The Red Algal Genus 

Audouinella Bory 

Nemaliales: Acrochaetiaceae) 

from North Carolina 

SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES • NUMBER 22 
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SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES-NUMBER 22 

The Red Algal Genus Audouinella Bory 

(Nemaliales: Acrochaetiaceae) 

from North Carolina 

Craig W. Schneider 

SMITHSONIAN INSTITUTION PRESS 

City of Washington 

1983 
ABSTRAC T 

Schneider, Craig W. The Red Algal Genus Audouinella Bory (Nemaliales: 

Acrochaetiaceae) from North Carolina. Smithsonian Contributions to the Marine 

Sciences, number 22, 25 pages, 3 figures, 1983.—A monographic study of 

Audouinella in the Atlantic waters of North Carolina reports 15 taxa from 

coastal and continental shelf habitats. Three of these, A. affinis, A. hoytii, and 

A. ophioglossa, herein described as a new species, are endemic. Audouinella bispora 

and A. daviesii are reported from the Carolina flora for the first time. The 

taxonomy from several historical reports is elucidated, and taxonomic confusions in this complex are clarified. 

OFFICIAL PUBLICATION DATE is handstamped in a limited number of initial copies and is recorded 

in the Institution's annual report, Smithsonian Year. SERIES COVER DESIGN: Seascape along the 

Atlantic coast of eastern North America. 

Library of Congress Cataloging in Publication Data 

Schneider, Craig W. 

The red algal genus Audouinella Bory (Nemaliales: Acrochaetiaceae) from North Carolina. 

(Smithsonian contributions to the marine sciences ; no. 22) 

Bibliography: p. 

Supt. of Docs, no.: SI 1.41:22 

1. Audouinella—Classification. 2. Algae—North Carolina—Classification. I. Title. II. Series. 

QK569.A27S36 1983 589.4'1 83-600261 
Contents 

Page 

Introduction 1 

Materials and Methods 2 

Acknowledgments 2 

Genus Audouinella 2 

Key to the Species of Audouinella from North Carolina 2 

Audouinella affinis (Howe and Hoyt), new combination 4 

Audouinella bispora (B0rgesen) Garbary 5 

Audouinella botryocarpa (Harvey) Woelkerling 5 

Audouinella corymbifera (Thuret) Dixon 6 

Audouinella dasyae (Collins) Woelkerling 7 

Audouinella daviesii (Dillwyn) Woelkerling 8 

Audouinella densa (Drew) Garbary 8 

Audouinella hallandica (Kylin) Woelkerling 9 

Audouinella hoytii (Collins), new combination 10 

Audouinella hypneae (B0rgesen), new combination 11 

Audouinella infestans (Howe and Hoyt) Dixon 12 

Audouinella microscopica (Nageli) Woelkerling 12 

Audouinella ophioglossa, new species 13 

Audouinella saviana (Meneghini) Woelkerling 15 

Audouinella secundata (Lyngbye) Dixon 16 

Literature Cited 17 

Figures 1-3 20 

in 

The Red Algal Genus Audouinella Bory 

(Nemaliales: Acrochaetiaceae) 

from North Carolina 

Craig W. Schneider 

Introduction 

The Acrochaetiaceae has received a great deal 

of systematic attention in the last half century 

(Drew, 1928; Hamel, 1928b; Papenfuss, 1945, 

1947; Abbott, 1962, 1968; Feldmann, 1962; 

Woelkerling, 1971, 1972, 1973a, 1973b; Dixon 

and Irvine, 1977; Garbary, 1978, 1979). Nevertheless, many of the species reported or described 

from North Carolina have received little or no 

attention since their early notation here (Howe 

and Hoyt, 1916; Hoyt, 1920; Williams, 1948, 

1951). The only taxa updated in a recent systematic listing for North Carolina (Searles and 

Schneider, 1978) were those species taxonomically treated in other floras. Therefore, the North 

Carolina checklist contained both recent and antiquated taxonomy in this family, as well as several unsubstantiated reports. This monograph 

was undertaken to verify and update the taxonomic record. In the process of this work additional taxa have been discovered in the North 

Carolina flora. 

Several genera have been described in Acrochaetiaceae and species have been variously assigned to them on the basis of chromoplast morphology and presence or absence of sexual reproduction (for example, see Garbary, 1979). Because 

Craig W. Schneider, Department of Biology, Trinity College, Hartford, CT 06106. 

generic concepts vary from study to study and 

because frequently utilized generic characters 

have proved unreliable (Woelkerling, 1971, 

1973b), I find that it is more pragmatic and less 

confusing to follow the single-genus concept for 

the Acrochaetiaceae previously utilized by B0rgesen (1915) and first argued for by Drew (1928). 

This approach has recently been followed by 

Dixon and Irvine (1977) and Garbary (1979). 

The genus with historical precedence is Audouinella Bory (1823:340), and its usage requires some 

new nomenclatural combinations herein proposed. With the advent of further culture work in 

this family, it is possible that in the future we will 

be able to resegregate the species into more than 

one genus based on more reliable characters, such 

as life history characteristics (West, 1968, 1969; 

Stegenga, 1978; Stegenga and van Wissen, 1979). 

Generic confusion aside, species concepts have 

of late also come under more critical examination. 

Several a priori standard specific characters such 

as mode of attachment and reproductive state 

have been criticized as taxonomic criteria (Borsje, 

1973; Stegenga and Borsje, 1976, 1977; Stegenga 

and Vroman, 1976; Stegenga and Mulder, 1979; 

Stegenga and van Erp, 1979). Broad circumscriptions of species utilized by some (e.g., Woelkerling, 1971, 1972, 1973a, 1973b; Stegenga and 

Vroman, 1976) are not universally accepted (e.g., 

Dixon and Irvine, 1977; Kornmann and Sahling, 

1 
SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

1978; Humm, 1979). The cultural experiments of 

Stegenga and his associates have paired "distinct" 

species as heteromorphic generations of one taxon 

in this family. For the present, however, I continue to follow Woelkerling's classical interpretation of species concepts within Audouinella until a 

more complete understanding of species based 

upon life history and other features can be synthesized. Included in the scheme listed below, for 

example, is Audouinella dasyae, which is probably 

a gametophytic generation of either A. saviana, A. 

hypneae, or A. bispora, or possibly, all of them 

(Stegenga and Borsje, 1976). One of the asexual 

taxa mentioned below could possibly be the sporophytic generation of Scinaia complanata (Collins) 

Cotton (van den Hoek and Cortel-Breeman, 

1970), though that remains to be determined. 

MATERIALS AND METHODS.—Where possible, 

observations, measurements, and drawings were 

made using living field-collected material or liquid-preserved material. Dried specimens were 

wetted and removed from herbarium sheets using 

10% aqueous sodium carbonate. Permanent slides 

were produced by either mounting the specimens 

in 20% Karo syrup, 1% aniline blue, 1 N HC1, 

3%-4% formalin in a ratio of 95: 1:2:2, or by 

staining with 1% aniline blue, fixing with 1% 

HC1, and mounting in 20% Karo with phenol 

added as a preservative. Drawings were made 

using a Zeiss camera lucida. Aside from those 

listed below for DUKE and US, duplicates of 

some of the author's collection numbers have 

been deposited in AHFH, GALW, HBFH, 

MASS, MICH, NCU, NFLD, NHA, and WNC. 

Herbarium abbreviations follow Holmgren et al. 

(1981). Specimen measurements are accurate to 

0.1 micrometer. 

ACKNOWLEDGMENTS.—I thank Duke University 

for space and assistance at the Marine Laboratory 

and Dr. W. Kirby-Smith of the Reference Museum for sponsoring an offshore collecting trip 

and allowing access to the collections. I acknowledge the Duke University Cooperative Program 

in Oceanography as some of my earlier collections 

were made on R/ V Eastward cruises. I also thank 

fellow scientists (and the herbaria with which 

they are associated): Drs. R.B. Searles (DUKE), 

J.N. Norris (US), and D.F. Kapraun (WNC) for 

loans; Dr. L.G. Williams for sending specimens; 

and the Duke University Botany Department for 

access to that herbarium. Dr. M.H. Hommersand 

assisted with the literature and Dr. J.S. Ramus 

allowed use of his personal library. Ginny Schneider deserves special thanks for typing during my 

leave. Drs. Searles, Norris, and W.J. Woelkerling 

kindly reviewed the manuscript. This research 

was sponsored by Trinity College as a sabbatical 

leave. 

This monograph is dedicated to the memory of 

Dr. G.F. Papenfuss, both for his early work with 

the Audouinella complex and for his recent encouragement for me to unravel the taxa in North 

Carolina. 

Genus Audouinella 

Key to the Species of Audouinella from North Carolina1 

(Species alphabetically arranged in text) 

1. Plants attached to the substrate by an obvious, persistent, undivided basal 

spore which may give rise to one or several accessory cells, an endophytic 

1

 Several specimens from a population should be observed when utilizing the above key. 

