Haplogloia andersonii (Farlow) Levring (BC)

Haplogloia andersonii (Farlow) Levring sensu lato consists of four genetic groups in BC. Individuals typically have an obvious coating of long dark brown assimilatory filaments (Image A). However, these filaments can be sparse or even absent on older regions of thalli (more so for some genetic groups than others), which can be compounded when individuals are viewed removed from water (Image B). Individual thalli are a few to 36 cm tall, typically prolifically branched to 3-4 orders and light to dark brown in colour bearing various degrees of long relative to short assimilatory filaments. However, the four genetic groups can be differentiated at the anatomical level.

Haplogloia sp. 1GWS, has a predominance of long assimilatory filaments near the tips (Image C), which are ephemeral to varying degrees as you move down the thallus. The primary lateral cells out to an including the basal cell to the primary assimilatory filaments produce rhizoids, which in turn bear secondary assimilatory filaments (Image D). Away from the tip a mix of short and long assimilatory filaments are present with the former typically clavate and composed of 4-5 square to squat cells, the lowest 5 µm wide by 3-5 µm tall while the terminal cells are typically 8-9 µm wide by 6-9 µm tall. Cells of the long assimilatory filaments are 6-10 wide by 3-14 µm long being shortest near the base (Image E). Unilocular sporangia are small, 15-22 by 20-30 µm, and moderately oblong to subspherical in shape (Image F), with the associated short assimilatory filament cells becoming rectangular near their base (compressed), 2-3 µm wide by 6-8 µm long, and curving to overtop the sporangia to take on the appearance of paraphyses (Image G). This is by far our most common genetic group with 18 confirmed collections ranging from California (n = 1) to Haida Gwaii, typically low intertidal to mid intertidal in pools.

Sp. 2GWS is the most similar to the image in Kylin (1940, fig. 14) in which occasional long assimilatory filaments are distributed among the short assimilatory filaments (Image H), the latter being composed of 4-6 squat to square cells with an overall clavate habit (Image I). Again unilocular sporangia are nested among these short assimilatory filaments, are broadly oval to spherical as in Kylin (1940), but whereas his drawings suggest these structures approach 30 µm in diameter, those in sp. 2GWS were only 16-21 µm in dimensions (Image J). We have only two collections for this group, both from the west coast of Vancouver island, one mid intertidal and the other subtidal (5 m).

Sp. 3GWS has a mix of long (many damaged in processing giving the shaggy appearance), as well as short, 4-6 celled, clavate assimilatory filaments (Image K). Cells of the short assimilatory filaments were rectangular low on the filaments (seemingly regardless to the presence of unilocular sporangia), 3-4.5 µm wide by 4-9 µm tall, with some filaments terminating in swollen cells (Image L). The sporangia were again elongate but more obovate than in the other genetic groups, 16-18 µm wide by 26-30 µm tall (Image M). This genetic group has a superficial resemblance to Haplogloia kuckuckia Kylin, but the material did not rehydrate well and key anatomical features could not be confirmed. We have only two collections for this group, both from the west coast of Vancouver island, and both from the mid intertidal.

Sp. 4GWS was the ‘fuzziest’ species and near the tips appeared to have exclusively long (many truncated from processing) assimilatory filaments (Image N). Closer inspection lower on the axes reveals definitive short and long assimilatory filaments (Image O), the former long with 7-10 cells, clavate in overall habit, the cells variously squat to rectangular and enlarging toward the tip (Image P), among which the unilocular sporangia were nested (Image Q). The latter were elongate oval, 24-30 µm wide X 44-50 µm tall being the largest of the four genetic groups (Image R). Our second most common genetic group, we have seven collections ranging from California (n = 2) to Haida Gwaii, from subtidal (5 m) to mid intertidal pools.

The occurrence of two species for this genus in BC, Haplogloia andersonii (Farlow) Levring and Haplogloia kuckuckii Kylin, as well as reports of Papenfussiella callitricha (Rosenvinge) Kylin, have been regarded with uncertainty (Gabrielson & Lindstrom 2018). This conundrum is made more complicated as we have uncovered four allied genetic groups in BC. A further complication is that the four nest within the current concept of Papenfussiella in preliminary phylogenetic analyses (Image S). Individuals in group sp. 4GWS can be particularly dark brown and fuzzy possibly accounting for records of Papenfussiella callitricha (Rosenvinge) Kylin in BC. Individuals from two of the groups have been collected in California to date (type locality of Haplogloia andersonii (Farlow) Levring), while all four genetic groups occur in southern BC close to the type locality in WA for Haplogloia kuckuckii Kylin. Considerable taxonomic work remains for this complex in our flora.

