smithsonian contributions to botany • number 105

Smithsonian Institution

Scholarly Press

Smithsonian Institution

Scholarly Press

The Mesophotic,

Coral Reef–Associated,

Marine Algal Flora of

Puerto Rico, Caribbean Sea

David L. Ballantine, 

Hector Ruiz Torres,

and Nilda E. Aponte
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smithsonian contributions to botany • numb e r 1 0 5

The Mesophotic, 

Coral Reef–Associated, 

Marine Algal Flora of 

Puerto Rico, Caribbean Sea

David L. Ballantine, 

Hector Ruiz Torres, 

and Nilda E. Aponte

WASHINGTON, D.C.

2016
ABSTRACT

Ballantine, David L., Hector Ruiz Torres, and Nilda E. Aponte. The Mesophotic, Coral Reef-Associated, 

Marine Algal Flora of Puerto Rico, Caribbean Sea. Smithsonian Contributions to Botany, number 105, viii 

+ 41 pages, 5 figures, 3 tables, 2 appendixes, 2016. — Deepwater open-circuit scuba, dredging, submersible, 

and technical mixed-gas (closed-circuit) rebreather diving collections of marine benthic algae made over the 

last approximately 30 years in Puerto Rico are summarized in this account. In total, 186 taxa (166 identified 

to species) comprising 60% Rhodophyta, 11% Phaeophyceae, and 29% Chlorophyta are reported from 

depths greater than 35 m. Eighty-nine of these (56% of taxa identified to species) from the Puerto Rican 

mesophotic are thought to be the deepest known distributional records for the species recognized. Forty-three 

species (8% of the entire benthic flora of Puerto Rico) are mostly or entirely restricted to depths greater than 

35 m. KEYWORDS: deepwater algae, mesophotic, Puerto Rico, tropical west Atlantic.

Cover images, from left to right: Edge of insular shelf, offshore from La Parguera; technical divers’ descent to 

the mesophotic; bottom habit, edge of insular shelf, “Weinberg,” offshore from La Parguera.

Published by SMITHSONIAN INSTITUTION SCHOLARLY PRESS

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Library of Congress Cataloging-in-Publication Data

Names: Ballantine, David L., author. | Ruiz Torres, Héctor J., author. | Aponte, Nilda, author. | Smithsonian Institution 

Scholarly Press, publisher.

Title: The mesophotic, coral reef-associated, marine algal flora of Puerto Rico, Caribbean Sea / David L. Ballantine, 

Hector Ruiz Torres, and Nilda E. Aponte.

Other titles: Smithsonian contributions to botany ; number 105. 0081-024X

Description: Washington, D.C. : Smithsonian Institution Scholarly Press, 2016. | Series: Smithsonian contributions to 

botany, ISSN 0081-024X ; number 105 | Includes bibliographical references and index.

Identifiers: LCCN 2016032644

Subjects: LCSH: Marine algae—Puerto Rico—Identification. | Marine algae—Ecology—Puerto Rico. | Marine plants—

Puerto Rico. | Benthic plants—Puerto Rico. | Deep-sea ecology—Puerto Rico.

Classification: LCC QK571.5.P9 B35 2016 | DDC 579.8/177097295—dc23 | SUDOC SI 1.29:105

LC record available at https://lccn.loc.gov/2016032644

ISSN: 1938-2812 (online); 0081-024X (print)

Publication date (online): 10 November 2016

Ó The paper used in this publication meets the minimum requirements of the American National Standard for Permanence of Paper for Printed Library Materials Z39.48–1992. 
Contents

LIST OF FIGURES v

LIST OF TABLES vii

INTRODUCTION 1

ENVIRONMENT AND HABITAT 2

MATERIAL AND METHODS 3

RESULTS 3

Rhodophyta 3

Heterokontophyta 15

Chlorophyta 17

DISCUSSION 23

ACKNOWLEDGMENTS 25

APPENDIX A: SUMMARY OF PUERTO RICAN MESOPHOTIC 

ALGAL SPECIMENS 27

APPENDIX B: COLLECTION LOCALITY DATA 31

REFERENCES 35

INDEX OF GENERA AND SPECIES 39

Figures

1. Map of Puerto Rico and associated islands showing collection locations 4

2. Encrusting deepwater rhodophyte species 5

3. Deepwater Rhodophyta species 9

4. Algal deepwater benthic habitats sampled 13

5. Deepwater Chlorophyta species 18

Tables

1. Algal species mostly depth-restricted to the mesophotic realm 

in Puerto Rico 23

2. Numbers of genera and species of families represented by four 

or more species reported from 35 m or greater in depth in Puerto Rico 24

Appendix A

A1. Identified mesophotic algal specimens from Puerto Rico 27

The Mesophotic, Coral Reef–Associated, 

Marine Algal Flora of Puerto Rico, 

Caribbean Sea

David L. Ballantine,1,4* Hector Ruiz Torres,2,4

and Nilda E. Aponte3,4

1 Department of Botany, National Museum 

of Natural History, Smithsonian Institution, 

Washington, D.C. 20560, USA.

2 HJR Reefscaping, P.O. Box 1126, Hormigueros, Puerto Rico 00660, USA.

3 Department of Biology, University of Puerto 

Rico at Bayamón, 170 Carretera 174, Bayamón, Puerto Rico 00956, USA.

4 Formerly at Department of Marine Sciences, 

University of Puerto Rico, Mayagüez, Puerto 

Rico 00681, USA.

* Correspondence: ballantined@si.edu

Manuscript received 16 February 2016; accepted 13 May 2016.

INTRODUCTION

The term “mesophotic” is somewhat difficult to define precisely, in part because its 

depth ranges vary geographically. Nevertheless, Hinderstein et al. (2010:248) characterized mesophotic communities as consisting of light-dependent corals and associated communities “typically found at depths ranging from 30–40 m and extending to over 150 m 

in tropical and subtropical regions. Due to difference in water transparency, the lower 

depth limit is to about 90 m in the Caribbean and to greater than 150 m in Hawaii. The 

dominant communities providing structural habitat in the mesophotic zone can be comprised of coral, sponge, and algal species.” Because of the decreased light at these depths, 

these environments have been referred to as “twilight zones” (Frick and Knauer, 1986; 

Pyle, 1996; Brokovich et al., 2008).

For the tropical and subtropical western Atlantic, published accounts of deep reefassociated algae are proportionately few, and thus, the benthic flora (>35 m) in the region 

remains poorly to incompletely known, largely because of logistical difficulties in working in water deeper than is safely accessible by scuba. Scuba collections are limited by 

decreased bottom time at depth and limits imposed on depth by modern scientific diving 

safety standards. Collection by dredging is a low-technology alternative that allows access to deeper-water environments; however, such collections are largely indiscriminate 

because they are conducted remotely from the surface. Dredge collections also suffer 

from the fact that the process is destructive to the community being sampled and may 

physically damage collections, particularly delicate forms. Thus, deeper-water studies 

have historically been largely hampered by existing technology. 

The advent of submersible and remotely operated vehicle (ROV) availability to the 

scientific research community has opened new avenues for examination of previously inaccessible communities. These devices afford direct access to deeper communities and 

allow for greater time spent at depth. Nevertheless, submersible- and ROV-based collections have their own shortcomings. In the first place, collections made by submersibles 

have limited fine-scale selectivity as the collector is normally meters away from the substratum collected and hence is unable to see small specimens from close range. Second, 

the submersible and ROV collection process is relatively crude, utilizing a mechanical arm 

that frequently results in damage to delicate forms and makes selective removal of closely 

adherent crustose forms difficult. Successful submersible and ROV collection is reliant 

on the competent handling of a manipulator arm and the capacity to store and segregate collections made at the bottom. One obvious restriction to the use of submersibles 
2 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

and ROVs is related to availability and expense of appropriate 

support vessels. Earle (1985) discussed the research potential of 

“one-atmosphere diving” suits as well as one-man microsubmersibles such as Deep Rover. However, work from these platforms to 

date has been largely limited to commercial applications. 

In water deeper than that safely accessible by scuba depth, the 

limitations mentioned above are largely ameliorated by technology that permits an experienced collector close-range access to the 

environment in which the algae live. Thus, the recently expanding 

utilization of trimix rebreather (closed-circuit) diving now allows 

substantially less obstructed access to this environment. Although 

still of limited practice, it has the advantage of allowing a collector 

(as is true for a scuba diver) immediate contact with the substratum, which allows selectivity in collection. One of the drawbacks 

to technical rebreather diving in general is the substantial time that 

a diver must train to be certified on the equipment. The equipment 

itself is also expensive, and a diving team requires infrastructure 

support (Sherman et al., 2013). Deep diving must also still be 

regarded as inherently dangerous. As might be expected, experienced algal collectors with technical dive training are few.

Historically, extensive systematic collection specifically aimed 

at deep-water algae in the tropical west Atlantic have been few. 

The earliest of these was that by Frederick (1963) on the Bermuda 

platform. Using mostly dredging collections at an average depth 

of 54 m during 1960 and 1961, Frederick recorded 97 species 

(exclusive of cyanobacteria), of which 91 were identified to species 

level. A large-scale scientific dredging program off the east coast 

of Brazil conducted under the auspices of Laboratório de Ciências 

do Mar (Lacimar) of the Universidade Federal de Pernambuco (see 

Ugadim and Pereira, 1978; Guimarães et al., 1981; see also additional references cited in the species accounts) involved numerous collections deeper than 35 m. A series of papers by Schneider 

and Searles (1973, 1975, 1976) as well as by Schneider (1974, 

1975, 1976) based mostly on scuba and dredging collections reported on the deep offshore flora of the Carolinas (east U.S. coast) 

and yielded a number of new algal species and species distributional records. Also in the western Atlantic, a series of submersible dives on the east Florida shelf (Eiseman, 1979; Eiseman and 

Blair, 1982; Hanisak and Blair, 1988a) resulted in a substantial 

increase in our knowledge of the deep algal flora of the region. 

More recently in the Bahamas, use of submersible-based dive collections also resulted in recognition of new algal species and new 

geographical records (Aponte and Ballantine, 2001; Ballantine 

and Aponte, 1996, 2002a, 2003, 2005). Dredging collections to 

90 m in the Flower Garden Banks National Marine Sanctuary 

(Gulf of Mexico) by Fredericq’s research group (reported in Gavio 

and Fredericq, 2003, 2005; Gavio et al., 2005) further resulted in 

the characterization of new algal species and new species records. 

More recently, Leichter et al. (2008) reported on distribution and 

spatial extent of algal communities off the Florida coast based on 

broad systematic groupings, using ROVs and scuba.

In the tropical western Atlantic (inclusive of Florida, the Bahamas, Jamaica, and Bermuda), only a handful of other submersiblebased studies have been conducted in reef environments in water 

greater than 50 m depth: these include the results of Lang (1974), 

Huston (1985), and Fricke and Meischner (1985) from Florida; 

Lang et al. (1975) and Liddell and Olhurst (1988) from Jamaica; 

Littler et al. (1985, 1986), Blair and Norris (1988), Liddell et al. 

(1997), and Norris and Olsen (1991) from the Bahamas; and Ballantine (1990) from Puerto Rico. Macintyre et al. (1991) reported 

a handful of algal species collected from Barbados on the basis of 

two submersible dives. Littler et al. (1985) reported the deepest 

known macroalga (to 268 m), an unidentified Corallinales. In a 

more recent paper based on deployment of a drop camera, Friedlander et al. (2014: “Deep Reefs,” paragraph 2) reported the presence of crustose coralline algae “from 312 m depth (and probably 

from 382 m)” at Pitcairn Island (Pacific Ocean), although specimens presumably were not obtained.

Other noteworthy deepwater algal reports include those by 

Dawes and Van Breedveld (1969) in the Florida Middle Grounds, 

Gulf of Mexico, who reported a number of benthic algal species 

and distributional records based on dredging and trynet collections to 73 m, and by Cheney and Dyer (1974) on the basis of 

scuba collections to 60 m. 

Systematics of deep-water algal flora in Puerto Rico effectively began with the publication of a new deep-water Halimeda

species variety (H. cryptica v. acerifolia) by Ballantine (1982) 

collected utilizing scuba. Wynne and Ballantine (1986) and Ballantine and Wynne (1987, 1988) reported three new species of 

benthic algae on the basis of scuba collections to 61 m. Subsequently, two species of a previously undescribed deepwater 

green alga, Verdigellas, were obtained utilizing dredging (Ballantine and Norris, 1994). Although these studies have provided 

insight into the diversity and distribution of deepwater species, 

initial trimix rebreather dives in southwest Puerto Rico have already yielded a handful of species new to science, particularly 

among calcified and noncalcified encrusting algae, as well as new 

geographical and depth records (including Ballantine and Ruiz, 

2005, 2008, 2010, 2011; Athanasiadis et al., 2013). 

ENVIRONMENT AND HABITAT

Mesophotic reef environments in general are principally 

noted for depth and, as a consequence, reduced light. For oceanic water off Brazil, there is a sharp decline in photosynthetically 

active radiation at 30 m, where less than 2% of surface irradiance reaches (Magalhães et al., 2015). In situ sea conditions are 

thought to be more stable in deeper water than in shallow water, 

with decreased disturbance and a lower fluctuation in water temperature; however, the presence of internal waves may periodically 

alter localized temperature regimes (Appeldoorn et al., 2016). 

Herbivory, which exerts a strong influence on algal populations 

through most of the euphotic zone, is less important in the mesophotic zone. For example, herbivorous fish populations have been 

shown to decline with depth in deepwater coral reef environments 

(Brokovich et al., 2010; Bejarano et al., 2014). Mesophotic reef 

habitats at the insular shelf break in south Puerto Rico are diverse 

in physical and geomorphic attributes. The bottom habitat may 

be slightly to steeply inclined and in some locales nearly vertical. 
NUMBER 105 • 3

The substrata available for colonization are either hard-bottom 

(primarily dead Agaricia plates) or sand; however, the extensive 

sand-bottom mesophotic habitats reported in Spalding (2012) 

were never seen in Puerto Rico. Downslope sediment transfer 

grooves, which are extensions of the spur and groove environment 

at the shelf break, are generally devoid of macroscopic biological 

growth. As a result, mesophotic community development is largely 

restricted to topographical highs (Sherman et al., 2010). Living 

coral cover is occasionally substantial; however, percent bottom 

cover is often dominated by algal and sponge growth. In terms of 

composition, few foliose algal species are present, and the flora 

is made up of mostly encrusting Corallinales species in addition 

to Peyssonneliaceae species; leafy Phaeophyta, including Dictyota

spp. and Lobophora spp.; and multispecies turf. Photographs of 

typical bottom habitat show the domination of benthic encrusting 

organisms at depths of 49 to 76 m (e.g., see in Figure 4).

MATERIAL AND METHODS

The present work is based on dredging and scuba collections 

and several submersible-based dives over the last three decades by 

the first author and the more recent deep trimix technical diving by 

the second author (see Sherman et al., 2013, for the deep technical 

diving protocols utilized). Species in this report were all collected in 

coastal waters of Puerto Rico (see Appendix A), with most effort concentrated on the southern coast (Figure 1; see also Appendix B). Additionally, collections have been conducted at Mona Island (66 km 

west of Puerto Rico) and at “El Seco,” southeast of Vieques Island 

(13 km east of Puerto Rico). Species of algae reported are limited to 

those collected at 35 m or greater depth. This is a somewhat arbitrary 

depth, partly because, for Puerto Rico, the depth is recognized as a 

transitional one between shallow and deeper affinities and is near the 

recommended limit for nondecompression open-circuit scuba diving 

on air. Voucher specimens of algae reported are deposited in the Herbario Marino Puertorriqueño of the Department of Marine Sciences, 

University of Puerto Rico (MSM), and/or in the algae collection of the 

U.S. National Herbarium, Smithsonian Institution. Authority names 

are according to Brummitt and Powell (1992). A total of 185 taxa of 

algae collected in water deeper than 35 m are listed below. An asterisk 

prior to entries represents what is considered to be the deepest (or 

equaling the deepest) recorded depths for that species. Species are 

ordered by presumed phylogenetic position (Guiry and Guiry 2016).

RESULTS

Rhodophyta

Florideophycideae, Corallinophycidae, Corallinales, Corallinaceae

*Amphiroa rigida J. V. Lamour. 

Specimens: DLB7357, 49 m; DLB7139, 50 m; DLB7552, 62 m; 

DLB7629, 70 m.

Amphiroa rigida is typically a plant of shallow water and 

reported to 11 m depth (Taylor, 1960).

*Amphiroa tribulus (J. Ellis et Sol.) J. V. Lamour. 

Specimens: DLB7358, 49 m; DLB7144, 50 m; DLB7489, 

61 m.

Amphiroa tribulus is also typically a plant of shallow water 

and reported to 18 m depth (Taylor, 1960).

