waters, but freshwater / gemma-bearing species cannot tolerate even moderate salinities. The genus is often found in a symbiotic partnership with fungi. Hildenbrandia has a remarkable tolerance to stresses including extreme temperatures, desiccation, and Ultra-violet light; it can be up and photosynthesizing near full capacity just minutes after being cooled to −17 °C or subjected to extreme salinities.

Reproduction

Sexual reproduction has never been observed in any Hildenbrandia species. It can reproduce by splitting into multiple colonies by fragmentation, or via stolons (i.e. sending out lateral branches) or gemmae.

Marine Hildenbrandia, on the other hand, reproduce by means of tetraspores that are produced within the thallus by conceptacles.

Systematics

The genus contains these species (this list is out of date):

• H. angolensis
• H. arracana
• H. canariensis
• H. crouanii
• H. crouaniorum
• H. dawsonii
• H. deusta
• H. expansa
• H. galapagensis
• H. kerguelensis
• H. lecannellieri
• H. lithothamnioides
• H. nardiana
• H. occidentalis
• H. pachythallos
• H. patula
• H. prototypus
• H. ramanaginaii
• H. rivularis
• H. rosea
• H. rubra
• H. sanjuanensis
• H. yessoensis

As of April 2022, the GBIF only accepts 6 species; Hildenbrandia crouaniorum J.Agardh, Hildenbrandia dawsonii, Hildenbrandia occidentalis Setch., 1917, Hildenbrandia rivularis (Liebman) J.Agardh, Hildenbrandia rubra (Sommerfelt) Meneghini and Hildenbrandia sanjuanensis.

Stonehenge

The presence of H. rivularis near Stonehenge has been put forward as a reason for the site's perceived mystical properties. Flint in the Blick Mead spring pools near to the henge takes on a pink hue a couple of hours after being taken out of water due to the presence of the algae. It is assumed that ancient hunter-gatherers would have seen the rocks as having magical properties and would have deemed the site worthy of interest.

References

1. Dethier, M. (1994). "The ecology of intertidal algal crusts: variation within a functional group". Journal of Experimental Marine Biology and Ecology. 177: 37–71. doi:10.1016/0022-0981(94)90143-0.
2. Sherwood, A.; Sheath, R. (2000). "Biogeography and systematics of Hildenbrandia (Rhodophyta, Hildenbrandiales) in Europe: inferences from morphometrics and rbcL and 18S rRNA gene sequence analyses". European Journal of Phycology. 35 (2): 143–152. doi:10.1080/09670260010001735731.
3. "Hildenbrandia Ben: Morphology". washington.edu.
4. Cabioch, J.; Giraud, G. (1982). "La structure hildenbrandioïde, stratégie adaptative chez les Florideés". Phycologia (in French). 21 (3): 308–315. doi:10.2216/i0031-8884-21-3-307.1.
5. ^ Pueschel, C. (1982). "Ultrastructural observations of tetrasporangia and conceptacles in Hildenbrandia (Rhodophyta: Hildenbrandiales)". European Journal of Phycology. 17 (3): 333–341. doi:10.1080/00071618200650331.
6. ^ Wayne Nichols, H. (1965). "Culture and development of Hildenbrandia rivularis from Denmark and North America". American Journal of Botany. 52 (1): 9–15. doi:10.2307/2439969. JSTOR 2439969.
7. ^ Sherwood, A. R.; Sheath, R. G. (2000). "Microscopic analysis and seasonality of gemma production in the freshwater red alga Hildenbrandia angolensis (Hildenbrandiales, Rhodophyta)". Phycological Research. 48 (4): 241–249. doi:10.1046/j.1440-1835.2000.00208.x. S2CID 84193742.
8. Pueschel, C. (1988). "Cell sloughing and chloroplast inclusions in Hildenbrandia rubra (Rhodophyta, Hildenbrandiales)". European Journal of Phycology. 23: 17–23. doi:10.1080/00071618800650021.
9. Burkhardt, Lotte (2022). Eine Enzyklopädie zu eponymischen Pflanzennamen [Encyclopedia of eponymic plant names] (pdf) (in German). Berlin: Botanic Garden and Botanical Museum, Freie Universität Berlin. doi:10.3372/epolist2022. ISBN 978-3-946292-41-8. Retrieved January 27, 2022.
10. Sherwood, A. R.; Sheath, R. G. (2003). "Systematics of the Hildenbrandiales (Rhodophyta): gene sequence and morphometric analyses of global collections". Journal of Phycology. 39 (2): 409–422. doi:10.1046/j.1529-8817.2003.01050.x. S2CID 86786840.
11. ^ Eloranta, P.; Kwandrans, J. (2004). "Indicator value of freshwater red algae in running waters for water quality assessment" (PDF). International Journal of Oceanography and Hydrobiology. XXXIII (1): 47–54. ISSN 1730-413X. Archived from the original (PDF) on 2011-07-27. Retrieved 2010-10-27.
12. ^ Sherwood, A. R.; Shea, T. B.; Sheath, R. G. (2002). "European freshwater Hildenbrandia (Hildenbrandiales, Rhodophyta) has not been derived from multiple invasions from marine habitats". Phycologia. 41: 87–95. doi:10.2216/i0031-8884-41-1-87.1. S2CID 84894072.
13. Saunders, G. W.; Bailey, J. C. (1999). "Molecular systematic analyses indicate that the enigmatic Apophlaea is a member of the Hildenbrandiales (Rhodophyta, Florideophycidae)". Journal of Phycology. 35: 171–175. doi:10.1046/j.1529-8817.1999.3510171.x. S2CID 84758802.
14. Garbary, D. (2007). "The Margin of the Sea". Algae and Cyanobacteria in Extreme Environments. Cellular Origin, Life in Extreme Habitats and Astrobiology. Vol. 11. pp. 173–191. doi:10.1007/978-1-4020-6112-7_9. ISBN 978-1-4020-6111-0.
15. Guiry, M.D.; Guiry, G.M. (2008). "Hildenbrandia". AlgaeBase. World-wide electronic publication, National University of Ireland, Galway. Retrieved 2009-04-18.
16. "Hildenbrandia Nardo, 1834". www.gbif.org. Retrieved 7 April 2022.
17. "Mesolithic settlement near Stonehenge: excavations at Blick Mead, Vespasian's Camp, Amesbury" (PDF). www.silversaffron.co.uk.
18. Jacques, David (2014). "Mesolithic settlement near Stonehenge: excavations at Blick Mead, Vespasian's Camp, Amesbury". Wiltshire Archaeological and Natural History Magazine. 107: 7–27.

External links

• Benjamin Weisgall. "Hildenbrandia spp. : the immortal red crust". FHL Marine Botany: an excellent, accessible overview of the genus.
• Images of Hildenbrandia at Algaebase

• Red algae genera
• Taxa named by Giovanni Domenico Nardo