Some characters, such as presence or absence of unicellular hairs, are variable and by themselves 

are not considered to be taxonomically significant. In practice, however, a combination of 

several characters, some apparently more stable than others, is useful in delineating taxa. This 

key reflects distinctions found in North Carolina specimens; it should also be helpful in 

distinguishing these "purported" taxa elsewhere. 
NUMBER 22 

branched filament, or a prostrate filamentous system 2 

Plants attached to the substrate by a multicellular prostrate system, the 

basal spore once septate and persistent, or not obvious, obscured by 

divisions in the earliest stages 8 

2. Plants minute, less than 250 jum tall A. microscopica 

Plants larger, greater than 500 jum tall 3 

3. Basal spores consistently globose or subglobose, obviously larger than erect 

filaments 4 

Basal spore ovoid to panduriform, globose or subglobose, the same size or 

only a few micrometers larger than erect filaments 7 

4. Basal spore producing a single penetrating endophytic branched filament 

in addition to one or more erect filaments 5 

Basal spore remaining undivided or occasionally producing a few accessory 

cells 6 

5. Basal spore 12-15 jum diam., erect filaments 8-13 jum diam.; dioecious, 

spermatangia forming corymbose clusters on lateral branches 

A. corymbifera 

Basal spore 7.5-12.5 /xm diam., erect filaments 3-5 jum diam.; monoecious, 

spermatangia paired, produced on opposite or whorled branches in a 

long series A. ophioglossa 

6. Monosporangia 5.0-7.5 jum diam., 11-15 jum long A. hoytii 

Monosporangia 10-18 jum diam., 18-27 jum long A. affinis 

7. Basal spore remaining undivided, ovoid, elongate to rectangular, occasionally with a flared disciform base A. hallandica 

Basal spore, pyriform to panduriform, producing a prostrate filamentous 

system A. dasyae 

8. Plants mostly endophytic or endozoic, upright axes less than 150 jum 

tall A. infestans 

Plants mostly epiphytic, epizoic, or saxicolous, upright axes greater than 

200 jum tall 9 

9. Plastids with more than one pyrenoid per cell A. botryocarpa 

Plastids with only one pyrenoid per cell 10 

10. Sporangia, for the most part, borne in clusters of 3 or more on branched 

stalks at the base of lateral branches A. daviesii 

Sporangia, for the most part, borne singly or in pairs, not clustered at the 

base of lateral branches 11 

11. Monosporangia 9-13 jum long 12 

Monosporangia 15-40 jum long 14 

12. Monosporangia 5-6 jum diam.; bisporangia, if present, 7-12 jum diam., 

12-18 jum long A. bispora 

Monosporangia 6-10 jum diam.; bisporangia not present 13 

13. Plants producing only single monosporangia, erect filaments to 1 mm tall 

arising from a uniseriate prostrate filament to small compact pseudoparenchymatous disc A. hypneae 

Plants producing single or 2-3 seriate monosporangia, erect filaments to 

0.5 mm tall arising from a wide-spreading, much-branched prostrate 

system, often with a persistent septate basal spore A. densa 
SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

14. Plastids stellate, parietal or axial, unicellular hairs common but not 

present on all plants, plants arising from an orbicular pseudoparenchymatous disc that is often later obscured by proliferating filaments ... . 

A. secundata 

Plastids lobate, parietal or axial, unicellular hairs absent or rare, but 

terminal cells occasionally taper sharply to the tips, plants not arising 

from an orbicular pseudoparenchymatous disc in early stages 15 

15. Prostrate system a widespreading mass of uniseriate filaments from an 

enlarged central pyriform to panduriform cell that is persistent but 

often obscured in older plants, uniseriate prostrate filaments giving rise 

to numerous erect filaments A. dasyae 

Prostrate system a compact, often penetrating mass of uniseriate filaments, 

usually coalesced centrally to form a pseudoparenchymatous disc, basal 

spore not persistent A. saviana 

Audouinella affinis (Howe and Hoyt), 

new combination 

FIGURE \a,b 

Acrochaetium affine Howe and Hoyt, 1916:118, pi. 15.—Hoyt, 

1920:471, pi. 69.—Searles and Schneider, 1978:100. 

Audouinella alariae sensu Kapraun, 1980:40, figs. 15-17. [Not 

(Jonsson) Woelkerling, 1973b:541, figs. 1-9] 

DESCRIPTION.—Plants epiphytic or epiendophytic, to 4 mm tall, caespitose, arising from a 

persistent, large, globose to subglobose spore, 14-

26 jum diam. including an obvious wall 2.5-5.0 

jum thick, one half again as big as the cells which 

emanate from it; if embedded in host tissue, the 

spore becoming vertically elongate to subpyriform or panduriform, if not, producing a subcylindric obtuse or truncate foot that penetrates to 

24 jum, the extended spore 20-33 jum high; spore 

usually remaining simple, occasionally developing a few smaller accessory cells, rarely producing 

short prostrate irregularly branched filaments up 

to 5 cells long; basal spore, and occasionally 

accessory cells or basal filament cells, producing 

1-4 erect filaments, cells cylindrical, 5-14 jum 

diam., 17-78 jum long, each with a parietal lobate 

plastid and one pyrenoid; for the most part plants 

immediately branched from the distal end of the 

first cell of the erect filament, branching subdichotomous or subtrichotomous, appearing rigid 

below, more flexuous above, branching mostly 

subdichotomous to distinctly lateral, ultimate 

branches elongate-virgate, 3.0-5.5 jum diam., 

gradually tapering toward the apices, often terminating with inconspicuous hairs. 

Monosporangia infrequent, lateral, secund, or 

less commonly terminal on upper portions of the 

plant, sessile or on one-celled pedicels, ovoid, 

obovoid, or oblong, 10-18 jum diam., 18-27 jum 

long, occasionally on gametophytic plants. 

Monoecious, spermatangia in proximity to sessile 

carpogonia, lateral or terminal, sessile or pedicellate, solitary or in groups of 2 or 3; carposporophytes common, 3-12 spored, carpospores 8-25 

jum diam., 13-28 jum long. 

TYPE-LOCALITY.—Onslow Bay, North Carolina, USA. 

HOLOTYPE.—US 56005. 

DISTRIBUTION.—Endemic to North Carolina, as 

far as currently known. 

NORTH CAROLINA SPECIMENS STUDIED.—Onslow Bay, reef, 11 Aug 1914, Radcliffe (Holotype, 

US 56005; Isotypes, US 33567, 33584, 33646; 

Isotypes, DUKE 111, 1458). Beaufort, Radio Is. 

jetty, 5 Jul 1960, Aziz (DUKE 109). Cape Lookout jetty, 23 Jul 1961, Aziz (DUKE 102, 108). 

Beaufort, Radio Is. jetty, 4 Aug 1961, Aziz 

(DUKE 103-107, 110, 120, 121). Onslow Bay, 

reef, 2 Aug 1971, Searles (DUKE 13190). Onslow 

Bay, reef, 20 Jul 1973, Schneider 718 (DUKE 

12606, 12716; US 50776; CWS). Onslow Bay, 

reef, 11 Aug 1981, Schneider 2711 (DUKE 16525; 

US 88906; CWS). Beaufort, Radio Is. jetty, 20 
NUMBER 22 

Jul 1982, Schneider 2867 (CWS). 

REMARKS.—This species is an obvious epiphyte 

of Dictyota, Sargassum, and other algae from deep 

(to 30 m) offshore and from shallow subtidal 

habitats in the Beaufort and Wilmington areas, 

at times turning the host alga red with a complete 

or marginal covering of its fine red filaments. To 

date, found from June to August. 

Hoyt (1920) listed several morphological characters useful in distinguishing Audouinella affinis 

from the earlier described A. hoytii. Although they 

are generally helpful, I find the size of monosporangia as the most reliable in the separation of 

these two closely allied taxa. A. affinis appears 

related to A. alariae (Jonsson) Woelkerling 

(1973b), but can be distinguished by the production of accessory cells from the basal spore and 

the larger monosporangia. A comparison of type 

material is needed to clarify the limits of species 

with persistent globose basal spores. 

Audouinella bispora (B0rgesen) Garbary 

FIGURE Ic-f 

Audouinella bispora (B0rgesen) Garbary, 1979:490. 

Chantransia bispora B0rgesen, 1910:178, fig. 1. 

DESCRIPTION.—Plants epiendophytic to 1.5 mm 

tall, arising from an irregularly ramified, spreading prostrate system that remains superficial or 

penetrates host tissue, prostrate cells cylindrical 

to irregular, to 17.5 jum in greatest dimension, all 

capable of producing an erect filament; erect 

filaments radially to irregularly branched and 

rebranched, infrequent below, more above, the 

branches sharply to narrowly angled from the 

axes that bear them, tapering toward the apices; 

cells cylindrical, 2.5-8.0 jum diam., 10.0-32.5 jum 

long in the main axes, 1.0-3.0 jum diam., in the 

ultimate segments, occasionally ending in long 

multicellular, lightly pigmented hair-like extensions, unicellular hairs unknown; each cell containing a well-developed parietal plastid with a 

single large pyrenoid. 

Monosporangia terminal or lateral on short 

branches to adaxial on longer axes, sessile or on 

one-celled pedicels, single or paired, narrow, 

ovoid to oblong, 5-6 jum diam., 9.5-12.5 jum long; 

bisporangia situated as monosporangia, single or 

paired, broad, ovoid to ellipsoidal, 7.5-11.5 jum 

diam., 12.5-17.5 jum long. Gametangia unknown. 

TYPE-LOCALITY.—St. Thomas, Virgin Islands. 

HOLOTYPE.—C. 

DISTRIBUTION.—North Carolina, St. Thomas 

(Virgin Islands). 

NORTH CAROLINA SPECIMENS STUDIED.—Onslow Bay, reef, 30 June 1972, Searles (DUKE 

16526). Onslow Bay, reef, 5 May 1981, Handsel, 

BLM 65818086 (Duke Marine Lab Reference 

Museum). 

REMARKS.—Collected to date from water 25-

30 m deep in Onslow Bay, May-June. This is the 

first report of this taxon outside the type-locality. 