*Hydrolithon abyssophila Athanas., D. L. Ballant. et H. Ruiz 

FIGURE 2I

Specimens: DLB7691, 49 m; Athanas.PR135A, 73 m; DLB7824, 

73 m; DLB7746, 76 m.

Known from 30 to at least 76 m, the species was recently described from an insular shelf habitat on the south coast of Puerto 

Rico (Athanasiadis et al., 2013). The species is extremely abundant and is probably to be found at mesophotic depths throughout the Caribbean and, perhaps by cover, is the most common 

benthic eukaryotic organism in the Caribbean at depths greater 

than 50 m (Ballantine and Ruiz, unpublished).

Hydrolithon farinosum (J. V. Lamour.) Penrose et Y. M. Chamb. 

var. chalicodictyum (W. R. Taylor) Serio

Specimens: DLBs.n., 55 m; DLB1711, 58 m; DLB1746, 

DLB7506, 61 m.

Schneider and Searles (1997) reported the species in Bermuda to a maximum depth of 27 to 40 m. The deepest known 

record for the species remains that of Frederick (1963) from Bermuda at 64 m.

*Jania adhaerens J. V. Lamour.

Specimen: DLB7385, 50 m.

This extremely broadly distributed species is mostly found 

in shallow-water environments (Taylor, 1960). It has been reported to 30 m depth in Florida by Hanisak and Blair (1988a) 

and to 35 m depth in Onslow Bay, North Carolina (Schneider 

and Searles, 1991).

*Jania cubensis Mont. ex Kütz. 

Specimens: DLB4190, 36 m; DLB7359, 49 m; DLB7488, 61 m; 

DLB7576, DLB7620, 70 m.

As Haliptilon cubense (Mont. ex Kütz.) Garbary et H. W. 

Johans, Ballantine and Aponte (2005) and Yoneshigue-Valentin 

et al. (2004) reported the species to 46 and 60 m in the Bahamas 

and Brazil, respectively.

*Jania subulata (J. Ellis et Sol.) Sond. 

Specimen: DLB3736, 36–46 m.

Jania subulata is typically known from shallow-water habitats (Littler and Littler, 2000).

Nemaliophycidae, Nemaliales, Galaxauraceae

Dichotomaria marginata (J. Ellis et Sol.) Lam.

Specimen: DLB7494, 61 m.
4 

• SMITHSONIAN CONTRIBUTIONS TO BOTANY

FIGURE 1. Map of Puerto Rico showing sampling sites. The depth contour surrounding the island represents a depth of 50 m. Map by Alice Tangerini for Smithsonian Institution.
NUMBER 105 • 5

Reported by Taylor (1960) as having been dredged to 55 m. 

The deepest known record for the species remains that of Frederick (1963) from Bermuda at 62 m.

*Dichotomaria obtusata (J. Ellis et Sol.) Lam. 

Specimen: DLB7490, 61 m.

Dichotomaria obtusata was reported to 27 m depth by 

Schneider and Searles (1973) from the offshore flora of North 

Carolina. Galaxaura obtusata var. major W. R. Taylor (1960) 

was reported as having been dredged to 53 m depth. MateoCid et al. (2013) reported collections from 50 to 56 m from 

Campeche Banks, Mexico, and Frederick (1963) reported the 

species to a depth of 59 m from Bermuda.

*Galaxaura rugosa (J. Ellis et Sol.) J. V. Lamour. 

Specimens: DLB7153, 50 m; DLB3104, 55–90 m; DLB7628, 

70 m.

Taylor (1960) indicated that G. rugosa is a shallow-water 

alga that had also been collected to 18 m depth.

*Tricleocarpa fragilis (L.) Huisman et R. A. Towns. 

Specimens: DLB7549, 62 m; DLB7557, 70 m.

FIGURE 2. Encrusting deepwater rhodophyte species. (A) Peyssonnelia gigaspora (DLB7637); sporangial sori are denoted by arrowheads. 

These tetrasporangia are the largest reproductive algal spores known (scale bar = 1.0 cm). (B) Peyssonnelia flavescens (DLB5883; scale bar = 

2.0 cm). (C) Peyssonnelia iridescens (DLB7116; scale bar = 1.0 cm). (D) Peyssonnelia incomposita (DLB7777; scale bar = 1.0 cm). (E) Peyssonnelia sp. 3 (DLB7411; scale bar = 5 mm). (F) Peyssonnelia sp. 2 (DLB7286; scale bar = 2.0 cm). (G) Ethelia sp. (DLB6355; scale bar = 

2.0 cm). (H) Polystrata fosliei (DLB7912; scale bar = 2.0 cm). (I) Hydrolithon abyssophila (Athanasiadis PR135A; scale bar = 2.0 cm). Photos 

by Hector Ruiz Torres.
6 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

Taylor (1960) also indicated that T. fragilis (as Galaxaura 

cylindrica (J. Ellis et Sol.) J. V. Lamour.) is a shallow-water alga; 

however, Dawes and Mathieson (2008) reported that the species 

has been dredged to 30 m in Florida.

Rhodymeniophycidae, Bonnemaisoniales, Bonnemaisoniaceae

*Asparagopsis taxiformis (Delile) Trevis. 

Specimens: DLB7362b, DLB7722, 49 m; DLB7384, 50 m; 

DLB7645, 70 m.

Asparagopsis taxiformis in Puerto Rican deepwater habitats 

is represented by the Falkenbergia stage (or tetrasporophyte life 

history alternate of A. taxiformis). This growth phase is typically seen in shallow-water environments in the Caribbean and 

is encountered only occasionally in deep water. The species was 

reported to 49 m in Bermuda by Frederick (1963), from 50 to 60 

m in Brazil (Magalhães et al., 2015), and to 63 m in the Pacific 

by Gilmartin (1960).

Ceramiales, Callithamniaceae

Aglaothamnion cordatum (Børgesen) Feldm.-Maz. 

Specimens: DLB4429, 37 m; DLB8077, 50 m.

The species (as Callithamnion cordatum Børgesen) has 

been collected by Schneider (1980) offshore North Carolina to 

32 m depth and to 74 m by Hanisak and Blair (1988a) from 

Florida.

*Crouania pumila Gavio, Reyes-Gómez et M. J. Wynne 

Specimen: DLB7904, 55 m.

Reported for Puerto Rico by Ballantine et al. (2015), only 

small, sterile epiphytic plants were collected. The holotype specimen was from 16 m depth in Colombia (Gavio et al., 2013), and 

other collections were made by them at 1–2 and 37 m.

*Crouanophycus latiaxis (I. A. Abbott) Athanas. 

Specimen: DLB7173, 50 m.

Depth information was not provided for the species (as Antithamnionella latiaxis; Abbott, 1979) from the holotype location in St. Croix, U.S. Virgin Islands; however, it is presumed to 

have been collected in shallow-water reef environments. Bucher 

and Norris (1995) reported the species (also as A. latiaxis) to a 

maximum depth of 12 m from Martinique.

*Seirospora occidentalis Børgesen 

Specimens: DLB3706, 36 m; DLB1214, DLB1818, 40 m; 

DLB1802, DLB7337, 50 m; DLB7503, 61 m; DLB7881, 

82 m; DLB2158, 87 m.

Seirospora occidentalis is a fairly common element in the 

deepwater turf flora of Puerto Rico.

*Seirospora viridis Aponte et D. L. Ballant. 

Specimen: DLB1721, 58 m.

This species is currently known only from Puerto Rico, 

where it was described from shelf edge habitats at 18 m depth 

(Aponte and Ballantine, 1995).

Ceramiaceae

*Antithamnion antillanum Børgesen

Specimens: DLB7156, 50 m; DLB7883a, 82 m.

Macintyre et al. (1991) reported the species (as A. lherminieri 

(P. Crouan et H. Crouan) Bornet ex Nasr) to 74 m in Barbados.

*Antithamnion decipiens (J. Agardh) Athanas.

Specimens: DLB1212, 40 m; DLB7362a, DLB7720, 49 m; 

DLB7405, 50 m; DLB7689, 64 m; DLB7915, 70 m; 

DLB7883b, 82 m.

Antithamnion decipiens was reported to 76 m in the Bahamas (Ballantine and Aponte, 2005). As A. ogdeniae I. A. Abbott, 

the species was known from 70 m in Hawaii (Agegian and Abbott, 1985).

Antithamnionella breviramosa (E. Y. Dawson) E. M. Woll. 

Specimen: DLB4419, 37 m.

The original report of the species from Puerto Rico was 

based on specimens collected at intermediate depths, to ~30 m 

(Ballantine and Wynne, 1986). Antithamnionella breviramosa

has been reported to a depth of 87 m in Florida by Hanisak and 

Blair (1988a).

*Antithamnionella graeffei (Grunov) Athanas. 

Specimen: DLB7774, 61 m.

Antithamnionella graeffei was reported to 45 m from 

Onslow Bay, North Carolina (Schneider, 1984), and Børgesen 

(1945) indicated that the species (as Antithamnion flagellata

Børgesen) occurred to 55 m in Mauritius.

Balliella pseudocorticata (E. Y. Dawson) D. N. Young 

Specimens: DLB3693, DLB7959, 36 m; DLB7717, 49 m; 

DLB7423, 50 m; DLB1720, 58 m.

As a taxonomic synonym, Bakothamnion curassavicum

C. Hoek (1978) was based on material collected between 10 and 

60 m in coral reef habitats at Curaçao. Balliella pseudocorticata

has also been reported from 27 to 40 m in Bermuda (Schneider 

and Searles, 1997).

*Ceramium bisporum D. L. Ballant. 

Specimens: DLB3714, 46–55 m; DLB7170, 50 m; DLB4726, 

60 m; DLB8025, 70 m; DLB3195, 80 m; DLB3188, 80–

100 m.

In Puerto Rico, the species is found exclusively in deep 

water (see Ballantine, 1990); however, C. bisporum was recently 

reported as a shallow-water epiphyte in the Mediterranean Sea 

(Sartoni and Boddi, 2002) and also in Colombia by Rincon-Diaz 

et al. (2014) at 12.5 m.
NUMBER 105 • 7

Ceramium leptozonum M. A. Howe

Specimen: DLB8096, 50 m.

Ceramium leptozonum was first reported from Puerto Rico 

by Ballantine et al. (2011). Frederick (1963) reported the species 

to 62 m in Bermuda, and Hanisak and Blair (1988a) reported C. 

leptozonum to 92 m in Florida.

*Ceramium nitens (C. Agardh) J. Agardh 

Specimen: DLB7492, 61 m.

Ceramium nitens is a very common element of the shallowwater flora in Puerto Rico, frequently found in coral reef 

environments.

Ceramium spp. 

Specimens: DLB1209, DLB7392, 50 m; DLB7888, 82 m.

*Gayliella transversalis (Collins et Herv.) Cho et L. McIvor

Specimens: DLB7554b, 62 m; DLB8018, 70 m.

The species is based on Ceramium transversalis Collins et 

Herv. (1917) from Bermuda without report of depth. As Ceramium flaccidum (Harvey ex Kütz.) Ardiss., the species was 

reported to 33 m by Hanisak and Blair (1988a) from Florida, 

and as C. tranversale, Børgesen (1918) indicated that it was 

found in the intertidal in the U.S. Virgin Islands. Schneider and 

Searles (1991) indicated that the species (as C. byssoideum

Harv.) was collected to 21.5 m depth offshore Onslow Bay, 

North Carolina.

*Perikladosporon abaxiale D. L. Ballant. et Aponte 

Specimen: DLB2173a, 87 m.

The species was described on the basis of specimens from 

the Bahamas collected at a depth of 76 m and from a submersible 

collection in Puerto Rico (Ballantine and Aponte, 2005). These 

represent the only collections known for the species.

Dasyaceae

Dasya sp.

Specimen: DLB8079, 50 m.

*Dictyurus occidentalis J. Agardh 

Specimen: DLB3732, 36–46 m.

Reported by Taylor (1960) as having been dredged to a 

depth of 30 m. Guimarães et al. (1981) also reported specimens 

dredged from 40 m in Brazil.

*Heterosiphonia crispella (C. Agardh) M. J. Wynne 

Specimens: DLB7073, 52 m; DLB8072, 70 m.

As Heterosiphonia wurdemannii (Baily ex Harvey) Falkenberg, this species was reported to 30 m depth from the offshore 

North Carolina flora (Schneider and Searles, 1973), to 58 m in 

Bermuda by Frederick (1963), and to 65 m in Hawaii (Agegian 

and Abbott, 1985). 

Heterosiphonia sp.

Specimens: DLB8097, 50 m; DLB8017, 70 m.

This entity represents an undescribed species that superficially resembles H. crispella, differing by the nature of its nearcomplete axial cortication.

Delesseriaceae

*Apoglossum gregarium (E. Y. Dawson) M. J. Wynne 

Specimens: DLB1779, 61 m; DLB3122, 100 m.

In the original report of this species from Puerto Rico, collections were reported from 18 to 30 m (Ballantine and Wynne, 

1985). The species has also been reported from 50 m depth in the 

Mediterranean (Tsiamis and Bellou, 2010). 

*Augophyllum wysorii S.-M. Lin, Fredericq et Hommersand 

Specimen: DLB7441, 49 m.

Augophyllum wysorii was described from Caribbean Panama (Lin et al., 2004) at 12–15 m depths. The species was originally reported from Puerto Rico by Ballantine et al. (2015).

*Branchioglossum prostratum C. W. Schneid. 

Specimen: DLB1745, 61 m.

The species is based on collections off North Carolina at 60 m 

(Schneider, 1974). Eiseman (1979) reported the species from 48 m 

on the east Florida continental shelf. Branchioglossum prostratum was first reported from Puerto Rico by Ballantine and Wynne 

(1987). Despite the fact that the species is only rarely found in water 

shallower than 25 m in Puerto Rico, a shallow-water collection 

from American Samoa was reported by Skelton and South (2007).

*Frikkiella pseudoprostrata (D. L. Ballant. et M. J. Wynne) M. 

J. Wynne et C. W. Schneid.

Specimens: DLB7074, 46 m; DLB1794, DLB7429, 50 m; 

DLB7902, 55 m; DLB1733, 58 m; DLB1755, 61 m; 

DLB7057, 52 m; DLB7889, 82 m.

Frikkiella pseudoprostrata is based on Branchioglossum 

pseudoprostratum D. L. Ballant. et M. J. Wynne (1987), where it 

was originally reported from 50 m depth in Puerto Rico. It is also 

known to 76 m in the Bahamas (Ballantine and Aponte, 2005).

Frikkiella searlesii M. J. Wynne et C. W. Schneid.

Specimen: DLB1733, 61 m.

Wynne and Schneider (1996) separated Frikkiella from Branchioglossum on the basis of the origin of lateral branches and the 

origin of tertiary cell rows. They discerned that the original circumscription of Branchioglossum pseudoprostratum included two 

separate species (i.e., F. pseudoprostrata and F. searlesii). The generitype was collected from 31 m depth in Mona Island, Puerto Rico 

(Ballantine and Wynne, 1987). The species has also been collected 

to 40 and 53 m in Bermuda (Wynne and Schneider, 1996), to 60 m 

in Brazil (Peruzzi de Oliveira and Yoneshigue-Valentin, 2014), and 

to 76 m in the Bahamas by Ballantine and Aponte (2005).
8 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

*Hypoglossum anomalum M. J. Wynne et D. L. Ballant. 

FIGURE 3C

Specimens: DLB3705, 36 m; DLB1823, 40 m; DLB1789, 

DLB8093, 50 m; DLB1719, 55 m; DLB4685, 55–60 

m; DLB7455, 57 m; DLB7455, 61 m; DLB7954, 67 m; 

DLB8023, DLB8057, 70 m; DLB3200, 80 m; DLB7680, 

82 m; DLB2164, 87 m; DLB3127, 91 m.

Hypoglossum anomalum, originally described from specimens collected at depths of 30 to 60 m in Puerto Rico (Wynne 

and Ballantine, 1986), is an extremely common element in deepwater habitats in the Caribbean. The species is also known to 

91 m in the Bahamas (Ballantine and Aponte, 2005). A shallowwater collection was reported by Skelton and South (2007) from 

American Samoa as well as by Wynne (2001) from Oman (Arabian Sea).

*Hypoglossum caloglossoides M. J. Wynne et Kraft 

FIGURE 3D

Specimens: DLB7740, 49 m; DLB7149, DLB7189, 50 m; 

DLB7903, 55 m; DLB7569, 70 m; DLB7602, 77 m.

Hypoglossum caloglossoides was reported from 61 m depth 

in the Bahamas by Ballantine and Aponte (2005) and originally 

from Puerto Rico to 70 m depth (Ballantine et al., 2009). The 

holotype of the species, however, was collected at 11 m depth 

(Wynne and Kraft, 1985) and is also known from shallow water 

in the Red Sea and the Mediterranean (Hoffman and Wynne, 

2015).

*Hypoglossum hypoglossoides (Stackh.) Collins et Herv.