Image A. Individual of Haplogloia sp. 4GWS with an extensive coating of dark long assimilatory filaments (mid intertidal pool on rock, Bamfield, BC; GWS003383).

Image B. Individual of Haplogloia sp. 2GWS removed from water and appearing devoid of a covering of long assimilatory filaments (subtidal (5 m) on rock, Tahsis, Rosa Harbour, Esperanza Channel, BC; GWS010206).

Image C. Abundant long assimilatory filaments near the branch tips in sp. 1GWS (drift, ‘Lands End’ at Pachena Bay, BC; GWS008107; rehydrated, aniline blue stained).

Image D. Rhizoids (R) descending from lateral vegetative cells and the basal cell of an assimilatory filament, cells of which can bear short (S) and long (L) assimilatory filaments (sp. 1GWS; mid intertidal pool on rock, Raft Cove, Vancouver Island, BC; GWS004725; rehydrated, aniline blue stained).

Image E. Close up of a 5-celled short (S) assimilatory filament and of cells of a long assimilatory filament (L) in sp. 1GWS (mid intertidal pool on rock, Pachena Beach, Bamfield, BC; GWS008286; rehydrated, aniline blue stained).

Image F. Unilocular sporangia borne low on the short assimilatory filaments in sp. 1GWS (GWS004725; rehydrated, aniline blue stained).

Image G. Close up of the unilocular sporangia and short assimilatory filaments in sp. 1GWS (GWS004725; rehydrated, aniline blue stained).

Image H. Occasional long assimilatory filaments observed among the dominate short assimilatory filaments in sp. 2GWS (GWS010206; rehydrated, aniline blue stained).

Image I. Typically spherical unilocular sporangia nested among the 4-6 celled short assimilatory filaments in sp. 2GWS (GWS010206; rehydrated, aniline blue stained).

Image J. Close up of the spherical unilocular sporangia and short assimilatory filaments in sp. 2GWS (GWS010206; rehydrated, aniline blue stained).

Image K. View of the short and (typically broken) long assimilatory filaments at lower magnification for sp. 3GWS (mid intertidal on rock, Pachena Beach, Bamfield, BC; GWS003382; rehydrated, aniline blue stained).

Image L. Close up of the short assimilatory filaments some with inflated terminal cells in sp. 3GWS (GWS003382; rehydrated, aniline blue stained).

Image M. Close up of the unilocular sporangia in sp. 3GWS (GWS003382; rehydrated, aniline blue stained).

Image N. Squash mount displaying numerous long (many truncated from processing) assimilatory filaments in sp. 4GWS (mid intertidal on rock, Pachena Beach, Bamfield, BC; GWS003383; rehydrated, aniline blue stained).

Image O. Squash mount displaying numerous long (many truncated from processing), but also possibly a few distinct short assimilatory filaments (sp. 4GWS; GWS003383; rehydrated, aniline blue stained).

Image P. Closer inspection clearly reveals short assimilatory filaments that typically have more cells than observed in the other genetic groups (sp. 4GWS; GWS003383; rehydrated, aniline blue stained).

Image Q. Unilocular sporangia nested among curved and straight short assimilatory filaments for sp. 4GWS (subtidal (3 m) on rock, Seal Pt., Haida Gwaii, BC; GWS036829; rehydrated, aniline blue stained).

Image R. Close up of the unilocular sporangia in sp. 4GWS (GWS036829; rehydrated, aniline blue stained).

Image S. Preliminary phylogenetic tree using rbcL including three of the four groups of Haplogloia from BC nested among species of Papenfussiella (RaxML analyses completed in Geneious with GTR+I+G, partitioning by codon and 500 bootstrap replicates). Note that sp. 3GWS, not included in the rbcL analyses, is closest to sp. 1GWS (Haplogloia andersonii in this figure) in COI-5P.