The North Carolina specimens perfectly fit the 

descriptions of B0rgesen (1910, 1915), and I do 

not hesitate to place them under Audouinella bispora. The questions raised by B0rgesen (1915) as 

to whether this plant is indeed a form of Audouinella hypneae (B0rgesen) Schneider cannot be ascertained without additional comparative material of the latter named species (see discussion of 

A. hypneae). Audouinella bispora occasionally produces bispores and has narrower monospores than 

reported for A. hypneae, a feature also seen in 

North Carolina plants. Nevertheless, it is interesting to note that in Garbary's (1979) cluster analysis, A. bispora was found to be quite similar to A. 

seriata (B0rgesen) Garbary, a species I consider a 

synonym of A. hypneae in this paper. Audouinella 

hypneae was not treated by Garbary (1979). 

The description of Audouinella bispora by B0rgesen is broadened here based on the variation 

seen within North Carolina plants, but no characteristics were found that contradict the earlier 

descriptions. 

Audouinella botryocarpa (Harvey) Woelkerling 

FIGURES 1^-/ 

Audouinella botryocarpa (Harvey) Woelkerling, 1971:37.— 

Searles and Schneider, 1978:100. 

Callithamnion botryocarpum Harvey, 1854:563. 

DESCRIPTION.—Plants epiphytic, epiendophytic 
SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

or saxicolous to 1 cm tall, caespitose, arising from 

an irregularly branched prostrate system with 

entangled filaments free from one another or 

coalesced into a pseudoparenchymatous disc, occasionally supplemented by corticating rhizoids 

descending cells or lower portions of erect filaments; base giving rise to several erect filaments, 

much and irregularly to unilaterally branched, 

sometimes terminating abruptly in multicellular 

hair-like extensions or gradually tapering toward 

the tips; cells of main axes cylindrical (10—) 15— 

20(-30) jum diam., 30-120 jum long and 6-15 jum 

diam. in the ultimate segments, each cell containing a single parietal lobate plastid with one to 

several pyrenoids, unicellular hairs unknown. 

Monosporangia terminal on short branches 

and adaxial, near the bases of branches, secund 

or scattered, sessile or on one-celled pedicels, single, paired, or rarely in groups of 3 to 5, ovoid, 

12-28 jum diam., 18-43 jum long; tetrasporangia 

cruciate, with range of dimensions, shape, and 

position similar to monosporangia, in general 

somewhat larger than monosporangia on the 

same plant. Sexual plants unknown in the Carolinas. 

TYPE-LOCALITY.—King George's Sound, western Australia. 

HOLOTYPE.—Harvey 324, travelling collection; 

not in TCD. 

DISTRIBUTION.—North Carolina, Australia, 

New Zealand. 

NORTH CAROLINA SPECIMENS STUDIED.—Beaufort, Radio Is. jetty, 2 Mar 1974, Schneider 829 

(DUKE 13250, 13251, 13264; CWS). Beaufort, 

Radio Is. jetty, 16 Feb 1975, Searles 75-1-1 

(DUKE 14178; CWS). Beaufort, Radio Is. jetty, 

14 Sep 1981, Schneider 2720 (DUKE 16524; CWS). 

Beaufort, Pivers Is. seawall, 10 Nov 1981, Schneider 

2746 (DUKE 16590; US 88908; CWS). Beaufort; 

Pivers Is. seawall, 7-8 Dec 1981, Schneider 2753 

(DUKE; US 88909; CWS). Beaufort, Radio Is. 

jetty, 20 Jul 1982, Schneider 2864 (DUKE; US 

88910; CWS). 

REMARKS.—Known only as an obvious epiphyte of Codium from the Pivers Island seawall 

and Radio Island jetty, Beaufort, but probably 

more widespread. Collected July-March. 

This taxon is at present known in the Atlantic 

Ocean only from North Carolina and also represents the only member of the acrochaetoid complex with more than one pyrenoid per cell (Figure 

1^) from the east coast of North America (Searles 

and Schneider, 1978). Specimens have been compared with type material by W.J. Woelkerling. 

The only plants thus far collected in North 

Carolina are tetrasporic and/or monosporangiate. Monosporangia, found in all collections, 

are large, falling in the upper size ranges of those 

listed by Woelkerling (1970, 1971). Many of the 

monosporangia are terminal on short branches, 

some later becoming subtended by the short lateral filaments that produce them, becoming displaced to a lateral position. Later the monosporangia are seen as clustered near the base of 

branches, presumably due to the subtended filaments forming beneath monosporangia. Tetrasporangia have been found only in December 

collections. Plants from New Zealand and Australia are no longer than 6 mm, while those in 

the Carolina region reach 1 cm, by far the largest 

of the acrochaetoids from this area. Multicellular 

hair-like extensions have thus far been observed 

only from the September Radio Island populations; none of these minor differences warrants a 

separation from the taxon previously known only 

from the Southern Hemisphere, Audouinella 

botryocarpa. 

Audouinella corymbifera (Thuret) Dixon 

Audouinella corymbifera (Thuret in Le Jolis) Dixon in Parke 

and Dixon, 1976:590. 

Chantransia corymbifera Thuret in Le Jolis, 1863:107 [in part]. 

Acrochaetium corymbiferum (Thuret in Le Jolis) Batters, 

1902:59.—Hoyt, 1920:473. 

Acrochaetium bornetii Papenfuss, 1945:313.—Searles and 

Schneider, 1978:100. 

DESCRIPTION.—Plants epiendophytic, 2-3 mm 

tall, arising from a persistent, large, globose spore, 

12-15 jum diam., which produces an irregular, 

contorted branched filament that deeply penetrates host tissue with cells 8-13 jum diam., 25-40 

jum long; basal spore and occasional cells of the 

internal filament giving rise to erect branched 

axes, secund to alternate with branches more 
NUMBER 22 

abundant above and not often rebranched; cells 

of erect filaments cylindrical, 7-16 jum diam., 22-

55 jum long, each cell containing a well-developed 

parietal lobed plastid with a single pyrenoid, 

unicellular hairs unknown. 

Monosporangia terminal on short branches or 

lateral, near the bases of branches, sessile or on 

one-celled pedicels, single, ovoid, 8-10 jum diam., 

15-18 jum long, occasionally associated with gametangia. Dioecious, spermatangia lateral on 

main axes, formed in a corymbose cluster on a 1-

2 celled pedicel, colorless, 4-5 jum diam.; carpogonia lateral on a main axis, on a 1-2 celled 

pedicel near the bases of branches, producing a 

corymbose carposporophyte with terminal carposporangia, ovoid, 9-15 jum diam., 14-18 jum 

long. 

TYPE-LOCALITY.—Belle-Ile-en-Mer, Atlantic 

France. 

HOLOTYPE.—Cannot be located (Dixon and 

Irvine, 1977:88). 

DISTRIBUTION.—North Carolina, Bermuda, 

Mediterranean, England, France. 

NORTH CAROLINA SPECIMENS STUDIED.—Beaufort harbor, 20 May 1907, Hoyt (DUKE 135; US 

33648). 

REMARKS.—Abundant sexually reproductive 

plants found on one specimen of Dasya from 

Beaufort harbor, to date collected only once, in 

May. 

For a clarification of the correct epithet refer 

to Dixon and Irvine (1977). 

Audouinella dasyae (Collins) Woelkerling 

FIGURE \m-p 

Audouinella dasyae (Collins) Woelkerling, 1973b:545, figs. 10-

31.—Searles and Schneider, 1978:100.—Kapraun, 

1980:40 [in part]. 

Acrochaetium dasyae Collins, 1906:191.—Aziz, 1967:408. 

Acrochaetium robustum sensu Williams, 1948:690; 1951:153.— 

Taylor, 1960:315 [in part]. [?Not B0rgesen, 1915:40, figs. 

38-40.] 

DESCRIPTION.—Plants epiphytic, to 3 mm tall, 

caespitose, arising from a basal spore and widespreading uniseriate, prostrate filamentous axis it 

produces; original spore elongated pyriform to 

panduriform, not greater in size than the cells 

which it produces, rarely remaining simple, often 

becoming obscured by descending rhizoids and 

entangled prostrate filamentous axes; creeping 

prostrate system giving rise to several erect filamentous, moderate to much and irregularly 

branched, often tapering to the tips, cells cylindrical, 6-12(-16) jum diam., 25-70(-90) jum long 

in lower portions, 6-10 jum diam., 15-60 jum long 

in the ultimate segments, each having a single 

parietal plastid with one pyrenoid, hairs unknown. 

Monosporangia mostly single, occasionally in 

pairs, secund to scattered, sessile or on one-celled 

pedicels, ovoid, 7—12(—16) jum diam., 16-27 jum 

long, borne on vegetative or gamete-producing 

plants. Monoecious or dioecious, spermatangia 

terminal or lateral on specialized short lateral 

branched axes, globose to ovoid, 2-4 jum diam., 

3-5 jum long; carpogonia scattered, sessile or 

stalked; carposporophytes with numerous terminal and lateral ovoid carposporangia, 9-12 jum 

diam., 16-24 jum long. 

TYPE-LOCALITY.— Woods Hole, Massachusetts, USA. 

HOLOTYPE.—FH. 

DISTRIBUTION.—Florida to Prince Edward Island (discontinuous; not reported from South 

Carolina and Georgia). 

NORTH CAROLINA SPECIMENS STUDIED.—Cape 

Lookout jetty, 16 Jun 1947, Williams (CWS). 

Cape Lookout jetty, 14 Aug 1962, Aziz (DUKE 

125). Beaufort, Pivers Is., 6 Apr 1963, Aziz 

(DUKE 158). Onslow Bay, reef, 16 Jul 1968, 

Searles (CWS). Beaufort, Radio Is. jetty, 8 May 

1971, Schneider 172a (CWS). Onslow Bay, reef, 11 

Aug 1981, Schneider 2687 (DUKE 16523; CWS). 