Specimens: DLB3691, 36 m; DLB7430, 50 m; DLB1750a, 

61 m; DLB2157a, 87 m.

Hypoglossum hypoglossoides was reported from Bermuda 

at 37 m by Schneider (2000) and at 50–56 m at Campeche 

Banks, Mexico, by Mateo-Cid et al. (2013).

*Hypoglossum rhizophorum D. L. Ballant. et M. J. Wynne 

Specimens: DLB1832, 40 m; DLB7402, DLB7737, 49 m; 

DLB7147, DLB7193, 50 m; DLB7905, 55 m; DLB1718, 

58 m; DLB1750b, 61 m; DLB7486, 63 m; DLB3206, 

68 m; DLB7594, 77 m; DLB2157b, 87 m.

The species was originally described from deepwater habitats in Puerto Rico to 61 m depth (Ballantine and Wynne, 1988). 

Schneider (2000) reported the species from Bermuda to a maximum of 47–50 m.

*Hypoglossum simulans M. J. Wynne, I. R. Price et D. L. Ballant. 

Specimens: DLB3696, 36 m; DLB4728, 60 m; DLB1783, 61 m.

The holotype of H. simulans is from 30 m at Guadeloupe, 

French West Indies (Wynne et al., 1989).

Hypoglossum tenuifolium (Harv.) J. Agardh 

Specimens: DLB1717, 58 m; DLB1743, DLB7772, 61 m; 

DLB2156, 87 m.

Reported to 83 m from Brazil (Cordeiro-Marino and Guimarães, 1981) and from 90 m in Florida by Hanisak and Blair 

(1988a).

*Martensia pavonia Hering 

Specimens: DLB3703, 36 m; DLB7027, 46 m; DLB7364, 

49 m; DLB7318, 50 m; DLB1772, 61 m; DLB7484, 63 m; 

DLB8004, 70 m; DLB7882, 82 m.

Martensia pavonia is another common deepwater algal turf 

component. It was reported to 40 m depth by Hanisak and Blair 

(1988a).

*Myriogramme prostrata (E. Y. Dawson, Neushul et Wildman) 

M. J. Wynne 

FIGURE 3B

Specimens: DLB7051, 52 m; DLB7948, 67 m.

Myriogramme prostrata was originally reported from 

Puerto Rico at 11 m depth by Ballantine et al. (2004).

*Nitophyllum adhaerens M. J. Wynne 

Specimens: DLB7958, 36 m; DLB7032, 46 m; DLB7142, 

DLB7346, 50 m; DLB449, 70 m; DLB7683, 82 m; 

DLB8162, 90 m.

Schneider (2000) reported Nitophyllum adhaerens from Bermuda to 50 m, and Ballantine and Aponte (2005) also reported the 

species from the Bahamas to a maximum depth of 50 m. Wynne 

(1997), in part on the basis of Puerto Rican specimens, reported 

the species from the Bahamas at 61 m depth. The holotype, however, was from shallow water at Quintana Roo, Mexico.

*Taenioma nanum (Kütz.) Papenf.

Specimen: DLB7812, 49 m.

Typically, a shallow-water entity (Taylor, 1960), the species 

has also been collected at the Puerto Rican insular shelf edge at 

21 m depth (Ballantine, unpublished).

Rhodomelaceae

*Amansia multifida J. V. Lamour.

Specimen: DLB8457, 55 m

Amansia multifida is typically a shallow-water alga. Littler 

and Littler (2000) reported that the alga lives from the intertidal 

to a depth of 5 m.

Chondria spp.

Specimens: DLB4204, 36 m; DLB7344, DLB7368, DLB7744, 

49 m; DLB7053, 52 m; DLB7887, 77 m.

These Chondria records probably represent multiple species.

*Herposiphonia secunda (C. Agardh) Ambronn. 

Specimen: DLB7499, 61 m.

The species is a member of the deepwater algal turf in Puerto 

Rico. It has been reported to 31 m in the Bahamas (Ballantine 
NUMBER 105 • 9

FIGURE 3. Deepwater Rhodophyta algal species. (A) Hymenoclonium serpens (DLB0720; scale bar = 100 μm). (B) Myriogramme prostrata

(DLB6029; scale bar = 100 μm). (C) Hypoglossum anomalum (DLB7569; scale bar = 200 μm). (D) Hypoglossum caloglossoides (DLB7569; 

scale bar = 200 μm). (E) Rhododictyon bermudense (DLB7527; scale bar = 1.0 mm). (F) Cresia opalescens (DLB6343; scale bar = 1.0 mm). (G) 

Botryocladia iridescens (DLB6307; scale bar = 2.0 mm). Photos by Hector Ruiz Torres.
1 0 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

and Aponte, 2005) and to 35 m in Bermuda (Schneider and Searles, 1997). The species was also identified from Brazil at 52 m by 

Guimarães et al. (1981).

Herposiphonia sp.

Specimen: DLB7222, 52 m.

*Laurencia intricata J. V. Lamour.

Specimens: DLB4195, 36 m; DLB4684, 55–60 m.

Laurencia intricata is a common shallow-water species (Littler and Littler, 2000), although Taylor (1960) reported that the 

species had been dredged to a depth of 36 m, and Mateo-Cid et 

al. (2013) recorded the species from 40 to 49 m at Campeche 

Banks, Mexico.

*Lophosiphonia cristata Falkenb.

Specimen: DLB8107, 49 m.

Listed by Taylor (1960) as a shallow-water species, L. cristata is a member of the deep-water algal turf in Puerto Rico.

Polysiphonia spp.

Specimens: DLB7285, 46 m; DLB7365, 49 m; DLB7398, 50 m; 

DLB8015, 70 m.

These Polysiphonia records probably represent multiple 

species.

Wrightiella tumanowiczi (Gatty ex Harv.) F. Schmitz

Specimens: DLB7436, DLB7389, 50 m; DLB4807, 55 m.

Puerto Rican inshore plants typically reach to over 30 cm 

in height, whereas the deepwater specimens are highly reduced, 

to 5 cm or less. The species has also been collected to 40 m 

depth offshore North Carolina (Schneider and Searles, 1975) 

and 50–56 m from the Campeche Banks, Mexico (Mateo-Cid 

et al., 2013). 

Sarcomeniaceae

Cottoniella filamentosa (M. Howe) Børgesen

Specimen: DLB3130, 91 m.

Taylor (1960) reported the species to a depth of 92 m.

*Platysiphonia caribaea D. L. Ballant. et M. J. Wynne 

Specimens: DLB3698, 36 m; DLB3713, 46–55 m; DLB1710, 

58 m; DLB1776, 61 m; DLB3199, 80 m; DLB3193, 80–

100 m.

This species was originally reported from Puerto Rico at a depth 

of 18 m as well as from 29 m in Quintana Roo, Mexico (Ballantine 

and Wynne, 1985). The 100 m depth record exceeds the 92 m depth 

reported by Ballantine and Aponte (2005) for the Bahamas.

Wrangeliaceae

Diplothamnion jolyi C. Hoek 

Specimens: DLB3735, 36–46 m; DLB1811, 50 m.

The species is based on collections from Curaçao, Netherlands Antilles, to a depth of 55 m (Hoek, 1978) and is also 

known to 32 m from Bermuda (Schneider and Searles, 1997). 

A previous account of the species from Puerto Rico was based 

on occurrence in wave-exposed habitats in very shallow water 

(Wynne and Ballantine, 1985).

*Grallatoria reptans M. Howe

Specimens: DLB3699, 36 m; DLB7936, 49 m; DLB7052, 52 m; 

DLB7899, 55 m.

Grallatoria reptans is typically a shallow-water alga in areas 

of high turbulence (Taylor, 1960); the species is also reported to 

55 m from Florida (Hanisak and Blair, 1988a). 

*Griffithsia heteromorpha Kütz

Specimens: DLB7372, 49 m; DLB7054, 52 m.

The species was reported by Schneider (2004) to a maximum depth of 37 m in Bermuda, and Hanisak and Blair (1988a) 

reported a collection of this species to 38 m in Florida. 

Haloplegma duperreyi Mont. 

Specimen: DLB7757, 49 m.

Ballantine and Aponte (2005) reported the species to 61 m 

in the Bahamas. The species is also known from Hawaii in deep 

water at 60 m (Agegian and Abbott, 1985).

Rhododictyon bermudense W. R. Taylor 

FIGURE 3E

Specimens: DLB7527, 63 m; DLB7574, 70 m; DLB7763, 

DLB7846, 82 m; DLB2177, DLB8145, 87 m; DLB8164, 

90 m; DLB8147, 91 m.

Ballantine and Aponte (2005) collected this exclusively 

deepwater species at a depth of 107 m in the Bahamas. The species was also reported to 100 m by Frederick (1963).

*Spermothamnion investiens (P. Crouan et H. Crouan) Vickers 

Specimens: DLB4120, 55 m; DLB1714, 58 m; DLB1771, 61 m; 

DLB7854, 82 m; DLB2168, 87 m.

Reported by Taylor (1960) to a depth of 27 m, S. investiens

var. cidaricola Børgesen was reported by Schneider (1976) to a 

maximum depth of 54 m in the Carolinas. 

*Spermothamnion cf. macromeres Collins et Herv.

Specimen: DLB7854, 82 m.

Reported as a shallow-water species by Taylor (1960).

*Spongoclonium caribaeum (Børgesen) M. J. Wynne 

Specimens: DLB3728, 36–46 m; DLB4119, 55 m; DLB2155, 87 m.

As Mesothamnion caribaeum Børgesen, the species was reported 

to 30 m depth in St. John, U.S. Virgin Islands (Børgesen, 1917). 

*Wrangelia bicuspidata Børgesen 

Specimens: DLB7282, 46 m; DLB7360, DLB7711, 49 m; 

DLB7317, DLB8088, 50 m; DLB7070, 52 m; DLB1723, 
NUMBER 105 • 1 1

58 m; DLB4686, 55–60 m; DLB1763, 61 m; DLB7974, 

62 m; DLB7478, 63 m; DLB7953, 67 m; DLB7562, 

DLB7631, 70 m; DLB7593, 77 m; DLB7678, 82 m.

Wrangelia bicuspidata is another common component of 

the deepwater turf community in Puerto Rico. Taylor (1960) reported that the species has been dredged from 40 m depth.

*Wrangelia gordoniae Bucher, D. L. Ballant. et J. N. Norris

Specimens: DLB7035, 46 m; DLB7742, 49 m; DLB7315, 

DLB7426, 50 m; DLB7445, 57 m; DLB4724, 60 m; 

DLB7470, 61 m; DLB7974, 62 m; DLB7825, 73 m.

Bucher et al. (2014), in describing the species in part on 

the basis of the above Puerto Rican collections, reported the 

species to a maximum depth of 67–73 m at San Salvador Island, Bahamas, equaling the depth of the deepest Puerto Rican 

collection.

Gigartinales, Cystocloniaceae

*Hypnea volubilis Searles in C. W. Schneid. et Searles

Specimen: DLB7897, 55 m.

Hypnea volubilis was previously reported from Puerto Rico 

(Ballantine and Aponte, 1997) to a depth of 60 m. It has also 

been reported off the east coast of Florida to 40 m by Eiseman 

(1979).

Kallymeniaceae

*Meredithia? caribaea D. L. Ballant, H. Ruiz et J. N. Norris 

Specimens: DLB7947, 67 m; DLB7579, 70 m.

Meredithia? caribaea, only recently described from offshore 

habitats in Puerto Rico (Ballantine et al., 2015), is known from 

two deepwater collections as well as from a midshelf algal plain 

at 17 m. 

*Meredithia pulchella D. L. Ballant, H. Ruiz et J. N. Norris

Specimens: DLB7308, 50 m; DLB7590, DLB7639, 70 m.

The entity first reported from Puerto Rico between 14 and 

70 m by Ballantine et al. (2011) was referred to Kallymenia limminghei Montagne. Schneider et al. (2014) indicated that on the 

basis of molecular evidence, the Puerto Rican entity was an undescribed species of the genus Meredithia. The species was subsequently described by Ballantine et al. (2015).

Rhizophyllidaceae

Contarinia sp. 

Specimens: DLB7921, 49 m; DLB7839, 5 m; DLB7892, 55 m; 

DLB7950, DLB7956, 67 m; DLB7813, 73 m; DLB7891, 

DLB8186, 82 m.

This noncalcified crust was common in the Puerto Rican 

deepwater collections and was sequenced by G. W. Saunders’ lab 

(University of New Brunswick, Fredericton, Canada) and listed 

as Contarinia sp. in Schneider et al. (2014).

Solieriaceae

*Agardhiella ramosissima (Harv.) Kylin 

Specimen: DLB4683, 55–60 m.

Agardhiella ramosissima has been reported as dredged to 

55 m by Taylor (1960).

*Flahaultia tegetiformans W. R. Taylor 

Specimens: DLB1709, 36 m; DLB1258, DLB1825, 40 m; 

DLB7440, 49 m; DLB1765, 61 m.

The species was previously reported from Puerto Rico by 

Ballantine and Aponte (1997) to a maximum depth of 61 m. A 

dredged specimen of F. tegetiformans from Jamaica was reported 

to 24 m depth by Taylor (1974).

Meristotheca gelidium (J. Agardh) E. J. Faye et Masuda

Specimen: DLB4188, 36 m.

Frederick (1963) reported the species as Eucheuma schrammii (P. et H. Crouan) J. Agardh from 64 m depth in Bermuda.

Solieria filiformis (Kütz.) P. W. Gabrielson

Specimen: DLB4209, 36 m.

Schneider and Searles (1991) reported that the species has 

been dredged from 29 to 45 m at Onslow Bay, North Carolina.

*Wurdemannia miniata (Spreng.) Feldmann et Hamel 

Specimen: DLB7832, 52 m; DLB7532, 63 m.

A member of the deepwater turf community in Puerto Rico, 

the species is typically known as being from shallow water (Taylor, 1960), although Hanisak and Blair (1988a) reported W. miniata from 26 to 58 m in Florida.

Nemastomatales, Nemastomataceae

*Predaea laciniosa Kraft

Specimens: DLB7505, DLB7520, 61 m; DLB7472, 63 m; 

DLB7778, 70 m.

The species as known from Puerto Rico is based on the 

above collections (Ballantine et al., 2009). The Puerto Rico report remains the species’ only occurrence in the Atlantic Ocean. 

Predaea laciniosa was originally described from the Great Barrier Reef, Australia, with the holotype having been collected at 

12 m depth (Kraft, 1984). The species is otherwise broadly reported from the Pacific, to 10 m in Hawaii (Abbott, 1999), as 

well as from the Indian Ocean, with depth records of 21 m in 

Oman and Yemen (Schils and Coppejans, 2002).

Predaea sp. 

Specimen: DLB7686, 64 m.

Schizymeniaceae

*Titanophora incrustans (J. Agardh) Børgesen 

Specimen: DLB4698, 60 m.
1 2 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

Littler and Littler (2000) indicated a depth range to 15 m 

for the species, and Mateo-Cid et al. (2013) reported Titanophora incrustans from 50 to 56 m at Campeche Banks, Mexico.

Gracilariales, Gracilariaceae

*Gracilaria isabellana Gurgel, Frederica et J. N. Norris 

Specimens: DLB3751, 36–46 m; DLB4124, 55 m; DLB4687, 

55–60 m; DLB4735b, 60 m.

The species is based on shallow-water collections in St. Croix, 

U.S. Virgin Islands (Gurgel et al., 2004).

Peyssonneliales, Etheliaceae

Ethelia sp.

FIGURE 2G

Specimens: DLB7529, 63 m; DLB7573, 70 m; DLB7814, 73 m; 

DLB7858, 82 m.

These specimens are recognized to represent an undescribed 

species, and their description is pending. 

Peyssonneliaceae

*Peyssonnelia boergesenii Weber Bosse

Specimens: DLB7401, 49 m; DLB7323, 50 m; DLB7471, 

61 m.

Peyssonnelia boergesenii is more typically a shallow-water 

alga (Taylor, 1960; Guimarães and Fujii, 1999). It had previously 

been reported from 0.5 and 22 m in Puerto Rico (Ballantine and 

Ruiz, 2005).

*Peyssonnelia flavescens D. L. Ballantine et H. Ruiz 

FIGURE 2B

Specimens: DLB7166, 50 m; DLB8053, 70 m; DLB7819, 73 m; 

DLB7872, 82 m.

The original and only report of the species is from Puerto 

Rico by Ballantine and Ruiz (2005).

*Peyssonnelia gigaspora D. L. Ballant. et H. Ruiz

FIGURE 2A

Specimens: DLB7830, 52 m; DLB7468, DLB7521, 61 m; 

DLB7477, 63 m; DLB7943, DLB7945, 67 m; DLB7612, 

DLB7637, DLB7907, 70 m; DLB7816, 73 m; DLB7859, 

DLB7890, 82 m; DLB8160, 90 m; DLB8150, 91 m.