REMARKS.—Epiphytic on Dasya, Sargassum, and 

other hosts from shallow and deep subtidal environments, April-August in the area. 

After an examination of Williams's collections 

of Acrochaetium robustum, I find those specimens 

easily fall within the circumscription of Audouinella 

dasyae (Woelkerling, 1973b); any slight variation 

in their holdfasts is probably due to the differences in surface textures of the host plants. An 

examination of Collins type material of A. robus-
8 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

turn is necessary to test whether the two taxa are 

indeed conspecific. Stegenga and Borsje (1976) 

have found that A. dasyae alternates with a heteromorphic tetrasporophyte provisionally identified as A. saviana; however, they felt it premature 

to deal with the nomenclatural problems arising 

from their study. 

North Carolina plants are often found with an 

early formed basal system of the ovoid to pyriform 

spore and one smaller accessory cell from which 

an upright filament rises obliquely (Figure \m,n). 

Audouinella daviesii (Dillwyn) Woelkerling 

FIGURE 2a-c 

Audouinella daviesii (Dillwyn) Woelkerling, 1971:28, figs. 7, 

22. 

Conferva daviesii Dillwyn, 1809:73, suppl., pi. F. 

Audouinella hallandica sensu Kapraun, 1980:41, fig. 26 [in 

part]. [Not Audouinella hallandica (Kylin) Woelkerling, 

1973a:82.] 

DESCRIPTION.—Plants epiphytic, epiendophytic, epizoic, or epiendozoic, to 6 mm tall, caespitose, arising from spreading to penetrating 

branched filamentous systems, centrally coalesced 

into irregular pseudoparenchymatous discs or entangled fungiform masses, original spores not persistent; erect filaments sparsely to freely and irregularly to unilaterally branched, branches often 

ending in long multicellular hair-like projections, 

3-4 jum diam., 60-120 jum long; cells of main axes 

cylindrical (6-)9-12(-20) jum diam., (8-) 15-50 

(-70) jum long, and only slightly tapering to tips, 

which are 7.5-10.0 jum diam. where extensions 

not formed, each cell containing a single parietal 

lobate plastid with one pyrenoid; unicellular hairs 

unknown. 

Monosporangia terminally clustered in groups 

of 3 or more on branched stalks or paired on 1-2 

celled stalks adaxially on the lowermost cells of 

lateral branches, rarely single and scattered, 

ovoid, 7-13 jum diam., 8-20 jum long; tetrasporangia cruciate, paired on unicellular pedicels, 

solitary or in groups of 3, in similar positions to 

monosporangia, 13-22 jum diam., 12-36 jum long. 

Sexual plants unknown in the Carolinas. 

TYPE-LOCALITY.—Bantry Bay, Ireland. 

HOLOTYPE.—NMW. 

DISTRIBUTION.— Nearly cosmopolitan. As yet, 

not known from South Carolina and Georgia. 

NORTH CAROLINA SPECIMENS STUDIED.— 

Wrightsville Beach, Masonboro Inlet jetty, 18 Jan 

1973, Kapraun (WNC). Wrightsville Beach, Masonboro Inlet jetty, Apr 1977, 6 Apr 1978, Kapraun 

(WNC; DUKE). Ft. Macon jetty, 30 Ju n 1982, 

Schneider 2831 (DUKE; US 88911; CWS). 

REMARKS.—To date, known only from the shallow subtidal of Masonboro Inlet jetty, Wrightsville Beach, January and April, and the intertidal 

of Fort Macon jetty, June. 

This is the first report of this taxon from North 

Carolina, although commonly known to the north 

(Woelkerling, 1973b). Only sporangiate plants 

have been found to date and these agree in all 

respects to recently published descriptions 

(Woelkerling, 1971, 1973b; Dixon and Irvine, 

1977). Multicellular hair-like projections were 

found only in the January collection and tetrasporangia only in April. 

Audouinella densa (Drew) Garbary 

FIGURE 2d-i 

Audouinella densa (Drew) Garbary, 1979:490. 

Rhodochorton densum Drew, 1928:168, pi. 38: figs. 17-24. 

Acrochaetium densum (Drew) Papenfuss, 1945:308.—Stegenga 

and Vroman, 1976:276, figs. 1-9. 

Acrochaetium thuretii sensu Kapraun, 1980:41, fig. 27. [Not 

Acrochaetium thuretii (Bornet) Collins and Hervey, 1917: 

98.] 

Acrochaetium hummii Aziz, 1965:59, pi. 8: figs. 1-6, pi. 9: figs. 

1-4, pi. 10: fig. 4 [nomen nudum]. 

DESCRIPTION.—Plants epiphytic or epizoic, 0.2-

0.3(-0.5) mm tall, caespitose, arising from an 

irregularly branched prostrate system, often originally developed from a persistent septate spore, 

occasionally coalesced to form a pseudoparenchymatous disc, especially centrally, giving rise to 

several erect filaments, sparsely to much 

branched, irregular, alternate, opposite and secund, usually with a moderate branching above, 

the branches sometimes tapering toward the tips, 

often ending in colorless unicellular hairs; cells of 

the upright filaments cylindrical to clavate, 4-8 
NUMBER 22 

(-13) jum diam., 7-18 jum long, each containing 

a single lobate to stellate plastid with a single 

pyrenoid. 

Monosporangia single or 2-3 seriate, terminal, 

alternate secund, rarely opposite, sessile or on 

one-celled pedicels, single or in pairs, rarely clustered; monosporangia ovoid to ovoid-ellipsoid, 7-

10 jum diam., 10-13 jum long. 

TYPE-LOCALITY.—Fort Point, San Francisco, 

California, USA. 

HOLOTYPE.—UC 294560 (Gardner 4607). 

DISTRIBUTION.—Atlantic: North Carolina, 

South Carolina, Brazil, Netherlands. Pacific: 

Mexico, California, Japan, Kurile Islands. 

NORTH CAROLINA SPECIMENS STUDIES.—Ft. Macon jetty, 27 Jun 1960, Aziz (DUKE 112). Beaufort, Radio Is. jetty, 15 Apr 1973, Kapraun (NCW). 

Wrightsville Beach, Masonboro Inlet jetty, 26 

Feb 1978, 20 May 1978, 10 Sep 1978, Kapraun 

(NCW). Ft. Macon jetty, 1 Jul 1978, Schneider 78-

12-2 (DUKE; US 88913; CWS). Wrightsville 

Beach, Masonboro Inlet jetty, 4 Apr 81, Searles 

(DUKE, slide). Beaufort, Pivers Is., 7-8 Dec 1981, 

Schneider 2760 (CWS). Beaufort, Radio Is. jetty, 

21 Jun 1982, Schneider 2790 (DUKE; US 88915; 

CWS). Ft. Macon jetty, 30 Jun 1982, Schneider 

2826 (DUKE; US 88912; CWS). 

REMARKS.—Epiphytic on various algae and animals, in particular Chaetomorpha aerea (Dillwyn) 

Kutzing, throughout the year from the intertidal 

and shallow subtidal of Cape Lookout jetty, Fort 

Macon jetty and the Pivers Island seawall, Beaufort and Masonboro Inlet jetty at Wrightsville 

Beach. Also a single collection from 4.5 m in Port 

Royal Sound, South Carolina. 

Germinating spores divide into two equal cells 

after attaching to a host (Figure 2d,e). One or 

both of these initials gives rise to an erect and/or 

prostrate filament, occasionally one initial remaining undivided for some time. The prostrate 

system consists of either irregular filamentous 

branched axes or becomes compacted into a pseudoparenchymatous disc (see Figure 2/). 

This taxon was first reported from North Carolina in an unpublished thesis as "Acrochaetium 

hummii" (Aziz, 1965). Stegenga and Vroman 

(1976) recognized the Aziz plants as members of 

a widespread heteromorphic acrochaetoid with 

seriate monosporangia tentatively identified by 

them as Acrochaetium densum (Drew) Papenfuss (= 

Audouinella densa [Drew] Garbary). In their report, 

Stegenga and Vroman (1976) call the report of 

gametangia in A. hummii by Aziz (1965) 

"strange." I could not find any sexual structures 

on Aziz's specimens (slides); therefore, to date 

only monosporangiate plants are known from 

here. This is further evidence that the North 

Carolina specimens belong in the A. densa complex outlined by Stegenga and Vroman (1976). 

They suggest A. microscopica as a possible gametophytic generation if seriate monosporangia are 

not considered taxonomically important at the 

species level. Additional study is needed to clarify 

this relationship. For the present, I consider seriate monosporangia taxonomically significant, 

and leave the plants under the oldest validly 

published name, A. densa. 

Audouinella hallandica (Kylin) Woelkerling 

Audouinella hallandica (Kylin) Woelkerling, 1973a:82, figs. 1-

4.—Schneider, 1976:138.—Searles and Schneider, 1978: 

100.—Kapraun, 1980:41, figs. 18-20 [in part]. 

Chantransia hallandica Kylin, 1906:123, fig. 8. 

Acrochaetium dufourii (Collins) B0rgesen, 1915:19.—Hoyt, 

1920:470, fig. 26.—Taylor, 1960:305. 

Acrohaetium sargassi Bdrgesen, 1915:17, figs. 7-10.—Taylor, 

1960:306. 

DESCRIPTION.—Plants epiphytic, to 1 mm tall, 

arising from an ovoid, obovate subglobose, disciform, elongated to rectangular basal spore; if 

disciform, the flared base of the spore to 20 jum, 

otherwise 5-8 jum diam. and the same size or 

slightly larger than cells of the filaments it produces; spore remaining simple producing 1-2(-3) 

erect filaments, branching irregularly lateral to 

secund, not closely set, beginning in the lowermost portions of the plant, occasionally tapering 

toward the apices, hairs unknown; lower cells 

cylindrical, 3-7 jum diam., 10-30 jum long, each 

containing a single parietal lobate plastid with 

one pyrenoid. 

Monosporangia single or paired, lateral to se-
10 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

cund, more rarely terminal, sessile or on onecelled pedicels, obovate to ovoid, 5-10 /xm diam., 

8-11 (-16) jum long. Dioecious, spermatangia opposite on short lateral branches, rare; carpogonia 

sessile, rare. 

TYPE-LOCALITY.—Hogardsgrund, Halland, 

Sweden. 

HOLOTYPE.—LD (See Woelkerling, 1973a:84). 

DISTRIBUTION.—North Carolina, South Carolina, Bermuda, Sargasso Sea, Florida to Venezuela, Brazil, Europe. 

NORTH CAROLINA SPECIMENS STUDIED (representative).—Ft. Macon jetty, 13 Jul 1908, Hoyt, 

Acrochaetium dufourii (Collins) Hoyt (Isotype, 

DUKE 136). Beaufort, Pivers Is., 25 Jun 1960, 

Aziz (DUKE 113, 142). Beaufort, Shackleford 

jetty, 29 Jul 1961, Aziz (DUKE 143). Ft. Macon 

jetty, 5 Aug 1961, Aziz (DUKE 154-157). Beaufort, Shackleford jetty, 6 Aug 1961, Aziz (DUKE 

144, 149-153). Beaufort, Shackleford jetty, 12 

Aug 1962, Aziz (DUKE 145-147). Cape Lookout 

jetty, 8 Ju n 1975, Schneider 947 (DUKE 15077, 

15133; CWS). Onslow Bay, wreck, 9 Jun 1975, 

Schneider 998a (CWS). Wrightsville Beach, Masonboro Inlet jetty, 11 Nov 1977, Kapraun (WNC). 

Onslow Bay, reef, 11 Aug 1981, Schneider 2683 

(CWS). 

REMARKS.—Known as an epiphyte of Dictyota 

and Sargassum from shallow subtidal habitats near 

Beaufort and Wilmington, and on a variety of 

red and brown algae from deep water (to 35 m) 

environs, June-November. 

This taxon is most easily recognized when attached to its host by the distinctive broad disc of 

cell-wall materials. The disc does not stain and 

the original spore remains as a simple elongated 

basal cell the same size or slightly larger than the 

cells of erect filaments it produces. 

A recent study by Stegenga and Borsje (1977) 

demonstrates that a plant provisionally determined as Audouinella hallandica is the gametophyte 

of a heteromorphic plant whose sporophyte is 

tentatively determined as Acrochaetium polyblastum 

(Rosenvinge) B0rgesen. No formal taxonomic 

changes for this species complex were offered, 

however. 

Audouinella hoytii (Collins), new combinaton 

FIGURE 2j 

Acrohaetium hoytii Collins, 1908:134.—Hoyt, 1920:470, figs. 

27, 28.—Williams, 1948:690; 1951:153.—Searles and 

Schneider, 1978:100.—Wiseman, 1978:29. 

DESCRIPTION.—Plants epiphytic, 0.2-1.3 mm 

tall, arising from a persistent, large, globose spore, 

9-15 (-28) jum diam. including an obvious wall 2-

3 jum thick; spore superficial to slightly embedded 

in host tissue, elongating to 30 jum, being obviously broader than the cells it produces, remaining simple, rarely forming one or a few 

accessory cells, producing 1—2(—4) erect filaments; 

erect branching infrequent to frequent, often secund, ultimate branches elongated, tapering to 

the apices for the most part, 2-3 jum diam., hairs 

unknown; cells of the main axes cylindrical, 5-7 

jum diam., 10-20 jum long, each with a single 

parietal plastid and a large central pyrenoid. 

Monosporangia secund or lateral on upper portions of the plant, sessile or on one-celled pedicels, 

oblong, 5.0-7.5 jum diam., 11-15 jum long. Monoecious, spermatangia lateral and terminal on 

short lateral branching systems; carpogonia sessile, lateral; carposporophytes developing as short 

branching systems lateral on main axes or near 

the base of branches, with 4-16 terminal carpospores, 4-9 jum diam., 7.0-13.5 jum long. 

TYPE-LOCALITY.—Fort Macon jetty, vicinity 

Beaufort, North Carolina, USA. 

HOLOTYPE.—US 33650. 

DISTRIBUTION.—North Carolina, South Carolina. 

NORTH CAROLINA SPECIMENS STUDIED.—Ft. 

Macon jetty, 20 Sep 1905, Hoyt (US 43499). Ft. 

Macon jetty, 12 Oct 1905, Hoyt (Holotype, US 

33650). Ft. Macon jetty, 15 Aug 1906, Hoyt 

(DUKE 137; US 43500a, 33651). Beaufort harbor, 5 Oct 1906, Hoyt (DUKE 1460; US 43500b). 

Ft. Macon jetty, 13 Jul 1908, Hoyt (DUKE 1461; 

US 33652, 2803 as P.B.-A. 1540). Beaufort, Shackleford jetty, 14 Jul 1938, Blomquist (DUKE 133). 

Cape Lookout jetty, 28 Aug 1946, Williams 

(CWS). Ft. Fisher outcropping, July 1955, Rho 
NUMBER 22 11 

(DUKE 139). Beaufort, Shackleford jetty, 4 Jul 

1960, Aziz (DUKE 119, 122, 123). Beaufort, Radio Is. jetty, 5 Jul 1960, Aziz (DUKE 148). Beaufort, Pivers Is., 18 Jul 1974, Searles (DUKE 16091). 

Beaufort, Radio Is. jetty, 13 Jun 1978, Schneider 

78-4-5b (CWS). Beaufort, Duncan breakwater, 27 

J u n 1978, Schneider 78-10-20 (CWS). Beaufort, 

Radio Is. jetty, 20 Sep 1981, Schneider 2738 

(DUKE 16522; US 88914; CWS). Beaufort, Radio Is. jetty, 21 Ju n 1982, Schneider 2789 (DUKE; 

US 88915; CWS). Beaufort, Radio Is. jetty, 20 

Jul 1982, Schneider 2861 (DUKE; US 88916; 

CWS). 

REMARKS.— Known as an epiphyte of Dictyota, 

Padina, and other seaweeds from shallow subtidal 

habitats in the Beaufort area, June-October. 

Although similar in morphology to Audouinella 

alariae (Jonsson) Woelkerling (1973b) from New 

England and Europe, main axes, carposporangia, 

and monosporangia of A. hoytii are one-half the 

size of the first mentioned taxon. A further comparison is necessary to clarify the relationship of 

these two species. Audouinella hoytii has also been 

linked to A. pectinata (Kylin) Papenfuss by Woelkerling (1971). 

Audouinella hypneae (B0rgesen), new 

combination 

FIGURE 2k-l 

Chantransia hypneae B0rgesen, 1909:2, fig. 2. 

Acrochaetium hypneae (B0rgesen) B0rgesen, 1915:51, figs. 53, 

54. 

Acrochaetium seriatum B0rgesen, 1915:32, figs. 25-28.—Williams, 1948:690—Searles and Schneider, 1978:100. 

DESCRIPTION.—Plants epiphytic or epiendophytic to 1 mm tall arising from small compact 

to spreading filamentous prostrate system composed of one or more irregularly bent, branched 

filaments that can be coalesced centrally into a 

pseudoparenchymatous disc; each cell of the disc 

or ramified filaments capable of producing an 

erect filament; erect branches little to radially 

and much branched below, alternate, often pectinate secund above, cells cylindrical, 6-10 jum 

diam., 16-22 jum long below, tapering to 4-7 jum 

diam., 16-30 jum long in the ultimate segments, 

each containing a single parietal plastid with an 

obvious pyrenoid, occasionally bearing unicellular hyaline hairs to 60 jum long. 

Monosporangia single, rarely paired, lateral in 

long secund series or rarely terminal, occasionally 

scattered, sessile or on one-celled pedicels, oblongovoid, 6-10 jum diam., 9-13 jum long. Gametangia 

unknown. 

TYPE-LOCALITY.—St. Thomas, Virgin Islands. 

HOLOTYPE.—C. 

DISTRIBUTION.—North Carolina, South Carolina, Georgia, Bermuda, Florida, Virgin Islands, 

Brazil, East Indies. 

NORTH CAROLINA SPECIMENS STUDIES.—Ft. Macon jetty, 15 Jul 1907, Hoyt (US 42553). Cape 

Lookout jetty, 28 Sep 1946, Williams (CWS). 

Beaufort, Shackleford jetty, 4 Jul 1960, Aziz 

(DUKE 124). Cape Lookout jetty, 8 Aug 1961, 

Aziz (DUKE 186-187). Onslow Bay, reef, 11 Jul 

1976, Searles (CWS). Onslow Bay, reef, 15 Jul 

1981, Handsel, BLM 65818144, 65818145, 

65818148 (Duke Marine Lab Reference Museum). Onslow Bay, reef, 28 Jul 1981, Amsler, 

BLM 65818152 (Duke Marine Lab Reference 

Museum). Onslow Bay, reef, 11 Aug 1981, Schneider 2712 (DUKE 16523; CWS). Cape Lookout 

jetty, 5 Jul 1982, Schneider 2841 (DUKE; US 

88917; CWS). 

REMARKS.—On a variety of plant and animal 

hosts from Beaufort area jetties and on Arthrocladia, Spatoglossum, and other hosts from deep offshore, July-September. 