The original and only report of the species is from Puerto 

Rico by Ballantine and Ruiz (2010). The species is very common 

in deepwater coral reef habitats.

Peyssonnelia inamoena Pilg. 

Specimens: DLB7278, 46 m; DLB1229, 49 m; DLB7817, 73 m.

Ballantine and Aponte (2005) reported the species to 91 m 

depth in the Bahamas, and Hanisak and Blair (1998a) reported 

the species in Florida to 98 m.

*Peyssonnelia incomposita D. L. Ballant. et H. Ruiz 

FIGURE 2D

Specimens: DLB7830, 52 m; DLB7987, 62 m; DLB7777, 

DLB8039, 70 m; DLB7818, 73 m; DLB7859, 82 m.

The original and only report of the species is from Puerto 

Rico by Ballantine and Ruiz (2011).

*Peyssonnelia iridescens D. L. Ballant. et H. Ruiz 

FIGURES 2C, 4C

Specimens: DLB7277, 46 m; DLB7116, DLB 7759, 49 m; 

DLB7316, DLB7329, 50 m; DLB7518, DLB9769, 61 m; 

DLB7547, 62 m; DLB7474, DLB7530, 63 m; DLB7687, 

64 m; DLB7558, DLB7914, 70 m; DLB7609, 77 m; 

DLB7670, DLB7855, 82 m; DLB8143, 87 m.

The original and only report of the species is from Puerto 

Rico by Ballantine and Ruiz (2010). Because of its iridescence 

and loosely attached habit, the species is a visually conspicuous 

element of the Puerto Rican mesophotic community.

Peyssonnelia sp. 1

Specimens: DLB7440, 49 m; DLB7334, 50 m; DLB7454, 57 m; 

DLB7986, 62 m; DLB8043, 70 m.

The crust possesses hypothallial filaments that do not laterally cohere.

Peyssonnelia sp. 2 

FIGURE 2F

Specimens: DLB7286, 46 m; DLB7727, 49 m; DLB7444, 57 m; 

DLB7519, DLB7522, 61 m; DLB7479, 63 m.

Peyssonnelia sp. 3 

FIGURE 2E

Specimens: DLB7548, 62 m; DLB7570, DLB7606, DLB7624, 

DLB8042, 70 m; DLB7865, 82 m.

This species as well as the other above Peyssonnelia species 

are reasonably abundant on the deep Puerto Rican insular shelf; 

unfortunately, reproductive specimens were never collected for 

any of these taxa. They all probably represent additional undescribed species.

*Polystrata fosliei (Weber Bosse) Denizot 

FIGURE 2H

Specimen: DLB7912, 70 m.

Ballantine et al. (2011) reported this species from Puerto 

Rico in deep water and at 17 m depth. Kato et al. (2006) reported the species from Japanese coastal waters to be a species of 

shallow to intermediate depths.

Halymeniales, Halymeniaceae

Cryptonemia crenulata (J. Agardh) J. Agardh 

Specimens: DLB3701, 36 m; DLB1208, 40 m; DLB7026, 46 m; 

DLB436, DLB7756, 49 m; DLB7836, 52 m; DLB7452, 
NUMBER 105 • 1 3

FIGURE 4. Bottom habits at edge of insular shelf south of La Parguera (all 1 m2). (A) “Weinberg,” 47 m. Pinkish background growths are Corallinales, mostly Hydrolithon abyssophila (arrows); also present are leafy Lobophora (probably L. guadeloupensis) (arrow heads) and several small 

Halimeda sp. in the lower right; orange growths are sponges; and at the far upper left is a colony of Agaricia lamarcki Milne Edwards et Haime. 

(B) “Hole in the wall,” 47 m. The frame is dominated by a brown sponge (bottom); the large yellow incrustation at center is Peyssonnelia flavescens; algal turf (including Wrangelia spp.) is indicated by arrows; and leafy Lobophora (probably L. guadeloupensis) is shown by the arrowhead. 

(C) “Hole in the wall,” 67 m. Nearly free blades of Peyssonnelia iridescens dominate the center of the frame (arrows); crustose coralline algae 

(some covered with sediment) appear as pink encrustations; the deep maroon encrustation at lower left is Peyssonnelia sp.; and leafy Lobophora 

(probably L. guadeloupensis) is shown by the arrowhead. (D) El Hoyo Terrace, 59 m. Coralline algae are pinkish encrustations; maroon encrustations (arrow) are Peyssonnelia sp.; leafy Lobophora (probably L. guadeloupensis) are shown by arrowheads; and Halimeda cryptica is indicated by double arrowheads. (E) “Hole in the wall,” 76 m. Bottom is dominated by encrusting coralline red algae, Peyssonnelia sp., and sponges. 

(F) “El Precipicio,” 70 m. Bottom is similarly dominated by encrusting coralline red algae, Peyssonnelia sp. (arrow), and sponges; Halimeda 

cryptica is indicated by a double arrowhead; and a portion of an Agaricia plate is in the upper left corner. Photos by Hector Ruiz Torres.
1 4 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

57 m; DLB7525, 63 m; DLB8003, 70 m; DLB7623, 70 m; 

DLB7875, 82 m.

Ballantine and Aponte (2005) reported the species to 61 m 

in the Bahamas, and Hanisak and Blair (1988a) reported the species to 93 m from Florida.

Cryptonemia sp.

Specimen: DLB1800, 50 m.

Halymenia hancockii W. R. Taylor

Specimen: DLB8458, 55 m.

The species was described from dredged material at 24 m 

depth in Colombia (Taylor, 1942). Frederick (1963) reported the 

species to 64 m from Bermuda, and Hanisak and Blair (1988a) 

recorded the species from 34 to 93 m off the east coast of Florida.

Halymenia pseudofloresii Collins et M. Howe

Specimens: DLB4206, 36 m; DLB4692, 55–60 m.

Halymenia pseudofloresii was reported to 60 m (as Halymenia floresii (Clemente) Agardh) by Leichter et al. (2008) from 

Florida and to the same depth from Bermuda by Frederick 

(1963). Dawes and Van Breedveld (1969) also reported the species (as H. floresii) from Florida at 73 m. 

Rhodymeniales, Champiaceae

Champia parvula (C. Agardh) Harv.

Specimens: DLB7314, DLB7381, 50 m; DLB7056, 52 m; 

DLB7509, 61 m; DLB8071, 70 m.

Taylor (1960) indicated that the species was primarily restricted to shallow water; however, he also reported that dredged 

material had been collected to 37 m. Gilmartin (1960) reported 

C. parvula from 63 m in Eniwetok, and Joly and Yoneshigue 

Braga (1966) reported the species from 120 m in Brazil. LozadaTroche and Ballantine (2010) indicated that specimens identified as C. parvula in the western Atlantic may represent multiple 

cryptic species.

Champia vieillardii Kütz. 

Specimens: DLB1253, 40 m; DLB7377, DLB7928, 49 m; 

DLB7338, DLB7404, 50 m; DLB7081, 55 m.

Ballantine and Aponte (2005) reported the species to 61 m 

in the Bahamas. The species has also been reported from 55 to 

60 m at St. Paul Archipelago (Mid-Atlantic Ridge, Brazil) by 

Magalhães et al. (2015).

Faucheaceae

*Gloiocladia atlantica (Searles) R. E. Norris 

Specimens: DLB7464, 41 m; DLB7933, 40 m; DLB7151, 50 m; 

DLB7553, 62 m; DLB7556, 70 m; DLB3121, 100 m.

Originally reported from Puerto Rico (as Gloioderma atlanticum Searles) by Ballantine and Norris (1989) from a maximum 

depth of 36 m, Gloiocladia atlantica is relatively rarely reported 

in the tropical western Atlantic. Hanisak and Blair (1988a) reported the species from a maximum depth of 60 m in Florida, 

and Ballantine and Aponte (2005) reported the species to 76 m 

from the Bahamas.

Hymenocladiaceae

Asteromenia peltata (W. R. Taylor) Huisman et A. Millar 

Specimens: DLB3692, 36 m; DLB4689, 55–60 m; DLB4719, 60 m.

Reported from the Gulf of Mexico to 50–65 m by Gavio 

and Fredericq (2005) and to 64 m (as Halichrysis peltata (W. R. 

Taylor) P. Huvé et H. Huvé) in the Canary Islands by Haroun 

et al. (1993). Dawes and Van Breedveld (1969) reported the 

species as Fauchea peltata W. R. Taylor from Florida to 73 m. 

The deepest known report for the species is by Ballantine 

and Aponte (2005), who reported A. peltata to 76 m in the 

Bahamas. 

Lomentariaceae

Lomentaria sp. 

Specimen: DLB7378, 49 m.

Lomentaria divaricata (Durant.) M. J. Wynne was reported 

(as L. baileyana (Harv.) Farlow) to 71 m by Hanisak and Blair 

(1988a).

Rhodymeniaceae

*Botryocladia iridescens D. L. Ballant. et H. Ruiz

FIGURE 3G

Specimens: DLB7036, 47 m; DLB7335a, 50 m; DLB7563, 

70 m; DLB7853, 82 m.

The species was originally reported from moderate (23 m) 

depth habitats in Puerto Rico and Grenada (Ballantine and Ruiz, 

2008) and has not been elsewhere reported.

Botryocladia pyriformis (Børgesen) Kylin 

Specimens: DLB4674, 55–60 m; DLB4714, 60 m.

Botryocladia pyriformis was reported to 60 m in Florida 

by Leichter et al. (2008) and to 70 m by Gavio and Fredericq 

(2005). The deepest depth report for the species is 73 m by Frederick (1963) at Bermuda.

*Botryocladia spinulifera W. R. Taylor et I. A. Abbott 

Specimens: DLB1251, 40 m; DLB7037, 47 m; DLB1218, 

DLB7748, 49 m; DLB7326, 50 m; DLB7944, 67 m; 

DLB7618, 70 m; DLB7876, 82 m.

The species was originally reported from Puerto Rico by 

Ballantine (1985) to a maximum depth of 49 m as well as by 

Bucher et al. (1990) to a maximum depth of 40–49 m in Florida. 

Mateo-Cid et al. (2013) reported the species from 50 to 56 m at 

Campeche Banks, Mexico.
NUMBER 105 • 1 5

*Botryocladia wynnei D. L. Ballant.

Specimen: DLB7475, 63 m.

The type locality for the species is a 17-m-deep algal plain in 

Puerto Rico (Ballantine, 1985). Schneider and Searles (1991) reported the species from 30 to 35 m depth at Onslow Bay, North 

Carolina.

Chrysymenia cf. ventricosa (J. V. Lamour.) J. Agardh 

Specimen: DLB 8422, 37 m.

Ballantine et al. (2015) reinstated the presence of the species in Puerto Rico on the basis of a collection at 37 m. Taylor 

(1960:479) indicated that C. ventricosa was “probably a deepwater species” that has been dredged to 90 m, and Rodriguez y 

Femenías (1888) reported the species growing at 130 m at Puerto 

de Mahón (Menorca, Spain) in the Mediterranean.

Chrysymenia sp.

Specimen: DLB7403, 50 m.

Coelarthrum cliftonii (Harv.) Kylin 

Specimens: DLB7932, 49 m; DLB7339, DLB7425, 50 m; 

DLB450, 70 m.

Gavio and Fredericq (2005) also collected the species from 50 

to 70 m in the Gulf of Mexico, and Frederick (1963) reported the 

species to 73 m depth in Bermuda. The deepest known depth for 

the species is 76 m in the Bahamas (Ballantine and Aponte, 2005).

*Cresia opalescens D. L. Ballant., Lozada-Troche et H. Ruiz 

FIGURE 3F

Specimen: DLB7729, 49 m.

To date, Cresia opalescens is known only from its type locality in Puerto Rico, where it is most frequently collected in coral 

reef habitats at moderate depths (Lozada-Troche et al., 2010).

*Halichrysis corallinaria D. L. Ballant., G. W. Saunders et H. Ruiz 

Specimen: DLB7133, 50 m.

Halichrysis corallinaria was reported from Puerto Rico to a 

maximum depth of 23 m by Ballantine et al. (2007) and has not 

been collected elsewhere.

Leptofauchea? rhodymenioides W. R. Taylor 

Specimens: DLB7747, DLB7927, 49 m; DLB7305, DLB7328 

50 m; DLB7837, 52 m; DLB 7451, 57 m; DLB7590, 70 m; 

DLB7766, 82 m.

As no reproductive material was collected, this record is unconfirmed. Leptofauchea rhodymenioides was reported to 93 m 

by Hanisak and Blair (1988a) and 95 to 155 m in the Gulf of 

Mexico by Gavio and Fredericq (2005).

Other Rhodophyta

*Hymenoclonium serpens (P. Crouan et H. Crouan) Batters 

FIGURE 3A

Specimens: DLB7833, 52 m; DLB7453, 57 m; DLB2153, 87 m.

This species was first reported for Puerto Rico by Ballantine and Norris (1989), following the listing of Taylor (1960) 

as a species of uncertain assignment. The Puerto Rican entity is 

occasionally seen in water as shallow as 20 m depth. It forms 

prostrate crusts with elongate, branched vein-like cells to 53 

μm in length. Small irregularly shaped cells fill in the crust 

between the branched axes (see also Ballantine and Norris, 

1989:6, fig. 9). 

The species was originally described as Callithamnion serpens P. Crouan et H. Crouan (1859) from material dredged at 

Brest, France, at 20 m depth and later transferred to the genus 

Hymenoclonium by Batters (1895), who noted that his material 

was “dredged from deep water” in Plymouth Bay. Hymenoclonium serpens is now considered to be the tetrasporophytic life 

history alternate of Bonnemaisonia asparagoides (Woodward) 

C. Agardh, although Åsen (1980) reported the entity growing 

directly from its basophyte. The only two members of the Bonnemaisoniales in the western Atlantic (Wynne, 2011) are Asparagopsis taxiformis (Delile) Trevis. and Bonnemaisonia hamifera

Harv. The tetrasporophytic life history alternate of the former in 

the western Atlantic is well known to be the filamentous Falkenbergia hillebrandii (Bornet) Falkenberg. The life history alternate 

of the latter is Trailliella intricata Batters (1896).

Other possible relationships with Hymenoclonium include the life history alternate of Acrosymphyton purpuriferum

(J. Agardh) Sjöstedt. Cortel Breeman (1975) provided evidence 

that a H. serpens–like prostrate filamentous form represented 

the tetrasporophyte of A. purpuriferum. Although A. purpuriferum is not known from Puerto Rico, A. caribaeum (J. Agardh) 

Sjözsedt has been reported from the island (Ballantine and 

Aponte, 2002b). Another possibility is the carpospore germination product of Meredithia microphylla (J. Agardh) J. Agardh. 

Guiry and Maggs (1984), in investigating the life history of 

that species, reported that the tetrasporophyte germinated and 

formed a complex system of prostrate filaments (figs. 11–21) resembling the above-cited fig. 9 in Ballantine and Norris (1989). 

The genus Meredithia is represented in Puerto Rico by the abovelisted deepwater species M.? caribaea and M. pulchella.

Heterokontophyta

Phaeophyceae, Dictyotales, Dictyotaceae

Canistrocarpus cervicornis (Kütz.) De Paula et De Clerck

Specimens: DLB7743, 49 m; DLB7159, 50 m.

Ballantine and Aponte (2003) reported the species from the 

Bahamas as Dictyota cervicornis Kütz. from 77 m.

*Dictyopteris delicatula J. V. Lamour. 

Specimens: DLB3704, 36 m; DLB7029, 46 m; DLB7111, 49 m; 

DLB7333, 50 m; DLB7500, 61 m; DLB7561, DLB7630, 

DLB8006, 70 m; DLB7598, 77 m; DLB7878, 82 m.
1 6 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

This alga is extremely common in shallow-water coral reef 

environments throughout the Caribbean. It is also very common in deepwater turf communities and was previously reported to a depth of 61 m in the Bahamas by Ballantine and 

Aponte (2003) and to 74 m by Macintyre et al. (1991) from 

Barbados.

Dictyopteris justii J. V. Lamour. 

Specimens: DLB3716, 36–46 m; DLB4114, 55 m; DLB4717, 

60 m.

Dictyopteris justii was reported by Taylor (1960) to range 

from the intertidal to 40 m depth. It has also been reported from 

46 m depth in the Bahamas (Ballantine and Aponte, 2003). The 

deepest known record of the species is 110 m reported by Guimarães et al. (1981) from Brazil.

Dictyota bartayresiana J. V. Lamour.

Specimen: DLB7775, 61 m.

In the western Atlantic, Dictyota bartayresiana was also 

reported to 61 m by Ballantine and Aponte (2003) from the 

Bahamas and to 60 m from Brazil by Yoneshigue-Valentin et 

al. (2004). The deepest known depth record for the species is 

118 m from Johnston Atoll, Pacific Ocean (Agegian and Abbott, 1985).