North Carolina plants show variation in their 

attachment, ranging from the small compact, 

epiphytic disc illustrated for Acrochaetium seriatum 

(B0rgesen, 1915:32) to the short branched or 

simple prostrate epiphytic (A. gracile, B0rgesen, 

1915:26) or endophytic (A. hypneae, B0rgesen, 

1915:52) filament. In his discussion of A. hypneae, 

B0rgesen (1915:52) states, "If the basal part is left 

out of consideration it cannot be denied that the 

above, [newly] described Acrochaetium seriatum 

shows great likeness to this species. But while the 

present plant has an endophytic base the other is 

epiphytic." I have observed epiphytic, epi-endophytic and endophytic bases of this small spored 
12 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

acrochaetoid on the same host, Arthrocladia, from 

30 m offshore. Basal system variation has previously been used as a taxonomic criterion, even 

though this variation may have been due to the 

texture of the host. Recent studies have shown 

great variation within the attachment morphology for Audouinella daviesii (Dillwyn) Woelkerling, 

and other species (Woelkerling 1971, 1973b). I 

therefore propose that plants previously reported 

from North Carolina as Acrochaetium seriatum B0rgesen by Williams (1948) are the same as those I 

have observed. 

Because the single criterion that separates them 

appears unreliable, Acrochaetium seriatum is synonymized under the earliest available taxon, Audouinella hypneae. A recent report of somewhat 

larger spore dimensions for A. seriatum than shown 

in this and classical reports requires additional 

study. If the plants reported by Hall and Eiseman 

(1981) conform to B0rgesen's A. seriatum, then 

spore size distinction between A. hypneae and A. 

saviana is clouded, and further taxonomic reductions may be necessary. Until additional studies 

are completed, the key and description of A. 

hypneae in this report conform to Carolina specimens and classical reports (e.g., B0rgesen, 1915; 

Taylor, 1960). 

It is, at this stage, still possible to segregate 

Audouinella bispora from A. hypneae on the basis of 

size and type of sporangia. I expect however, as 

did B0rgesen, that these two taxa are one and 

that further collections and life history studies 

will bear this out. The necessity of a taxonomic 

study of the many B0rgesen Caribbean acrochaetoid species is obvious to me. Several of the small 

spored, partially or wholly endophytic or epiphytic plants will no doubt prove to be conspecific. 

At present, I consider the evidence sufficient to 

synonymize two of the species in this complex. 

Audouinella infestans (Howe and Hoyt) Dixon 

FIGURE 2m 

Audouinella infestans (Howe and Hoyt) Dixon in Parke and 

Dixon, 1976:590. 

Acrochaetium infestans Howe and Hoyt, 1916:116, pi. 14.— 

Hoyt, 1920:473, pi. 68. 

Kylinia infestans (Howe and Hoyt) Papenfuss, 1947:438.— 

Taylor, 1960:301. 

Colaconema infestans (Howe and Hoyt) Woelkerling, 1973a:89, 

figs. 5, 6.—Searles and Schneider, 1978:100. 

DESCRIPTION.—Plants endophytic or endozoic 

with emergent axes to 100 jum tall exclusive of 

hairs, internal axes consisting of tortuous, serpentine, or labyrinthine irregularly branched filaments, sparse and straight for considerable distances, often crowded between host cells or compacted and subparenchymatous, cells irregular to 

elongate, (2-)4-8(-13) jum diam., 6-60 jum long; 

emergent axes to 10 or more cells high but generally less, simple or sparingly and irregularly 

branched, occasionally bearing terminal hairs to 

170 jum long; cells cylindrical to ellipsoid, 3-7 jum 

diam., 6-30 jum long, containing a single parietal 

lobate plastid with a single pyrenoid. 

Monosporangia single or paired, terminal or 

lateral, often on the adaxial surface, sessile or on 

one celled pedicels, ovoid to ellipsoid, 4-6(-9) jum 

diam., 6-15 jum long. Gametangia unknown. 

TYPE-LOCALITY.—Onslow Bay, North Carolina, USA. 

HOLOTYPE.—US 56009. 

DISTRIBUTION.—North Carolina, Bermuda, 

Sargasso Sea, British Isles, Mediterranean, Japan. 

NORTH CAROLINA SPECIMENS STUDIES.—Onslow Bay, reef, 11 Aug 1914, Radclijfe, (Holotype, 

US 56009, Isotypes, US 33514, 33653, 336534, 

DUKE 140). Cape Lookout jetty, 8 Aug 1961, 

Aziz (DUKE, slide). Onslow Bay, wreck, 8 Jul 

1976, Searles (CWS). Beaufort, Radio Is. jetty, 13 

Jun 1978, Schneider 78-4-5a (CWS). Onslow Bay, 

reef, 15 Jul 1981, Searles BLM 65818144 (Duke 

Marine Lab Reference Museum). Onslow Bay, 

reef, 28 Jul 1981, Amsler BLM 65818152 (Duke 

Marine Lab Reference Museum). 

REMARKS.—Collections on various hydroids, 

Dictyota, and Sargassum from deep offshore habitats and shallow subtidal habitats near Beaufort 

and Wilmington, June-August. 

Audouinella microscopica (Nageli) 

Woelkerling 

Audouinella microscopica (Nageli) Woelkerling, 1971:33, figs. 

10, 23A.—Searles and Schneider, 1978:100.—Kapraun, 

1980:41, figs. 21, 22. 

Callithamnion microscopicum Nageli in Kutzing, 1849:640. 
NUMBER 22 13 

Acrochaetium trifilum (Buffham) Batters, 1902:58. 

Acrochaetium parvulum (Kylin) Hoyt, 1920:470, fig. 25. 

Acrochaetium compactum Jao, 1936:241, pi. 10: figs. 6-14.— 

Williams, 1948:690. 

Kylinia crassipes (B0rgesen) Kylin, 1944:13.—Taylor, 

1960:300. 

DESCRIPTION.—Plants minute epiphytes, 20-

150(-220) jum tall exclusive of hairs, arising from 

a persistent subglobose, isodiametric, to cylindrical spore with or without a thick cell wall, 5-15 

jum diam., close to the same size as the cells it 

produces; spore giving rise to 1-6 erect and/or 

lateral filaments, commonly arcuate, simple to 

much branched, secund, irregular or opposite, 

branches short, tapering; cells barrel-shaped, isodiametric to cylindrical, 3-10 jum diam., 3-11 jum 

long, having a single lobate to stellate plastid 

with one pyrenoid, often bearing hyaline hairs to 

50 jum long. 

Monosporangia terminal and secund to opposite, sessile or on one-celled pedicels, ovoid, 4-

10(-15) jum diam., 6-15(-22) jum long. 

TYPE-LOCALITY.—Torquay, England. 

HOLOTYPE.—L 940285 . . . 306. 

DISTRIBUTION.—Almost cosmopolitan in distribution. 

NORTH CAROLINA SPECIMENS STUDIED.—Beaufort Inlet, sea buoy, 27 Jul 1909, Hoyt (DUKE 

141; US 33655). Beaufort, Shackleford jetty, 29 

Nov 1940, Blomquist 11472 (DUKE 676, as Callithamnion polyspermum). Ft. Macon jetty, 27 Feb 

1971, Schneider 50A (CWS). Wrightsville Beach, 

Masonboro Inlet jetty, 9 Jun 1977, Kapraun 

(WNC). Beaufort, Radio Is. jetty, 20 Jul 1982, 

Schneider 2865 (CWS). 

REMARKS.—Known as an epiphyte on a variety 

of intertidal and shallow subtidal seaweeds from 

the Beaufort and Wilmington areas, throughout 

the year. Tetrasporangia have not been observed 

on North Carolina specimens and only rarely 

elsewhere (Woelkerling, 1972). 

Although some authors have chosen to retain 

the similar Audouinella microscopica and A. parvula 

(Kylin) Dixon as separate entities (e.g., Dixon 

and Irvine, 1977), the grounds for their separation 

seem unclear and unsubstantial in light of the 

morphological diversity shown for the A. microscopica complex based upon studies of several type 

collections (Woelkerling, 1972). Stegenga and 

Mulder (1979), however, find some of the members of this complex synonymized by Woelkerling 

(1972) belong in the separate genera based upon 

their utilization of a multiple generic scheme for 

the family. There is, however, no doubt that all 

previously and recently collected Carolina specimens easily fit into the circumscription of A. 

microscopica proposed by Woelkerling (1971, 1972, 

1973b), which for the present I continue to follow. 

Examination of type material oi A. parvula (Kylin, 

1906 as Chantransia parvula) is still necessary if a 

formal transfer is to be made. 

Audouinella microscopica is reported as Acrochaetium trifilum (Buffham) Batters for adjacent Virginia (Humm, 1979). That listing follows Aziz 

(1965), which finds A. parvulum synonymous with 

several Caribbean taxa, of which A. trifilum is the 

oldest available binomial. The synonyms included A. compactum Jao and A. crassipes (B0rgesen) B0rgesen, both of which also appear as synonyms of A. microscopica (Woelkerling, 1972). Aziz 

did not mention A. microscopica in his paper, and 

at that time it was not listed for the Southeast or 

Caribbean (Taylor, 1960), but it is clear that the 

two are the same, with A. microscopica having 

historical precedence over A. trifilum. I therefore 

reject usage of the latter name. 

Audouinella ophioglossa, new species 

FIGURE 3 

Plantae epiendophyticae, 1.0-2.2 mm elatae, 

spora persistenti magna globosa exorientes, 7.5-

12.5 jum diametro; spora basilis filum singulare 

unramosum aut ramosum furcatum irregulare 

contortum producens qui hospitem cum cellulas 

2-4 jum diametris profunde penetrans, vel basem 

unicellulam remanens; spora basalis 1-2 axes erectas ramosas producens, secundas ad alternas et 

oppositas, rami plures communes super; cellulae 

filorum erectorum cylindricae, 4-5 jum diametris, 

45-55 jum longae segmentis infernis, cellula omnis 

plastum singularem lobatum ad spiralem parietalem cum pyrenoide singulari inconspicua capiens; trichomata unicellularia terminalia vel lateralia, ad 2 mm, saepe cum ramis gametangiis 

consociata. 
14 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

Monosporangia terminalia in ramis brevibus 

aut lateralia, secunda ad alterna aut opposita, 

sessilia aut in pedicellis unicellularis, singularia 

vel binata, ovoidea ad ellipsoidea, 6.0-12.5 jum 

diametris, 12.5-20.0 jum longa, plerumque cum 

gametangiis consociata. Plantae monoeciae, rami 

spermatangiis quadriverticillati, oppositi vel minus plerumque ramus unicus ad uterque nodum 

in seriebus longis in partibus superis plantae; 

spermatangia binata in pedicellis unicelluris, globosa, 2.5 jum diametris; carpogonia lateralia, opposita vel juxta monosporangium, aliquando in 

verticillo ramorum spermatangiorum facientia, 

non nisi unum carposporophytum ad uterque 

nodum producentia, saepe in seriebus longis interrupts; carposporangia terminalia globosa, 10-

15 jum diametris. 

Specimen holotypicum a SCUBA lectum, 20 

VII 1973, 34°23.0'N, 76°36.0'W (North Carolina), in profunditatae 22 metrorum, in Dudresnaya 

crassa Howe, leg. R.B. Searles, in herbariis Universitatis Dukensis, No. 16585, 96; Isotypi DUKE, 

US 88918. 

DESCRIPTION.—Plants epiendophytic, 1.0-

2.2mm tall, arising from a persistent, large, globose spore, 7.5-12.5 jum diam., which produces a 

single unbranched or forked branched irregularly 

contorted filament that deeply penetrates host 

tissue with cells 2-4 jum diam., or remains as a 

unicellular base; basal spore giving rise to 1-2 

erect branched axes, secund to alternate and 

opposite, more common above; cells of the erect 

filaments cylindrical, 4-5 jum diam., 45-55 jum 

long in lower segments; each cell containing a 

single lobate to spiral parietal plastid with a single 

inconspicuous pyrenoid, unicellular hairs terminal or lateral, to 2 mm, often associated with 

gametangial branches. 

Monosporangia terminal on short branches or 

lateral, secund to alternate or opposite, sessile or 

on one-called pedicels, single or in pairs, ovoid to 

ellipsoid, 6.0-12.5 jum diam., 12.5-20.0 jum long, 

commonly associated with gametangia. Monoecious, spermatangial branches in a whorl of 4, 

opposite paired or less commonly one per node, 

in long series in the upper parts of the plant; 

spermatangia paired on one-celled pedicels, globose, 2.5 jum diam.; carpogonia lateral, opposite 

or adjacent to a monosporangium, occasionally 

forming in a whorl of spermatangial branches, 

producing only one carposporophyte per node, 

often in a long interrupted series; carposporangia 

terminal, globose, 10-15 jum diam. 

TYPE-LOCALITY.—Onslow Bay, North Carolina, USA. 

HOLOTYPE.—DUKE 16585. 

DISTRIBUTION.—Endemic to North Carolina as 

far as currently known. 

NORTH CAROLINA SPECIMENS STUDIED.—Onslow Bay, reef, 18 Sep 1972, Schneider 553e (CWS). 

Onslow Bay, reef, 19 Jul 1973, Schneider 665a 

(CWS). Onslow Bay, reef, 20 Jul 1973, Searles 

(Holotype, DUKE 16585; Isotypes, DUKE 

16586, 16587, US 88918). Onslow Bay, reef, 20 

Jul 1973, Schneider 716a (Topotypes, CWS). Onslow Bay, reef, 19 Jun 1980, Searles (DUKE 16589). 

REMARKS.—Audouinella ophioglossa is known 22-

35 m offshore on Dudresnaya crassa, June-September. The species is named for the general appearance of the basal spore and often forked endophytic filament, which resembles the head of a 

snake with extended tongue (see Figure 3b-d). Of 

the several species worldwide with a persistent, 

obvious basal spore which produces a deep penetrating endophytic filament, this newly described plant is most similar to Audouinella subtilissima (Kutzing) Garbary, an epiendophyte of 

Dudresnaya and other species along the French 

coast. The new taxon, A. ophioglossa, differs from 

A. subtilissima in that its branches do not taper 

toward the tips, in having blunt apices, a smaller 

filament diameter (4-5 vs. 6-7 jum), and conspicuous unicellular hairs (see Figure 3fh,j). Hamel 

(1928a) reports A. subtilissima as having some 

filaments end in pseudohairs that do not appear 

to be unicellular, a feature not illustrated by him 

or by Kutzing (1861). Hamel (1928a) describes 

and illustrates other characters of A. subtilissima 

that vary from the North Carolina plant: oval to 

oblong carposporangia, carpogonia often distal to 

spermatangia on the same branch, and laterally 

disposed spermatangial branches. In A. ophiog-
NUMBER 22 15 

lossa, the carpogonia are either proximal to spermatangial branches (Figure 3/), paired with one 

(Figure 3i), or are found as one branch in a whorl 

of 4, the remainder being male axes (Figure 3h). 

The verticillate arrangement of male branches in 

the new taxon (Figure 3hJ) is unique for this 

species cluster of Audouinella. In A. ophioglossa, 

monosporangia, male branches or carpogonia are 

commonly oppositely arranged (Figure 3fl,m), or 

a carpogonium is disposed opposite to a monosporangium (Figure 3/). Opposite spermatangia 

are found occasionally in Acrochaetium kuckuckianum Hamel (1928b), which has a similar mode 

of attachment to A. ophioglossa. Its size and dioecious habit still separate it clearly from the new 

species. 

From Audouinella corymbifera, the only other 

member from North Carolina having a persistent 

spore with a penetrating endophytic filament, A. 

ophioglossa, is distinguished by a smaller basal 

spore, narrower erect filaments, its monoecious 

habit and the disposition of spermatangial 

branches. 

Audouinella saviana (Meneghini) Woelkerling 

Audouinella saviana (Meneghini) Woelkerling 1973b:560, figs. 

56-60.—Schneider, 1976:138.—Searles and Schneider, 

1978:100. 

Callithamnion savianum Meneghini, 1840:511. 

Acrochaetium thuretii (Bornet) Collins and Hervey, 1917:98. 

Audouinella thuretii (Bornet) Woelkerling, 1971:36, figs. 12, 24. 

[Not Audouinella thuretii sensu Kapraun, 1980:41.] 

DESCRIPTION.—Plants epiphytic to epiendophytic, to 4 mm tall, arising from a compact, 

mostly superficial prostrate system of short, simple or branched filaments, free from each other 

to coalesced into an irregular pseudoparenchymatous disc, the original spore not recognizable; 

prostrate filaments the same size or distinctly 

larger than erect ones; erect filaments with irregular or alternate to secund branching, cells of 

main axes and branches cylindrical 7-14 jum 

diam., 20-60 jum long, sometimes 4-6 jum diam. 

in the ultimate branches, unicellular hairs absent 

or rare, each cell containing a single parietal 

lobate plastid. 

Monosporangia single or paired, adaxially secund to irregularly arranged, sessile or on 1 (or 2) 

celled elongate pedicels, ovoid to ellipsoid, 10-15 

jum diam., 18-27 jum long. Tetrasporangia cruciate, with shape and positions similar to monosporangia, often borne simultaneously, 17-24 jum 

diam., 26-34 jum long. Monoecious, spermatangia 

sparse, on short branchlets near the base of 

branches; carpogonia sparse and solitary on lower 

segments near the base of branches, carpospores 

9-13 jum diam., 18-21 jum long. 

TYPE-LOCALITY.—Genoa, Italy. 

HOLOTYPE.—FL. 

DISTRIBUTION.—Atlantic North America, Bermuda, Caribbean, Atlantic Europe, Mediterranean, Juan Fernandez Islands, Australia, California. 

NORTH CAROLINA SPECIMENS STUDIED (representative).—Beaufort, Duncan Breakwater, 19 

Oct 1940, Blomquist 11447A (DUKE 134, 1459), 

Ft. Macon jetty, 27 Jun 1960, Aziz (DUKE 114, 

116, 159). Marshallberg, Cowper Is., 29 Aug 1962, 

Wirtensen (DUKE 115). Beaufort, Bird Shoals, 10 

Jun 1971, Brauner (DUKE 12459). Wrightsville 

Beach, Figure 8 Is., Jun 1972, Kapraun (NCW). 

Onslow Bay, reef, 9 Jun 1974, Schneider 904 

(DUKE 13514; CWS). Beaufort, Radio Is. jetty, 

13 Jun 1978, Schneider 78-4-17 (CWS). Onslow 

Bay, wreck, 18 Jun 1978, Schneider 78-6-9 (DUKE 

16271; CWS). Beaufort, Radio Is. jetty, 14 Sep 

1981, Schneider 2727 (CWS). Cape Lookout jetty, 

5 Jul 1982, Schneider 2852 (CWS). 

REMARKS.—Epiphytic usually on Codium but 

also known on Dictyopteris and other coarse algae 

from shallow subtidal and deep offshore habitats 

to 30 m, May-October. 