Dictyota ciliolata Sond. ex Kütz.

Specimen: DLB3749, 36–46 m.

Dictyota ciliolata was reported by Taylor (1960) to generally be an alga of shallow water, ranging to 24 m depth, but 

was reported to 50 m by Schneider (1976). Suárez et al. (2015) 

reported that D ciliolata is known to a depth of 55 m; however, 

the depth is reported without citation or location.

*Dictyota humifusa Hörnig, Schnetter et Coppejans 

Specimens: DLB7028, 46 m; DLB7039, 46 m; DLB7138, 50 m; 

DLB7482, 63 m; DLB7572, 70 m; DLB7884, 82 m.

This alga is easily recognized because of its blue and green 

iridescent bands and is extremely abundant at the southwestern 

Puerto Rican insular shelf break in approximately 18 m of water. 

De Clerck (2003) indicated that the species typically lives in the 

shallow subtidal in the Indian Ocean.

*Dictyota jamaicensis W. R. Taylor

Specimen: DLB3719, 36–46 m.

Taylor (1960) reported that the species was limited to shallow water.

Dictyota pulchella Hörnig et Schnetter 

Specimens: DLB7160, DLB8090, 50 m; DLB7485, 63 m; 

DLB7504, 61 m; DLB8064, 70 m.

Dictyota pulchella was reported to a depth of 76 m in the 

Bahamas by Ballantine and Aponte (2003). This species, as 

D. divaricata Lamour., although not appearing in his species list, 

was nevertheless reported by Frederick (1963) as the most abundant brown alga on the Bermuda platform to 63 m.

Dictyota stolonifera E. Y. Dawson

Specimens: DLB7989, 62 m; DLB8063, 70 m.

The species was first reported from Puerto Rico and the 

Caribbean region by Ballantine et al. (2011). De Clerck (2003) 

indicated that the species ranged from shallow depths to intermediate (to 20 m) throughout its distribution, although Spalding 

(2012) reported D. stolonifera to 93 m in Hawaii.

*Lobophora canariensis (Sauv.) C. W. Vieria, De Clerck et Payri.

Specimens: DLB4179, 37–38 m; DLB1817, 40 m; DLB1798, 

DLB8082, 50 m; DLB3090, 55–90 m; DLB7491, 61 m; 

DLB7973, 62 m; DLB7868, 82 m.

For virtually every deep-water collection on which this report is based, Lobophora specimens were obtained. Until the 

work reported by Shultz et al. (2015), all of these Caribbean 

collections were assumed to be Lobophora variegata. On reexamination of deepwater herbarium specimens in MSM identified as L. variegata, both L. canariensis and L. guadeloupensis

(below) proved to be abundant. Shultz et al. (2015) described 

as a new segregate species Lobophora payriae N. E. Schultz, C. 

W. Schneid. et F. Rousseau on the basis of algae collected in Bermuda to depths of 35–36 m and from Guadeloupe. Vieira et al. 

(2016) subsequently regarded L. payriae as a synonym of Cutleria canariensis (Sauvageau) I. A. Abbott et J. M. Juisman and 

transferred the species to Lobophora. The present report is the 

first known occurrence in Puerto Rico. Some of the Puerto Rican 

specimens were extremely small, with flabella measuring 9.0 mm 

high and 10 mm across; larger specimens measured to 4 cm high 

and 5 cm broad. All were 3 cell layers thick and measured 33 to 

40 μm thick. Further examination of herbarium specimens will 

probably reveal that L. canariensis is not restricted to depths 

greater than 35 m in Puerto Rico.

Lobophora guadeloupensis N. E. Schultz, F. Rousseau et L. 

Le Gall

Specimens: DLB1817, 40 m; DLB3717, 40–50 m; DLB7136, 

49 m; DLB4130, 56 m; DLB4716, 60 m; DLB8055, 70 m.

Lobophora guadeloupensis represents a new species record for Puerto Rico. Flabella measured to 5.5 cm high and to 

8.0 cm across. In section, Puerto Rican specimens possessed a 

single medullary layer with two cortical rows both above and 

below, averaging 82 μm in thickness. In radial section, the cortical cells located immediately outside medullary cells are covered 

by two elongate outer cortical cells. Collections at MSM also 

reveal that L. guadeloupensis is found in Martinique (DLB3546) 

at 50 m and the Bahamas (DLB4341 and DLB4518) at 33–35 

and 46–53 m, respectively. The greatest known depth for the 

species is reported to be 135 m from Guadeloupe (Schultz et 

al., 2015). Once herbarium records attributable to L. variegata

in deep water are reexamined, the broader occurrence of both 
NUMBER 105 • 1 7

L. guadeloupensis and L. payriae will undoubtedly be confirmed, 

and the depth records reported here will be exceeded. 

Lobophora variegata (J. V. Lamour.) Womersley ex E. C. Oliveira 

Specimen: DLB4031, 36–37 m.

The species, as originally circumscribed, was broadly distributed worldwide. Vieira et al. (2014), employing molecular tools, 

concluded that species diversity in Pacific Lobophora was underestimated, and they suggested that Atlantic species required reexamination as well. In Puerto Rico, until the recent revision by Shultz 

et al. (2015), Lobophora variegata was thought to be a very common element from shallow nearshore environments and extending 

across the insular shelf into deeper water. It has been considered to 

have a broad worldwide tropical distribution; however Vieiria et 

al. (2016) concluded that it is restricted to the Caribbean.

Historical records of Lobophora variegata from deepwater 

must now be accepted with caution. The species was reported 

to a depth of 76 m in the Bahamas by Ballantine and Aponte 

(2003), although unattached thalli were regularly observed 

deeper than 100 m (DLB and NEA, unpublished observations). 

The species is also known from 90 m in San Salvador Island, 

Bahamas (Hanisak and Blair, 1988b). Littler and Littler (2000) 

further reported that species has been collected to 120 m. Although Agegian and Abbott (1985) reported a maximum depth 

of 140 m for the species at Johnston Atoll, Pacific Ocean, Spalding (2012) regarded these reports as probably being Distromium 

flabellata Womersley. In light of Shultz et al.’s (2015) treatment, 

we have confirmed only a single specimen deeper than 35 m to 

be L. variegata. Nevertheless, L. variegata is still probably abundant in shallower mesophotic environments.

Padina cf. sanctae-crucis Børgesen 

Specimen: DLB4441, 37 m.

Reported by Taylor (1960) as a plant of shallow water; 

however, Dawes and Van Breedveld (1969) reported the species 

from 73 m in Florida.

Stypopodium zonale (J. V. Lamour.) Papenf.

Specimens: DLB7141, DLB7176, 50 m.

Frederick (1963) reported the species to 64 m in Bermuda, 

and Littler and Littler (2000) reported a maximum depth of 80 m.

Sphacelariales, Sphacelariaceae

Sphacelaria sp. 

Specimen: DLB8019, 70 m.

Puerto Rican deepwater specimens were without vegetative 

propagules.

Ectocarpales, Scytosiphonaceae

Rosenvingea sp.

Specimen: DLB1215, 40 m.

The three known species of the genus from the Caribbean are all known from shallow water, although R. intricata

Børgesen is also known from a dredge collection at 35 m depth 

(Taylor, 1960).

Sporochnales, Sporochnaceae

Sporochnus bolleanus Mont.

Specimens: DLB3723, 36–46 m; DLB4117, 55 m.

Sporochnus bolleanus was reported to 59 m by Hanisak and 

Blair (1988a) and to 90 m by Taylor (1960).

Fucales, Sargassaceae

*Sargassum filipendula C. Agardh

Specimen: DLB4118, 55 m.

Sargassum filipendula was reported by Taylor (1960) to 

a depth of 33 m. Schneider (1975) also reported the species to 

55 m from the North Carolina continental shelf. 

Sargassum hystrix J. Agardh

Specimen: DLB8046, 70 m.

Littler and Littler (2000) reported that the species had been 

dredged to 137 m. This specimen may very well have been unattached and drifted from shallower water.

*Sargassum ramifolium Kütz.

Specimen: DLB4042, 36 m.

Littler and Littler (2000) indicated a shallow-water distribution for the species.

Sargassum sp.

Specimen: DLB7163, 50 m.

Chlorophyta

Palmophyllophyceae, Palmophyllales, Palmellopsidaceae

*Verdigellas fimbriata D. L. Ballant. et J. N. Norris

Specimen: DLB3711, 46–55 m; DLB3606, 55–65 m.

Verdigellas fimbriata, the generitype, had been collected 

only in deepwater habitats in Puerto Rico (Ballantine and Norris, 1994) prior to the report by Moura (2010) from Brazil.

Verdigellas peltata D. L. Ballant. et J. N. Norris 

FIGURE 5A

Specimen: DLB3710, 46–55 m; DLB3605, 55–65 m; DLB3189, 

80–100 m.

Ballantine and Aponte (2003) collected the species to 122 m 

from a submersible in the Bahamas. It is also known from a 

depth of 97 m in Florida (Hanisak and Blair, 1988a). Littler and 

Littler (2000) indicated a maximum depth of 157 m for the species, and Bravin et al. (1999) reported the species from Brazil at 
1 8 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

FIGURE 5. Deepwater Chlorophyta species. (A) Verdigellas peltata (DLB8029, DLB8029; scale bar = 1.0 cm). (B) 

Microdictyon boergesenii (DLB7422; scale bar = 5 mm). (C) Halimeda copiosa (DLB7502; scale bar = 1.0 cm). (D) 

Halimeda pumila (DLB7867; scale bar = 1.0 cm). (E) Rhipiliopsis profunda (DLB3607; scale bar =1.0 cm). (F) Udotea 

unistratea (DLB8047; scale bar = 1.0 cm). (G) Phyllodictyon pulcherrimum (DLB8056; scale bar = 1.0 cm). Photos by 

Hector Ruiz Torres.
NUMBER 105 • 1 9

a depth of 110 m. Although most collections are in water deeper 

than 50 m, Schneider et al. (2010) has reported the species from 

2 to 3 m in a shaded grotto in Bermuda.

Chlorophyceae, Ulvales, Gayraliaceae

*Gayralia oxysperma (Kütz.) K. L. Vinogr. ex Scagel, P. W. Gabrielson, Garbary, Golden, M. W. Hawkes, S. C. Lindstrom, 

J. C. Oliveira et Widd. 

Specimen: DLB7847, 82 m.

A few very small specimens were collected at the edge of the 

Puerto Rican insular shelf. In deepwater, small individuals were 

also reported in 49 and 60 m of water by Frederick (1963) from 

Bermuda.

Phaeophilaceae

*Phaeophila dendroides (P. Crouan et H. Crouan) Batters

Specimen: DLB2170, 65 m.

Reported from shallow water in the Florida Dry Tortugas 

by Taylor (1928). This deepwater specimen was identified by Dr. 

Ruth Nielsen.

Cladophorales, Anadyomenaceae

Anadyomene lacerata D. Littler et Littler 

Specimens: DLB8111, DLB3391, 40 m; DLB8465, 46 m;

DLB4677, 55–60 m; DLB7977, 62 m.

The species is based on a collection to 40 m at Monito Island in Puerto Rico (Littler and Littler, 1991). It is also known 

from 65 m in Brazil (Alves et al., 2011). The deepest known 

record for the species is also from Brazil at 180 m (YoneshigueValentin et al., 2004). 

Anadyomene saldanhae A. B. Joly et E. C. Oliveira 

Specimens: DLB3690, 36 m; DLB7957, 49 m; DLB3712, 

46–55 m; DLB3203, 68 m; DLB8182, 83 m; DLB3124, 

100 m.

Anadyomene saldanhae is a common member of deepwater 

turfs in mesophotic coral reef habitats. The species was originally 

described from Brazilian material, being dredged from 79 m (Joly 

and De Oliveira Filho, 1969 [1968]) and subsequently reported 

from Brazil to 91 m by Alves et al. (2011) as well as to 180 m by 

Yoneshigue-Valentin et al. (2004).

Anadyomene stellata (Wulfen) C. Agardh 

Specimens: DLB7331, DLB7174, 50 m; DLB4116, 55 m; 

DLB4733, 60 m; DLB3202, 68 m; DLB7581, 70 m; 

DLB7665, 82 m; DLB3190, 80–100 m; DLB3114, 100 m.

Although it is generally considered to be a shallow-water 

alga, Ballantine and Aponte (2003) collected the species to 46 m 

in the Bahamas, and Littler and Littler (1991) observed small 

clumps of the species to 91.5 m depth. The species is also known 

to 100 m in Bermuda (Frederick, 1963) and to about 160 m by 

Guimarães et al. (2008) and 180 m by Yoneshigue-Valentin et al. 

(2004) from Brazil.

Microdictyon boergesenii Setch. 

FIGURE 5B

Specimens: DLB1242, 49 m; DLB7422, 50 m; DLB4678, 

55–60 m; DLB4697, DLB7465, DLB7773, 61 m; 

DLB8040, 70 m; DLB7764, DLB7845, 82 m; DLB3120, 

100 m.

Microdictyon boergesenii is another very common element 

in the deepwater algal turf community in Puerto Rico, as well as 

in the Bahamas, where Ballantine and Aponte (2003) reported 

it to 77 m. Taylor (1928) reported the species to 159 m from 

Florida, and Littler and Littler (2000) reported the species from 

a depth of 160 m in the Caribbean region.

Siphonocladales, Boodleaceae

Cladophoropsis sp.

Specimens: DLB8033, 70 m; DLB7863, 82 m.

Schneider and Searles (1975) reported another Cladophoropsis species, C. membranacea, to 33 m on the Carolina continental shelf.

Phyllodictyon anastomosans (Harvey) Kraft et M. J. Wynne

Specimen: DLB3697, 36 m.

Phyllodictyon anastomosans is typically a species of shallow 

water in Puerto Rico and the Caribbean. However, the species is 

reported from 50 to 56 m in Campeche Banks, Mexico (MateoCid et al., 2013), and (as Struvea anastomosans (Harvey) Picc. 

et Grunov ex Picco.) to 61 m by Norris and Olsen (1991) from 

the Bahamas.

Phyllodictyon pulcherrimum J. E. Gray 

FIGURE 5G

Specimens: DLB8075, 50 m; DLB4715, 60 m; DLB8056, 70 m.

As pointed out by Norris and Olsen (1991), the species is 

typically a common shallow-water element. Nevertheless, they 

reported the species to 85 m in the Bahamas. As Struvea pulcherrima (J. E. Gray) Murray et Boodle, Dawes and Van Breedveld 

(1969) reported collections to 73 m from Florida, and Humm 

and Cerame-Vivas (1964) dredged the species from 60 to 120 m 

in the Gulf of Mexico and North Carolina.

Struvea elegans Børgesen 

Specimens: DLB7284, 46 m; DLB1224, 47 m; DLB4691, 

55–60 m; DLB1766, 61 m; DLB7578, 70 m.

Struvea elegans is mostly restricted to deepwater habitats in 

the western Atlantic. From the Bahamas, it has been collected to 

76 m (Ballantine and Aponte, 2003).
2 0 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

Cladophoraceae

*Cladophora coelothrix Kütz. 

Specimen: DLB7861, 82 m.

Norris and Olsen (1991) reported the species to 76 m at San 

Salvador, Bahamas; Ballantine and Aponte (2003) also reported 

Cladophora coelothrix from the Bahamas at 82 m at Lee Stocking Island, Bahamas.

Siphonocladaceae

Valonia macrophysa Kütz.

Specimen: DLB7857, 82 m.

Ballantine and Aponte (2003) reported the species, also 

typically a shallow-water species, to 76 m at Lee Stocking Island, 

and Norris and Olsen (1991) reported Valonia macrophysa to 92 

m at San Salvador Island, Bahamas.

Valonia ventricosa J. Agardh 

Specimens: DLB3748, 36–46 m; DLB7755, 49 m; DLB7165, 

50 m; DLB7991, 62 m; DLB3116, 100 m.

Eiseman and Blair (1982) reported the species from 69 m in 

the Gulf of Mexico, and Norris and Olsen (1991) reported it to 

90 m in the Bahamas. Valonia ventricosa is a common shallowwater element in the Puerto Rican and Caribbean algal flora. 

The deepest record for the species (as Ventricaria ventricosa (J. 

Agardh) J. L. Olsen et J. A. West) is ~160 m by Guimarães et al. 

(2008) from Brazil.

Bryopsidales, Bryopsidaceae

Bryopsis sp. 

Specimens: DLB7898, 55 m; DLB7815, DLB7828, 73 m.

*Derbesia osterhoutii (L. R. Blinks et A. C. H. Blinks) Page

Specimens: DLB7349, 49 m; DLB7087, 56 m; DLB7844, 

DLB7852, 82 m.