Dixon and Irvine (1977) and Garbary (1979) 

question the conspecificity of Audouinella thuretii 

(Bornet) Woelkerling with A. saviana as proposed 

by Woelkerling (1973b). In a numerical taxonomic approach to the Acrochaetiaceae, Garbary 

(1979) finds dissimilarity in some published descriptions of A. thuretii and A. saviana and states 

"that [Woelkerling's] conclusions are so different 

from those reached by other workers is perhaps a 

reflection of the limitations of dealing with her-
16 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

barium specimens of acrochaetioid algae." I reject 

the inference that taxa can be distinguished on 

the basis of written description alone, especially 

if these are not type descriptions, as is often the 

case in Garbary's (1979) report. Therefore, I continue to follow the results of Woelkerling (1973b) 

who, using herbarium specimens, found type material of the two taxa conspecific, and accept A. 

saviana as the epithet for North Carolina plants. 

This taxon may be an alternate generation of A. 

dasyae (see prior discussion and Stegenga and 

Borsje, 1976). 

Although I have not found unicellular hairs on 

specimens from the Carolinas, in conformity with 

those from New England (Woelkerling, 1973b), 

Hall and Eiseman (1981) reported them on plants 

during the winter months in Florida. 

Audouinella secundata (Lyngbye) Dixon 

Audouinella secundata (Lyngbye) Dixon in Parke and Dixon, 

1976:590.—Kapraun, 1980:41, figs. 23-25. 

Callithamnion daviesii var. secundatum Lyngbye, 1819:129, pi. 

41: figs. B4-B6. 

Colaconema secundata (Lyngbye) Woelkerling, 1973a:94, figs. 

7, 8.—Searles and Schneider, 1978:100. 

Acrochaetium virgatulum (Harvey) Bornet, 1904:xxii.—Williams, 1948:690; 1951:153. 

Kylinia virgatula (Harvey) Papenfuss, 1947:437.—Taylor, 

1962:214. 

DESCRIPTION.—Plants epiphytic or epizoic, to 

3 mm tall, scattered to caespitose, arising from a 

uni- to bistratose, generally compact pseudoparenchymatous disc, the original spore not persistent; disc giving rise to several erect, usually freely 

and irregularly branched with laterals often consisting of only 1-5 cells, with or without lateral 

and teminal unicellular hairs to 300 jum long; 

cells of main axes cylindrical 8-15 (-20) jum diam., 

15-100 jum long, each with an axial or parietal 

stellate plastid and single central pyrenoid. 

Monosporangia lateral or terminal on shorter 

lateral branches, sessile or pedicellate, single, 

paired or rarely in groups of 3, ovoid, (6-) 10-20 

jum diam., (10-)15-26(-32) jum long. 

TYPE-LOCALITY.—Kvivig, Faeroes Islands. 

HOLOTYPE.—C. 

DISTRIBUTION.—Atlantic coast of North America (not reported from Georgia), Sargasso Sea, 

Europe, Mediterranean, Canary Islands, Australia. 

NORTH CAROLINA SPECIMENS STUDIED.—Beaufort, Pivers Is., 10 Aug 1907, Hoyt (DUKE 188 D; 

US 7l721a,b). Beaufort, Shackleford jetty, 12 

Aug 1907, Hoyt (US 71721c). Ft. Macon jetty, 15 

Apr 1909, Hoyt (DUKE 188A; US 33656). Beaufort, Shackleford jetty, 19 Ju n 1960, Aziz (DUKE 

126, 131, 132). Beaufort, Bird Shoals, 10 Jun 

1971, Brauner (DUKE 12457, 12459). Beaufort, 

Bird Shoals, 20 Jun 1971, Brauner (DUKE 12480). 

REMARKS.—Epiphytic on a variety of algae 

intertidally, April-August. 

Although some authors maintain a specific distinction between Audouinella secundata and A. virgatula (Harvey) Dixon (e.g., Dixon and Irvine, 

1977; Kornmann and Sahling, 1978), I have observed morphological plasticity in North Carolina 

plants that corroborates the findings leading to 

the recent synonomy of the two taxa (Woelkerling, 1973a, 1973b). 

Easiest to distinguish in young stages, the prostrate system develops from an orbicular, parenchymatous group of cells. This disc may later 

proliferate as prostrate filamentous axes, obscuring the central disc and becoming more similar 

to later stages of Audouinella dasyae and A. saviana. 
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1962. Some Lza^ora-inhabiting Species of Acrochaetium. 

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Aziz, K.M.S. 

1965. Acrochaetium and Kylinia in the Southwestern North 

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Batters, E.A.L. 

1902. A Catalogue of the British Marine Algae. Journal 

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B0rgesen, F. 

1909. Some New or Little Known West Indian Florideae. Botanisk Tidsskrift, 30:1-19, 11 figures, 2 

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1915. The Marine Algae of the Danish West Indies, Part 

III: Rhodophyceae (1). Dansk Botanisk Arkiv, 

3(1 a): 1-80, 86 figures. 

Bornet, E. 

1904. Deux Chantransia corymbifera Thuret: Acrochaetium et 

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51 (supplement) :xiv-xxiii. 

Borsje, W.J. 

1973. The Life History of Acrochaetium virgatulum (Harv.) 

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Collins, F.S. 

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Collins, F.S., and A.B. Hervey 

1917. The Algae of Bermuda: Contributions of the Bermuda Biological Station for Research, number 68. 

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Dillwyn, L.W. 

[1809.] British Confervae. 87 pages, London. W. Phillips. 

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Drew, K.M. 

1928. A Revision of the Genera Chantransia, Rhodochorton, 

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Feldmann, J. 

1962. The Rhodophyta Order Acrochaetiales and its 

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Garbary, D. 

1978. On the Phylogenetic Relationships of the Acrochaetiaceae (Rhodophyta). British Phycological Journal, 13:247-254. 

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1981. The Seagrass Epiphytes of the Indian River, Florida I: Species List with Descriptions and Seasonal 

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Hamel, G. 

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Holmgren, P.K., W. Keuken, and E.K. Schosield 

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Humm, H.J. 

1979. The Marine Algae of Virginia. Special Papers in 

17 
18 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

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1980. An Illustrated Guide to the Benthic Marine Algae of 

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1978. Meeresalgen von Helgoland, iii -I- 289 pages. Hamburg: Biologische Anstalt Helgoland. 

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1976. Check-List of British Marine Algae—Third Revision. Journal of the Marine Biological Association, 

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1976. Spatial and Temporal Distributions of Benthic 

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1-14. 

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1960. Marine Algae of Eastern Tropical and Subtropical Coasts 

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van den Hoek, C , and A.M. Cortel-Breeman 

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1968. Morphology and Reproduction of the Red Alga 

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Williams, L.G. 

1948. Seasonal Alternation of the Marine Floras at Cape 

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Pearse and L.G. Williams, The Biota of the Reefs 

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NUMBER 22 19 

Wiseman, D.R. 

1978. Benthic Marine Algae. In R.G Zingmark, An Annotated Checklist of the Biota of the Coastal Zone of South 

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Woelkerling, W.J. 

1970. Acrochaetium botryocarpum (Harv.) J. Ag. (Rhodophyta) in Southern Australia. British Phycological 

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1971. The Morphology and Taxonomy of the Audouinella 

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Australian Journal of Botany, Supplement, 1:1-91, figures 1-27. 

1972. Studies on the Audouinella microscopica (Naeg.) 

Woelk. Complex (Rhodophyta). Rhodora, 74:85-

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1973a. The Audouinella Complex (Rhodophyta) in the 

Western Sargasso Sea. Rhodora, 75:78-101, figures 

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1973b. The Morphology and Systematics of the Audouinella 

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figures 1-83. 
2 0 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

FIGURE 1.—Audouinella affinis: a, surface view of a basal spore with a single erect filament; b, 

transverse section of Dictyota cells displaced by the penetrating spore. A. bispora: c, epiphytic and 

endophytic (broken lines) attachment filaments; d, bisporangia in various stages of development; 

ef, monosporangia. A. botryocarpa: g, detail of two cells showing lobate parietal plastids and 

associated multiple pyrenoids (clear); h,i, tetrasporangia; j , branch bearing monosporangia 

(some spent) and a single tetrasporangium; k,l, monosporangia. A. dasyae: m,n, axis rising on 

oblique angle from two-celled basal system; o, basal cells immersed in cell wall of Ceramium; p, 

maturing basal system with original panduriform basal cell still obvious. 
NUMBER 22 21 
2 2 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

FIGURE 2.—Audouinella daviesii: a-c, various arrangements of monosporangia. A. densa: d, e, early 

prostrate system showing septate spore; /, habit on Chaetomorpha, single and seriate monosporangia; g-i, single and 2-3 seriate monosporangia. A. hoytii: j , habit of a monosporangiate plant, 

showing obvious persistent basal spore. A. hypneae: k, basal system with erect filaments; /, 

monosporangia in series. A. infestans: m, endophytic system and emergent axis with monosporangia and a single unicellular hair. 
NUMBE

R 22 

23 
2 4 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

FIGURE 3.—Audouinella ophioglossa, new species: a, basal spore, without endophytic penetrating 

filament, adjacent to Dudresnaya cortical cells; b-e, basal spores with erect and forked penetrating 

filaments;/, habit, showing paired spermatangia (s) and a spent monosporangium (SM) opposite 

to a carpogonium (c); g, monosporangia (M) and a whorl of spermatangia; h,i, primarily male 

axis bearing carpogonia (c) with extended trichogynes, hair cell present; j , male axis with 

numerous unicellular hairs (H) among whorls of spermatangia; k, carpogonium with 4 spermatangia on trichogyne; /, opposite carpogonia; m, opposite or single monosporangia, some 

spent (SM) ; n-p, development of carposporophyte with carpospores (cs). 
NUMBER 22 25 

um

 ( a-b, e-f,h-p 



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