Derbesia osterhoutii is a life history alternate that in deepwater collections in Puerto Rico, and also, the Bahamas are represented by the Halicystis stage, where it was reported to 61 m 

by Ballantine and Aponte (2003).

Caulerpaceae

Caulerpa ambigua Okamura

Specimen: DLB1829, 40 m.

Despite the single collection referenced above, Caulerpa ambigua (as Caulerpa vickersiae Børgesen), although never abundant, is commonly found in the Puerto Rican deeper-water flora. 

Caulerpa ambigua was reported from the Gulf of Mexico to 37 

m (as Caulerpa vickersiae) by Dawes and Van Breedveld (1969) 

and from 51 m at Eniwetok (also as C. vickersiae) by Gilmartin 

(1960).

Caulerpa chemnitzia (Esper) J. V. Lamour. 

Specimens: DLB7971, 62 m; DLB8035, 70 m; DLB8054, 70 m.

This entity has had a confused taxonomic history. Species 

assignation follows Belton et al. (2014), who listed the Indian 

Ocean as the type locality while regarding C. racemosa var. peltata (J. V. Lamour.) Eubank and C. peltata Lamouroux (both 

from the Caribbean) as heterotypic synonyms. Taylor (1960) 

reported the species as C. peltata J. V. Lamour., having been 

dredged to over 30 m depth. The species has also been reported 

to 73 m in the Gulf of Mexico by Eiseman and Blair (1982) as 

well as to 73 m by Norris and Olsen (1991) in the Bahamas.

Caulerpa mexicana Sond. ex Kütz. 

Specimens: DLB3721, 36–46 m; DLB7276, 46 m; DLB7351, 

DLB7735, 49 m.

Caulerpa mexicana was reported as dredged to 73 m and 

“probably to 110 m” by Taylor (1960:141) in the tropical west 

Atlantic. In Pacific Hawaii, C. mexicana was reported to 129 m 

by Spalding (2012).

Caulerpa microphysa (Weber van Bosse) Feldmann 

Specimens: DLB1810, DLB8089, 50 m; DLB4121, 55 m; 

DLB7448, 57 m; DLB7952, 67 m; DLB7566, 70 m; 

DLB7877, 82 m.

The species has been reported from 61 to 70 m at San Salvador Island, Bahamas (Norris and Olsen, 1991) and to 110 m in 

Bermuda by Frederick (1963).

Caulerpa racemosa (Forssk.) J. Agardh

Specimens: DLB7158, DLB7179, 50 m.

In Puerto Rico, Caulerpa racemosa is a common species 

in exposed reef flat habitats and has been relatively rarely collected in deeper water. Taylor (1960) reported the species to 55 

m. In deep water, the species is also known from Hawaii at 60 m 

(Agegian and Abbott, 1985) and Eniwetok to 63 m (Gilmartin, 

1960).

*Caulerpa verticillata J. Agardh

Specimens: DLB7730, 49 m; DLB7421, 50 m; DLB7895, 55 

m; DLB7447, 57 m; DLB1233, DLB7571, DLB1715, 58 

m; DLB1748, 61 m; DLB7917, 70 m; DLB7600, 77 m; 

DLB3123, 100 m.

In Puerto Rico, Caulerpa verticillata is a very common 

shallow-water epiphyte on mangrove prop roots, moderately 

common in offshore algal plains at intermediate depths, as well 

as being a not uncommon element in deeper-water algal turfs. 

Norris and Bucher (1982) also indicated that the species lived 

both in shallow-water and deeper-water habitats to 12 m. 

*Caulerpa webbiana Mont. 

Specimens: DLB7135, 50 m; DLB3205, 68 m; DLB3117, 100 m.

Taylor (1960) indicated that the species is found from the 

littoral to a depth of 50 m.
NUMBER 105 • 2 1

Codiaceae

Codium isthmocladum Vickers ssp. clavatum (Collins et Herv.) 

P. C. Silva

Specimens: DLB4197, 36 m; DLB4681, 55–60 m.

Norris and Olsen (1991) reported the subspecies to 73 m in 

the Bahamas. The deepest known report of C. isthmocladum is 

75 m by Joly and Yoneshigue Braga (1966) from Brazil.

Codium taylorii P. C. Silva 

Specimens: DLB4199, 36 m; DLB7180, 50 m.

Taylor (1960) indicated that the species was limited to shallow water; however, Hanisak and Blair (1988a) also reported 

Codium taylorii to 50 m in Florida. The deepest report for the 

species is that by Suárez et al. (2015), who reported C. taylorii to 

a depth of 55 m; however, the depth is reported without citation 

or location.

Rhipiliaceae

Rhipiliopsis profunda (Eiseman et S. A. Earle ) J. N. Norris et 

S. Blair 

FIGURE 5E

Specimens: DLB8073, 50 m; DLB3607, 55–65 m; DLB3607, 

55–65 m; DLB8009, DLB8067, 70 m.

Rhipiliopsis profunda was reported to 92 m in the Bahamas 

by Norris and Olsen (1991).

Rhipiliopsis reticulata (C. Hoek) Farghaly et Denizot 

Specimens: DLB7718, DLB7733, 49 m; DLB7633, 70 m; 

DLB7870, 82 m.

The species, as Udotea reticulata Hoek (1978), was originally described from Curaçao as a coral reef–associated alga and 

was collected to 60 m. Norris and Olsen (1991) reported the 

species to 92 m in the Bahamas.

Rhipiliopsis stri (S. A. Earle et J. R. Young) Farghaly et Denizot

Specimens: DLB1234, 49 m; DLB1808, 165 m; DLB8010, 

70 m.

Rhipiliopsis stri was reported to a depth of 76 m in the Bahamas by Ballantine and Aponte (2003).

Udoteaceae

Avrainvillea asarifolia Børgesen

Specimens: DLB8099, 50 m; DLB8016, 70 m.

Taylor (1960) indicated that Avrainvillea asarifolia is a 

deepwater species and had been dredged to 90 m, although it is 

not uncommon at an offshore 17-m-deep algal plain in southwest Puerto Rico.

*Avrainvillea elliotii A. Gepp et E. Gepp

Specimen: DLB4115, 55 m.

Joly et al. (1965) reported the species (as A. atlantica A. B. 

Joly et Yamaguishi) from 2 to 3 m depth in Brazil. Dawes and 

Mathieson (2008) reported the species to 10 m from the Florida 

Keys.

Cladocephalus luteofuscus (P. Crouan et H. Crouan) Børgesen 

Specimen: DLB4127, 55 m.

Taylor (1960) reported the species to 72 m, and the species 

is also known from very shallow water, less than 1 m in depth in 

Bermuda (Schneider et al., 2010).

Halimeda copiosa Goreau et E. A. Graham 

FIGURE 5C

Specimens: DLB7140, 50 m; DLB7181, 50 m; DLB7550, 62 

m; DLB7559, 70 m; DLB7856, 82 m; DLB3125, 91 m; 

DLB3112, 100 m.

Ballantine and Aponte (2003) reported Halimeda copiosa to 

a maximum depth of 107 m in the Bahamas, where it commonly 

formed long pendant chains hanging from shelf wall ledges, as 

illustrated by Suárez et al. (2015:179, fig. 195). Blair and Norris (1988) commented that H. copiosa was the most common 

Halimeda species in the deep-water algal community at San Salvador, where they indicated a maximum depth of 152 m. 

Halimeda cryptica Colinvaux et E. A. Graham

Specimens: DLB7610, DLB446, DLB8070, 70 m; DLB7841, 

DLB7862, 82 m.

The species was reported between 61 and 152 m from San 

Salvador Island, Bahamas (Blair and Norris, 1988). Halimeda cryptica as var. acerifolia D. L.Ballant. (DLB1755, 61 m; 

DLB7638, 70 m) was initially collected in Puerto Rico (Ballantine, 1982) from 53 m. 

Halimeda discoidea Decne. 

Specimens: DLB4189, 36 m; DLB7145, 50 m; DLB4679, 

55–60 m; DLB8051, 70 m.

Blair and Norris (1988) and Littler and Littler (2000) reported maximum depths of 73 and 80 m, respectively, for the 

species in the tropical western Atlantic. Spalding (2012) reported 

H. discoidea to 86 m in Pacific Hawaii. The deepest known depth 

for the species is 115 m (Suárez et al., 2015); however, the depth 

is reported without citation or location.

Halimeda goreaui W. R. Taylor 

Specimens: DLB3688, 36 m; DLB7432, DLB7728, 49 m; 

DLB7049, 52 m; DLB7466, DLB7507, 61 m; DLB7473, 

63 m; DLB7946, 67 m; DLB8092, 70 m; DLB7826, 73 m.

Halimeda goreaui was described from Jamaica (Taylor, 

1962) on the basis of collections to a depth of 42 m. In Puerto 

Rico, H. goreaui is extremely common at the insular shelf edge 

break (18 m) and becomes less abundant with depth. The deepest known depth for the H. goreaui is 80 m (Suárez et al., 2015); 

however, the depth is reported without citation or location.
2 2 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

Halimeda gracilis Harv. ex J. Agardh 

Specimens: DLB4680, 55–60 m; DLB7502, 61 m; DLB7972, 

62 m; DLB8037, 70 m.

In Puerto Rico, Halimeda gracilis is rarely collected at 

depths shallower than 17 m. Joly et al. (1968) reported the species to a depth of 75 m in Brazil, and Blair and Norris (1988) collected the species to 122 m in the Bahamas. The deepest known 

reported record for Halimeda gracilis is that by Agegian and Abbott (1985), who reported the species to 125 m from Hawaii and 

Johnson Atoll.

*Halimeda hummii D. L. Ballant. 

Specimens: DLB7555, 62 m; DLB8044, 70 m.

The species was based on collections from both shallow and 

moderate depths in Puerto Rico (Ballantine, 1982) and is also known 

from moderate depths in Panama (Wysor and Kooistra, 2003).

Halimeda pumila Verbruggen, D. Littler et Littler

FIGURE 5D

Specimens: DLB7115, DLB7396, 59 m; DLB7867, 82 m.

Verbruggen et al. (2007) described the species on the basis 

of their Bahamian collections to 50 m depth and regarded the 

Littler and Littler (2000) report of Halimeda cryptica var. acerifolia (to a maximum depth of 174 m) to be H. pumila. The 

first report of the species in Puerto Rico was by Ballantine et al. 

(2011). 

Halimeda tuna (J. Ellis et Sol.) J. V. Lamour. 

Specimen: DLB8030, 70 m.

Taylor (1960) reported H. tuna to 80 m, and Blair and Norris (1988) reported the species to 100 m depth at San Salvador 

Island, Bahamas.

*Penicillus capitatus J. V. Lamour. 

Specimen: DLB3191, 80–100 m.

In Puerto Rico, this alga is extremely common in shallowwater seagrass habitats and in moderate-depth algal plain habitats. Sangil et al. (2010) reported the species growing in large 

stands to 50 m depth in the Canary Islands; however, deepwater Puerto Rico specimens were always collected as solitary 

individuals.

*Rhipocephalus oblongus (Decne) Kütz.

Specimen: DLB8469, 36 m.

The species was only recently recognized from Puerto Rico 

(Ballantine et al., 2015); however, it is typically found in shallow 

water (Taylor, 1960).

Rhipocephalus phoenix (J. Ellis et Sol.) Kütz.

Specimens: DLB3687, 36 m; DLB3389, 40 m; DLB7162, 

DLB8074, 50 m.

Rhipocephalus phoenix was reported to 60 m in the Florida 

Keys by Leichter et al. (2008). Taylor (1960) indicated a maximum depth of 72 m for the species. 

*Udotea abbottiorum D. Littler et Littler 

Specimen: DLB3744, 36–46 m.

This species is normally a plant of shallow water; the holotype is from 5 m depth (Littler and Littler, 1990).

*Udotea caribaea D. Littler et Littler

Specimen: DLB8455, 46–55 m.

Prior to the report from 55 m in Bermuda by Schneider et 

al. (2010), the species was typically reported from depths of less 

than 7 m. Ballantine et al. (2015) first reported the species from 

Puerto Rico. 

*Udotea conglutinata (J. Ellis et Sol.) J. V. Lamour. 

Specimen: DLB7971, 62 m.

Udotea conglutinata was reported to a depth of 40 m by 

Littler and Littler (1990), to 52 m from Bermuda by Schneider et 

al. (2010), and to a depth of 55 m by Dawes and Van Breedveld 

(1969) in the Florida Gulf of Mexico.

Udotea cyathiformis Decne. var. flabellifolia D. Littler et Littler 

Specimens: DLB4201, 36 m; DLB3745, 36–46 m; DLB8179, 45 m.

Udotea cyathiformis was reported to 56 m by Eiseman and 

Blair (1982) as U. cyathiformis f. infundibulum (J. Agardh) Littler et D. S. Littler. The maximum known depth for the species is 

73 m (Frederick, 1963). The deepwater collections from Puerto 

Rico all possessed a peltate flabellum. Flabellum filaments were 

free of lateral appendages, and those from stipe filaments were as 

illustrated by Littler and Littler (1990) for the variety.

Udotea dixonii D. Littler et Littler 

Specimens: DLB3390, 40 m; DLB4676, 55–60 m.

Littler and Littler (1990, 2000) characterized this species as 

typically being from deepwater habitats, with the holotype from 

46 m. Udotea dixonii has also been reported from 50 to 56 m 

by Mateo-Cid et al. (2013). The deepest report for the species 

is by Suárez et al. (2015), who reported it to a depth of 55 m 

(although without citation or location).

Udotea flabellum (J. Ellis et Sol.) J. V. Lamour. 

Specimens: DLB4186, 36 m; DLB4722, 60 m.

Normally a plant of shallow water (Littler and Littler 1990), 

this species has been reported by Taylor (1960) to have been 

dredged from greater than 70 m depth and reported to 40 m 

by Schneider (1976) from the North Carolina continental shelf. 

The deepest report of the species is to ~160 m from Brazil by 

Guimarães et al. (2008). 

*Udotea occidentalis A. Gepp et E. Gepp

Specimen: DLB8048, 70 m.

Udotea occidentalis is another typically shallow-water 

plant, although the species was reported to have been dredged 

to 36 m in St. Thomas, U.S. Virgin Islands (Littler and Littler, 

1990), and Mateo-Cid et al. (2013) reported the species from 50 

to 56 m from Campeche Banks, Mexico.
NUMBER 105 • 2 3

Udotea spinulosa M. Howe 

Specimen: DLB8440, 37–46 m.

The species was reported from Puerto Rico by Littler and 

Littler (1990) and has been reported as being dredged from over 

90 m depth (Taylor, 1960).

*Udotea unistratea D. Littler et Littler 

FIGURE 5F

Specimen: DLB8047, 70 m.

The holotype of U. unistratea was collected from Belize 

at 46 m depth (Littler and Littler, 1990), and Mateo-Cid et al. 

(2013) reported the species from 50 to 56 m from Campeche 

Banks, Mexico.

Dasycladales, Dasycladaceae

Neomeris annulata Dickie 

Specimen: DLB3742, 36–46 m.

Although this species is typically reported from shallowwater habitats, Taylor (1960) and Dawes and Mathieson (2008) 

indicated a maximum depth of 50 m for the species. Joly (1953) 

reported dredged specimens taken from between 30 and 50 m 

at Trindade Island, off the western coast of Brazil. The deepest 

known depth for Neomeris annulata is from Hawaii at 60 m 

(Agegian and Abbott, 1985).

Polyphysaceae

*Parvocaulis pusillus (M. Howe) S. Berger, Fettweiss, Gleissberg, Liddle, U. Richter, Sawitzky et Zuccarello

Specimens: DLBs.n., 40 m; DLB1222, 49 m.

Parvocaulis pusillus is based on collections from shallow 

water in Jamaica and the Bahamas (Howe, 1909).

DISCUSSION

Our collections of algae from water deeper than 35 m (listed 

above) comprise 59% Rhodophyta, 31% Chlorophyta, and 10% 

Phaeophyceae. Species reported with depth ranges collected are 

summarized in Appendix A. These 185 species constitute a 

third of the total algal flora known from Puerto Rico (Ballantine and Aponte, 2002b; Ballantine et al., 2015). Puerto Rican 

algal species restricted to depths greater than 35 m (Table 1) 

TABLE 1. Algal species mostly depth-restricted to the mesophotic realm in Puerto Rico, grouped by higher taxa. (Numbers in parentheses in column headings are counts of species listed.)

Rhodophyta (23) Phaeophyceae (3) Chlorophyta (17)

Botryocladia iridescens Dictyota stolonifera Anadyomene lacerata

Branchioglossum prostratum Lobophora canariensis Anadyomene saldanhae 

Ceramium bisporum Lobophora guadeloupensis Avrainvillea elliotii

Ceramium leptozonum Caulerpa ambigua

Ethelia sp. Cladocephalus luteofuscus

Flahaultia tegetiformans Cladophora coelothrix 

Frikkiella pseudoprostrata Halimeda copiosa

Frikkiella searlesii Halimeda cryptica

Gloiocladia atlantica Halimeda pumila 

Hydrolithon abyssophila Microdictyon boergesenii

Hypoglossum anomalum Rhipiliopsis profunda

Hypoglossum caloglossoides Rhipiliopsis reticulata

Hypoglossum rhizophorum Rhipiliopsis stri

Hypoglossum simulans Udotea dixonii

Perikladosporon abaxiale Udotea unistratea

Peyssonnelia gigaspora Verdigellas fimbriata

Peyssonnelia incomposita Verdigellas peltata

Peyssonnelia iridescens

Peyssonnelia sp. 1

Peyssonnelia sp. 2

Peyssonnelia sp. 3

Predaea laciniosa

Rhododictyon bermudense 
2 4 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

are dominated by members of Rhodophyta and Chlorophyta; 

only three species of Phaeophyceae listed are depth restricted. 

However, further examination of herbarium specimens of Lobophora canariensis and L. guadeloupensis is required to determine if those species are restricted to depths greater than 35 m 

in Puerto Rico. Lobophora canariensis, as L. payriae, is known 

from shallow water in Guadeloupe and Bermuda (Schultz et al., 

2015). Species such as Ceramium leptozonum, Flahaultia tegetiformans, Gloiocladia atlantica, Hypoglossum caloglossoides, 

Peyssonnelia incomposita, and Verdigellas peltata are mostly 

restricted to mesophotic depths in Puerto Rico and are rarely encountered in shallower water. Ceramium bisporum and Dictyota 

stolonifera, although restricted to the mesophotic zone in Puerto 

Rico, have been reported from shallow water from different geographical locations (Sartoni and Boddi, 2002; De Clerck, 2003; 

Rincon-Diaz et al., 2014).

Among each major division of algae, several families are 

disproportionately represented (Table 2). For example, among 

Rhodophyta, 27 species of Ceramiaceae (sensu lato), 17 species 

of Delesseriaceae (sensu lato), 11 species of Peyssonneliaceae 

(included here is one species of Etheliaceae), and 11 species of 

Rhodymeniaceae were identified. Among Phaeophyceae, Dictyotaceae comprised 14 species. Deepwater Chlorophyta included 7 

species of Caulerpaceae and 26 species of Udoteaceae. Although 

these two latter families were species rich and conspicuous because of their size, they were generally represented by only one 

or two individuals per collection. The most visually conspicuous 

members of the deepwater flora were coralline Rhodophyta (Hapalidiaceae) and Peyssonneliaceae species in addition to turfs. 

Many of the deepwater species, generally small statured, were 

identified as members of the species-rich and ubiquitous turf 

flora.

Three depth distributional groups may be thought of as 

comprising the mesophotic flora. Nearly half of the species (81) 

that are found in depths of 35 m or greater are found across 

the entire shelf, ranging from shallow nearshore habitats to 

the offshore mesophotic zone. There is a tendency for many 

of these species to drop out of the flora toward the shallower 

mesophotic region (for example, only approximately a quarter of these species are found in water deeper than ~60 m). A 

second distributional group consists of algae (~42 species) that 

are found ranging from intermediate depths to the mesophotic 

realm. More than half of these are also found at depths greater 

than ~60 m. An only slightly smaller group of algae (with a few 

minor exceptions), made up by 38 taxa identified to species (and 

accounting for 7% of the total Puerto Rican algal flora), appears 

to be restricted to water greater than 35 m in depth (see Table 

1). Thus, community composition gradually changes with depth, 

both by the elimination of shallow species and the inclusion of 

mesophotic-restricted species. It is important to note that some 

species included in Table 1 may be found in shallower habitats in 

locations other than Puerto Rico. 

Ballantine et al. (2010) reported that the principal calcium 

carbonate–producing organisms in the deep-water algal flora 

(>50 m depth) off the southwest coast of Puerto Rico were principally Peyssonnelia spp. and Corallinales. Similarly, many of the 

macroscopic species recognized at depths of 50 to 100 m in other 

studies (see Larkum et al., 1967; Littler et al., 1986; Aponte and 

Ballantine, 2001) were also dominated by Peyssonneliaceae and 

Corallinales. The coralline/Peyssonnelia dominance at mesophotic depths in tropical and warm temperate Atlantic regions 

may be a universal deepwater distributional pattern. 

 Hanisak and Blair (1988a) listed 157 species of algae from 

deeper than 40 m in Florida, and roughly a third of these species are the same as the above-reported deepwater species from 

Puerto Rico. Of these, only 20% had a deeper distribution in 

Florida than in Puerto Rico, possibly a function of greater water 

clarity in coastal Puerto Rico than coastal Florida.

On the basis of his study at Eniwetok (Pacific Ocean), 

Gilmartin (1960:218) concluded that there was not a “specifically distinct deep water algal flora” with a relatively large 

percentage of species also occurring in littoral and infralittoral 

environments. Conversely, Spalding (2012) indicated that 45% 

of the 76 total algae identified in Hawaiian mesophotic habitats 

were restricted to those depths. For the Puerto Rican mesophotic 

zone, the fact that there are 20 species (Rhodophyta and Chlorophyta) essentially restricted to the mesophotic realm (Table 1) 

and the fact that most Puerto Rican algal species are never found 

TABLE 2. Numbers of genera and species of families represented 

by four or more species reported from 35 m or greater in depth 

in Puerto Rico.

Family No. Genera No. Species

Rhodophyta

Ceramiaceae (sensu lato) 7 13

Dasyaceae 3 4

Callithamniaceae 4 5

Delesseriaceae (sensu lato) 9 15

Galaxauraceae 3 4

Hapalidiaceae 3 7

Halymeniaceae 3 4

Peyssonneliaceae and Etheliaceae 3 11

Rhodomelaceae 7 8

Rhodymeniaceae 7 11

Solieriaceae 4 4

Wrangeliaceae 7 9

Heterokontophyta

Dictyotaceae 6 14

Sargassaceae 1 4

Chlorophyta

Boodleaceae 3 4

Caulerpaceae 2 7

Cladophoraceae 3 4

Udoteaceae 7 26
NUMBER 105 • 2 5

in deep water are indicative that there is also a distinct deepwater 

flora in Puerto Rico. Within the mesophotic region, algal species composition changed with depth between approximately 30 

and 70 m. There was a fairly high similarity in species composition between 30 and 50 m (68%). The species similarity between 

30 and 70 m falls to 54%; in other words, roughly half of the 

species are cosmopolitan across these depths. There was also a 

fairly high similarity between 50 and 70 m (67%). This similarity would suggest an approximate depth of rapid change at 

about 50 m. 

Fifty-one percent of the algae reported herein (identified to 

species) are most likely the deepest known distributional depths 

for those species. This proportion may be slightly high as certainly not every depth record for each species has been located, 

and unfortunately, the literature is replete with species names accompanied by little data, particularly concerning depth. On the 

other hand, some of the deepwater algae reported herein at the 

generic level probably, or, in some cases, certainly, represent new 

species to science that would increase the proportion of obligate 

deepwater species. One frustration in working with algae from 

deep environments is that some species are rare enough that it is 

difficult to obtain more than a few individuals. Furthermore, it 

is not uncommon for the deep algae to be vegetative, lacking reproductive characters to support description. Frederick’s (1963) 

unpublished dissertation remarkably still includes many of the 

region’s deepest algal records despite the fact that the work was 

conducted over 50 years ago.

With the present report of Puerto Rican deepwater algae, 

we still feel that the mesophotic flora of the western Atlantic 

remains essentially only partially characterized. The geographical areas studied at any depth represent only a fraction of deepwater communities, and the percentage of deepwater floras 

studied in the Puerto Rico (and the Caribbean as a whole) is 

thus extremely small. Furthermore, even in areas that have been 

studied, the observation by Gilmartin (1960) broadly applies. He 

concluded that in his sampling at Eniwetok at depth, although 

species saturation curves were never calculated, probably not a 

single sampled site was adequately collected such that most species could be accounted for. This is probably true worldwide of 

most deepwater habitats sampled, which is understandable given 

limitations inherent in most deepwater sampling.

Many of the deeper-water studies to date have generally focused on a single systematic group (i.e., sponges, corals, algae) or 

have provided only broad systematic categories in characterization of deep communities. Historically, relatively few systematic 

collections of algae at any one site have been undertaken; thus, 

many records of the deepwater algae are based on incidental reports. Despite increased deepwater collection in recent decades 

with more sophisticated equipment, our overall understanding 

of deepwater algae is still limited. This limitation is due to several factors, one being that such studies have been limited to 

restricted geographical areas and another being that most of the 

region remains effectively unsampled. Virtually all deepwater 

excursions to date have resulted in the discovery of new species 

as well as important geographical distribution records. Further, 

given that the known mesophotic algal flora is rich floristically, it 

is virtually certain that sampling new mesophotic environments 

will continue to yield new species to science. 

ACKNOWLEDGMENTS

A number of funding sources have supported this research 

over the last three decades. These have included the Office of 

Research Coordination, University of Puerto Rico, Recinto 

Mayagüez; the National Marine Fisheries Service, which made 

collection time available aboard the submersible Johnson-­Sea-Link II and ship time available aboard the R/V Oregon II; and 

the National Oceanographic and Atmospheric Administration 

(NOAA), award number NA6NOS4780190, which funded 

the deep trimix technical diving. Kimberley Pugilise of NOAA’s 

Undersea Research Program kindly provided a digital copy of 

NOAA nautical chart 25640, from which Figure 1 was drafted. 

We thank Alice Tangerini, NMNH botanical illustrator, for 

drafting Figure 1. Finally, we thank James Norris, Smithsonian 

Institution, for nomenclatural discussions and Heather Spalding 

and Michael Wynne for their reading of the manuscript and for 

providing constructive criticism.

Appendix A: Summary of Puerto Rican 

Mesophotic Algal Specimens

TABLE A1. Identified mesophotic algal specimens from Puerto Rico and their depth ranges.

Species Mesophotic depth range (m)

Rhodophyta

Agardhiella ramosissima 55–60

Aglaothamnion cordatum 55–60

Amansia multifida 55

Amphiroa rigida 49–70

Amphiroa tribulus 49–61

Antithamnion antillanum 50–82

Antithamnion decipiens 40–82

Antithamnionella breviramosa 37

Antithamnionella graeffei 61

Apoglossum gregarium 61–100

Asparagopsis taxiformis 49–70

Asteromenia peltata 36–60

Augophyllum wysorii 49

Balliella pseudocorticata 36–58

Branchioglossum prostratum 61

Botryocladia iridescens 47–82

Botryocladia pyriformis 55–60

Botryocladia spinulifera 40–82

Botryocladia wynnei 63

Ceramium bisporum 46–100

Ceramium leptozonum 50

Ceramium nitens 61

Champia parvula 50–70

Champia vieillardii 40–55

Chrysymenia cf. ventricosa 37

Coelarthrum cliftonii 49–70

Cottoniella filamentosa 91

Cresia opalescens 49

Crouania pumila 55

Crouanophycus latiaxis 50

Cryptonemia crenulata 36–82

(Continued)
2 8 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

Dichotomaria marginata 61

Dichotomaria obtusata 61

Dictyurus occidentalis 36–46

Diplothamnion jolyi 36–50

Flahaultia tegetiformans 36–61

Frikkiella pseudoprostrata 46–82

Frikkiella searlesii 61

Galaxaura rugosa 50–70

Gayliella transversalis 62–70

Gloiocladia atlantica 41–100

Gracilaria isabellana 36–60

Grallatoria reptans 36–55

Halichrysis corallinaria 50

Griffithsia heteromorpha 49–52

Haloplegma duperreyi 49

Halymenia hancockii 55

Halymenia pseudofloresii 36–60

Herposiphonia secunda 61

Heterosiphonia crispella 52–70

Hydrolithon abyssophila 49–76

Hydrolithon farinosum 55–61

Hymenoclonium serpensa 52–87

Hypnea volubilis 55

Hypoglossum anomalum 36–91

Hypoglossum caloglossoides 49–77

Hypoglossum hypoglossoides 36–87

Hypoglossum. rhizophorum 40–87

Hypoglossum simulans 36–61

Hypoglossum tenuifolium 58–87

Jania adhaerens 50

Jania cubensis 36–70

Jania subulata 36–46

Leptofauchea? rhodymenioides 49–82

Laurencia intricata 36–60

Lophosiphonia cristata 49

Martensia pavonia 36–82

Meredithia? caribaea 67–70

Meredithia pulchella 50–70

Meristotheca gelidium 36

Myriogramme prostrata 52–67

Nitophyllum adhaerens 36–90

Perikladosporon abaxiale 87

Peyssonnelia boergesenii 49–61

Peyssonnelia flavescens 50–82

Peyssonnelia gigaspora 52–91

Peyssonnelia inamoena 46–73

Peyssonnelia incomposita 52–82

Peyssonnelia iridescens 46–87

TABLE A1. (Continued)

Species Mesophotic depth range (m)

Rhodophyta
NUMBER 105 • 2 9

Platysiphonia caribaea 36–100

Polystrata fosliei 70

Predaea laciniosa 61–70

Rhododictyon bermudense 63–91

Seirospora occidentalis 36–87

Seirospora viridis 58

Solieria filiformis 36

Spermothamnion investiens 55–87

Spermothamnion cf. macromeres 82

Spongoclonium caribaeum 36–87

Taenioma nanum 49

Titanophora incrustans 60

Tricleocarpa fragilis 62–70

Wrangelia bicuspidata 46–82

Wrangelia gordoniae 46–73

Wrightiella tumanowiczi 50–55

Wurdemannia miniata 52–63

Heterokontophyta

Canistrocarpus cervicornis 49–50

Dictyopteris delicatula 36–82

Dictyopteris justii 36–60

Dictyota bartayresiana 61

Dictyota ciliolata 36–46

Dictyota humifusa 46–82

Dictyota jamaicensis 36–46

Dictyota pulchella 50–70

Dictyota stolonifera 62–70

Lobophora canariensis 37-82

Lobophora guadeloupensis 40–70

Lobophora variegata 36–37

Padina cf. sanctae-crucis 37

Stypopodium zonale 50

Sporochnus bolleanus 36–55

Sargassum filipendula 55

Sargassum hystrix 70

Sargassum ramifolium 36

Chlorophyta

Anadyomene lacerata 40–62

Anadyomene saldanhae 36–100

Anadyomene stellata 50–100

Avrainvillea asarifolia 50–70

Avrainvillea elliotii 55

Caulerpa ambigua 40

Caulerpa chemnitzia 62–70

Caulerpa mexicana 36–49

Caulerpa microphysa 50–82

Caulerpa racemosa 50

(Continued)

Species Mesophotic depth range (m)

Rhodophyta
3 0 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

Caulerpa verticillata 49–100

Caulerpa webbiana 50–100

Cladocephalus luteofuscus 55

Cladophora coelothrix 82

Codium isthmocladum 36–60

Codium taylorii 36–50

Derbesia osterhoutii 49–82

Gayralia oxysperma 82

Halimeda copiosa 50–100

Halimeda cryptica 70–82

Halimeda discoidea 36–70

Halimeda goreaui 36–73

Halimeda gracilis 55–70

Halimeda hummii 62–70

Halimeda pumila 59–82

Halimeda tuna 70

Penicillus capitatus 80–100

Microdictyon boergesenii 49–100

Neomeris annulata 36–46

Parvocaulis pusillus 40–49

Phaeophila dendroides 65

Phyllodictyon anastomosans 36

Phyllodictyon pulcherrimum 50–70

Rhipiliopsis profunda 50–70

Rhipiliopsis reticulata 49–82

Rhipiliopsis stri 49–70

Rhipocephalus oblongus 36

Rhipocephalus phoenix 36–50

Struvea elegans 46–70

Udotea abbottiorum 36–46

Udotea caribaea 46–55

Udotea conglutinata 62

Udotea cyathiformis 36–45

Udotea dixonii 40–60

Udotea flabellum 36–60

Udotea occidentalis 70

Udotea spinulosa 37–46

Udotea unistratea 70

Valonia macrophysa 82

Valonia ventricosa 36–100

Verdigellas fimbriata 46–65

Verdigellas peltata 46–100

aThe species is listed as a form genus.

TABLE A1. (Continued)

Species Mesophotic depth range (m)

Chlorophyta
Locations of collection sites sampled around Puerto Rico are provided here, grouped 

by geographic area. Numbers for sites correspond to collection site numbers in Figure 1.

EAST

1. Vieques “El Seco” [approximately 18°7′N, 65°15′W]

70 m: DLB8038–8053, DLB8092–8094

SOUTH

2. Southeast, Puerto Rico, National Oceanic and Atmospheric Administration R/V

Seward Johnson collected from DSV Johnson-­Sea-­Link [approximately 17°59′N, 

65°49′W ]

66 m: DLB2321

3. Grappler Bank (SE Puerto Rico), collected by University of Puerto Rico technical diving team and dredged from R/V Oregon II [17°47′56.63′′N, 65°54′39.53′′W]

55–90 m: DLB3089–3104, DLB3201–3206, DLB8381–8382

100 m: DLB3111–3124

4. Edge of insular shelf, offshore Salinas [approximately 17°52′N, 66°18′W]

49 m: DLB1245–1249

5. Edge of insular shelf, Ponce [approximately 17°56′N, 66°40′W]

55 m: DLB8254

6. Edge of insular shelf, Guanica, collected in submersible Johnson-­Sea-­Link [approximately 17°56′N, 66°50′W]

50 m: DLB2152–2179

7. Edge of insular shelf, “Baranca” [approximately 17°54′N, 66°59′W]

49 m: DLB7432–7442

61 m: DLB7522–7523

70 m: DLB7615–7616

Appendix B: Collection Locality Data
3 2 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

8. Edge of insular shelf, “Weinberg” [17°53.423′N, 66°59.320′W]

50 m: DLB7314–7347, DLB7380–7394, DLB7403–7409

9. Edge of insular shelf, “Buoy” [17°53.292′N, 66°59.872′W]

36 m: DLB8430

37 m: DLB1705–1709

40 m: DLB1208–1215, DLB1250–1258, DLB1817–1832

46 m: DLB7026–7033

49 m: DLB435–436, 1218–1242, DLB7011–7124

50 m: DLB1786–1813, DLB7305–7308

52 m: DLB7048–7059, DLB7070–7073

55 m: DLB7080–7087

58 m: DLB1710–1738, DLB7514–7517

61 m: DLB1740–1783, DLB7518–7521

70 m: DLB446–450

10. Edge of insular shelf, “Black Wall” [17°53.090′N, 

67°00.863′W]

46 m: DLB7074, DLB7276–7287

49 m: DLB7800

57 m: DLB7444–7455

70 m: DLB7635–7645

82 m: DLB7675–7685, DLB7866–7891

11. Edge of insular shelf, “Hole in the Wall” [17°53.077′N, 

67°1.315′W]

37 m: DLB7219–7225 

41 m: DLB7464 

49 m: DLB7709–7746, DLB7811–7812, DLB7956–7957

50 m: DLB7420–7431 

55 m: DLB7892–7905 

61 m: DLB7465–7471 

63 m: DLB7472–7486

67 m: DLB7942–7955

70 m: DLB7618–7634

73 m: DLB7813–7828

77 m: DLB7590–7609

91 m: DLB8160–8169

12. Edge of insular shelf, “El Hoyo” [17°52.537′N, 67°2.648′W]

46 m: DLB8719

49 m: DLB7348–7379, DLB7747–7760, DLB7395–7402,

DLB7920–7936

50 m: DLB8177

52 m: DLB7829–7839, DLB8181

61 m: DLB7772–7775

62 m: DLB7547–7555

63 m: DLB7525–7532, DLB7686–7692

70 m: DLB7610–7614, DLB7776–7779, DLB7906–7918

73 m: DLB7646

82 m: DLB7664–7674, DLB7761–7771, DLB7840–7848, 

DLB7850–7865, DLB8192–8186 

87 m: DLB8143–8146 

91 m: DLB8147–8154

13. Edge of insular shelf, “El Precipico” [17°52.510′N, 

67°2.929′W]

61 m: DLB7487–7513

70 m: DLB7556–7581

14. Edge of insular shelf, South La Parguera, dredged R/V Oregon II (station 46003) [approximately 17°52′N, 67°6′W ]

40 m: DLB2903–2916

WEST

15. Tourmaline Reef [approximately 18°6′N, 67°24′W]

37 m: DLB8422–8424

16. West of Mayaguez, dredged R/V Oregon II (station 45998) 

[approximately 15°11′N, 67°24′W]

91 m: DLB2896–2902, DLB3125–3130

17. Bajo de Sico [approximately 18°13′N, 67°21′W]

50 m: DLB7133–7199

MONA ISLAND

18. East Mona Island [approximately 18°6′N, 67°50′W] 

46–55 m: DLB8455–8456, DLB8468

61–82 m: DLB8470

19. Southeast Mona Island [approximately 18°5′N, 67°50′W]

70 m: DLB8026–8037

20. South Mona Island [approximately 18°5′N, 67°51′W]

62 m: DLB7971–7993

21. South Mona Island [18°3′N, 67°53′W]

50 m: DLB8095–8101

70 m: DLB8003–8025 

80–100 m: DLB3188–3200

22. South Mona Island, dredged from R/V Isla Magueyes [approximately 18°3′N, 67°55′W]

45–65 m: DLB3605–3607, DLB3710–3715

23. Mona Island, Carabinero [18°3′54.93′′N, 67°55′36.65′′W] 

35–37 m: DLB3686–3708, DLB8469

50 m: DLB8073–8091

24. Mona Island (wall) [approximately 18°4′N, 67°56′W]

70 m: DLB8054–8072

25. West Mona Island [approximately 18°6′N, 67°57′W]

46 m: DLB8465–8467
NUMBER 105 • 3 3

26. West Monito [18°9′30′′N, 67°56′99′′W]

40 m: 3389–3391

NORTH COAST

27. Punta Aguyereada, Aguadilla [approximately 18°33′N, 67°8′W]

36 m: DLB4178–4212

60 m: DLB4694–4698, DLB4712–4733

28. Isabela, dredged R/V Isla Magueyes [approximately 18°32′N, 

67°1′W]

37 m: DLB4410–4441

55–60 m: DLB4673–4693

29. Guajataca, Quebredillas, dredged [approximately 18°30′N, 

66°56′W]

55 m: DLB8457–8459

30. Punta Caracoles, Arecibo, dredged [approximately 18°29′N, 

66°42′W]

36 m: DLB4031–4074

55 m: DLB4114–4129

31. Islote, Arecibo [approximately 18°30′N, 66°36′W]

55 m: DLB4802–4808

37–46 m: DLB8440–8441

32. Between Punta Cerro Gorda and Punta Fraile (north coast), 

dredged [approximately 18°30′N, 66°18′W]

36–46 m: DLB3716–3751

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Wynne, M. J., I. R. Price, and D. L. Ballantine. 1989. Distinctions between Hypoglossum barbatum Okamura, H. minimum Yamada and H. simulans sp. nov. 

(Delesseriaceae, Rhodophyta). Phycologia 28:28–38. doi:10.2216/i0031-

8884-28-1-28.1.

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Index of Genera and Species

Acrosymphyton, 15

caribaeum, 15

purpuriferum, 15

Agardhiella, 11, 27

ramosissima, 11, 27

Agaricia, 13

lamarcki, 13

Aglaothamnion, 6, 27

cordatum, 6, 27

Amansia, 8, 27

multifida, 8, 27

Amphiroa, 3, 27

rigida, 3, 27

tribulus, 3, 27

Anadyomene, 19, 23, 29

lacerata, 19, 23, 29

saldanhae, 19, 23, 29

stellata, 19, 23, 29

Antithamnion, 6, 28

antillanum, 6, 28

decipiens, 6, 28

flagellata, 6

lherminieri, 6

ogdeniae, 6

Antithamnionella, 6, 28

breviramosa, 6, 28

graeffei, 6, 28

latiaxis, 6

Apoglossum, 7, 27

gregarium, 7, 27

Asparagopsis, 6, 15, 27

taxiformis, 6, 15, 27

Asteromenia, 14, 27

peltata, 14, 27

Augophyllum, 7, 27

wysorii, 7, 27

Avrainvillea, 21, 23, 29

asarifolia, 21, 23, 29

elliotii, 21, 23, 29

Bakothamnion, 6

curassavicum, 6

Balliella, 6, 27

pseudocorticata, 6, 27

Bonnemaisonia, 15

asparagoides, 15

hamifera, 15

Botryocladia, 9, 14, 15, 23, 27

iridescens, 9, 14, 23, 27

pyriformis, 14, 27

spinulifera, 14, 27

wynnei, 15, 27

Branchioglossum, 7, 23, 27

prostratum, 7, 23, 27

pseudoprostratum, 7, 28

Bryopsis sp., 20

Callithamnion, 6, 15

cordatum, 6

serpens, 15

Canistrocarpus, 15, 29

cervicornis, 15, 16, 29

Caulerpa, 20, 23, 29

ambigua, 20, 23, 29

chemnitzia, 20, 29

mexicana, 20, 29

microphysa, 20, 29

peltata, 20

racemosa, 20, 29

verticillata, 20, 30

vickersiae, 20

webbiana, 20, 30

Ceramium, 6, 7, 23, 24, 27

bisporum, 6, 23, 24, 27

byssoideum, 7

flaccidum, 7

leptozonum, 7, 23, 24, 27

nitens, 7, 27

spp., 7
4 0 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

Ceramium (continued)

transversale, 7

transversalis, 7

Champia, 14, 27

parvula, 14, 27

vieillardii, 14, 27

Chondria, 8

spp., 8

Chrysymenia, 15, 27

cf. ventricosa, 15, 27

sp., 15

Cladocephalus, 21, 23, 30

luteofuscus, 21, 23, 30

Cladophora, 20, 23, 30

coelothrix, 20, 23, 30

Cladophoropsis, 19

membranacea, 19

sp., 19

Codium, 21, 30

isthmocladum, 21, 30

 ssp. clavatum, 21

taylorii, 21, 30

Coelarthrum, 15, 27

cliftonii, 15, 27

Contarinia sp., 11

Cottoniella, 10, 27

filamentosa, 10, 27

Cresia, 9, 15, 27

opalescens, 9, 15, 27

Crouania, 6, 27

pumila, 6, 27

Crouanophycus, 6, 27

latiaxis, 6, 27

Cryptonemia, 12, 14, 27

crenulata, 12, 27

sp., 14

Dasya sp., 7

Derbesia, 20, 30

osterhoutii, 20, 30

Dichotomaria, 3, 5, 28

marginata, 3, 28

obtusata, 5, 28

Dictyopteris, 15, 16, 29

delicatula, 15, 29

justii, 16, 29

Dictyota, 3, 15, 16, 23, 24, 29

bartayresiana, 16, 29

cervicornis, 15

ciliolata, 16, 29

divaricata, 16

humifusa, 16, 29

jamaicensis, 16, 29

pulchella, 16, 29

stolonifera, 16, 23, 24, 29

Dictyurus, 7, 28

occidentalis, 7, 28

Diplothamnion, 10, 28

jolyi, 10, 28

Ethelia sp., 5, 12, 23

Eucheuma, 11

schrammii, 11

Falkenbergia, 6, 15

hillebrandii, 15

Fauchea, 14

peltata, 14

Flahaultia, 11, 23, 24, 28

tegetiformans, 11, 23, 24, 28

Frikkiella, 3, 7, 23, 28

pseudoprostrata, 3, 7, 23, 28

searlesii, 3, 7, 23, 28

Galaxaura, 5, 6, 28

cylindrica, 6

obtusata var. major, 5

rugosa, 5, 28

Gayliella, 7, 28

transversalis, 7, 28

Gayralia, 19, 30

oxysperma, 19, 30

Gloiocladia, 14, 23, 24, 28

atlantica, 14, 23, 24, 28

Gracilaria, 12, 28

isabellana, 12, 28

Grallatoria, 10, 28

reptans, 10, 28

Griffithsia, 10, 28

heteromorpha, 10, 28 

Halichrysis, 14, 15, 28

corallinaria, 15, 28

peltata, 14

Halicystis, 20

Halimeda, 13, 18, 21, 22, 23, 30

copiosa, 18, 21, 23, 30

cryptica, 13, 21, 23, 30

 var. acerifolia, 21, 22

discoidea, 21, 30

goreaui, 21, 30

gracilis, 22, 30

hummii, 22, 30

pumila, 18, 22, 23, 27, 30

tuna, 22, 30

Haliptilon, 3

cubense, 3

Haloplegma, 10, 23

duperreyi, 10, 23

Halymenia, 14, 28

hancockii, 14, 28

pseudofloresii, 14, 28

Herposiphonia, 8, 10, 28

secunda, 8, 28

sp., 10

Heterosiphonia, 7, 28

crispella, 7, 28

sp., 7

wurdemannii, 7

Hydrolithon, 3, 5, 13, 23, 28

abyssophila, 3, 5, 13, 23, 28

farinosum, 3, 28

 var. chalicodictyum, 3

Hymenoclonium, 9, 15, 28

serpens, 9, 15, 28

Hypnea, 11, 28

volubilis, 11, 28

Hypoglossum, 8, 9, 23, 24, 28

anomalum, 8, 9, 23, 28

caloglossoides, 8, 9, 23, 24, 28

hypoglossoides, 8, 28

rhizophorum, 8, 23, 28

simulans, 8, 23, 28

tenuifolium, 8, 28

Jania, 3, 28

adhaerens, 3, 28

cubensis, 3, 28

subulata, 3, 28

Kallymenia, 11

limminghei, 11

Laurencia, 10, 28

intricata, 10, 28

Leptofauchea, 15, 28

rhodymenioides, 15, 28

Lobophora, 3, 13, 16, 17, 23, 24, 29

canariensis, 16, 23, 24, 29

guadeloupensis, 13, 16, 17, 23, 24, 29

payriae, 16, 17, 24

variegata, 16, 17, 29

Lomentaria, 14

baileyana, 14

divaricata, 14

sp., 14

Lophosiphonia, 10, 18, 28

cristata, 10, 18, 28

Martensia, 8, 28

pavonia, 8, 28

Meredithia, 11, 15, 28

caribaea, 11, 28

microphylla, 15

pulchella, 11, 28

Meristotheca, 11, 28

gelidium, 11, 28

Mesothanmion, 10

caribaeum, 10

Microdictyon, 18, 19, 23, 30

boergesenii, 18, 19, 23, 30

Myriogramme, 8, 9, 28

prostrata, 8, 9, 28

Neomeris, 23, 30

annulata, 23, 30

Nitophyllum, 8, 28

adhaerens, 8, 28

Padina, 17, 29

cf. sanctae-crucis, 17, 29

Parvocaulis, 23, 30

pusillus, 23, 30

Penicillus, 22, 30

capitatus, 22, 30

Perikladosporon, 7, 23, 28

abaxiale, 7, 23, 28

Peyssonnelia, 5, 6, 12, 13, 23, 24, 28

boergesenii, 12, 28

flavescens, 5, 6, 12, 13, 28
NUMBER 105

• 4 1

gigaspora, 5, 12, 23, 28

inamoena, 12, 28

incomposita, 5, 12, 23, 24, 25, 28

iridescens, 5, 6, 12, 13, 23, 28

sp. 1, 12, 23

sp. 2, 5, 12, 23

sp. 3, 5, 12, 23

Phaeophila, 19, 30

dendroides, 19, 30

Phyllodictyon, 18,19, 30

anastomosans, 19, 30

pulcherrimum, 18, 19, 30

Platysiphonia, 10, 29

caribaea, 10, 29

Polysiphonia, 10

spp., 10

Polystrata, 5, 12, 29

fosliei, 5, 12, 29

Predaea, 11, 23, 29

laciniosa, 11, 23, 29

sp., 11

Rhipiliopsis, 18, 21, 23, 30

profunda, 18, 21, 23, 30

reticulata, 21, 23, 30

stri, 21, 23, 30

Rhipocephalus, 22, 30

oblongus, 22, 30

phoenix, 22, 30

Rhododictyon, 9, 10, 23, 29

bermudense, 9, 10, 23, 29

Rosenvingea sp., 17

Sargassum, 17, 29

filipendula, 17, 29

hystrix, 17, 29

ramifolium, 17, 29

sp., 17

Seirospora, 6, 29

occidentalis, 6, 29

viridis, 6, 29

Solieria, 11, 29

filiformis, 11, 29

Spermothamnion, 10, 29

investiens, 10, 29

 var. cidaricola, 10

cf. macromeres, 10, 29

Sphacelaria sp., 17

Spongoclonium, 10, 29

caribaeum, 10, 29

Sporochnus, 17, 29

bolleanus, 17, 29

Struvea, 19, 30

elegans, 19, 30

Stypopodium, 17, 29

zonale, 17, 29

Taenioma, 8, 29

nanum, 8, 29

Titanophora, 11, 12, 29

incrustans, 11, 12, 29

Trailliella, 15

intricata, 15

Tricleocarpa, 5, 29

fragilis, 5, 6, 29

Udotea, 21, 22, 30

abbottiorum, 22, 30

caribaea, 22, 30

conglutinata, 22, 30

cyathiformis, 22, 30

 f. infundibulum, 22

 var. flabellifolia, 22

dixonii, 22, 30

flabellum, 22, 30

occidentalis, 22, 30

reticulata, 21

spinulosa, 22, 30

unistratea, 18, 22, 30

Valonia, 20, 30

macrophysa, 20, 30

ventricosa, 20, 30

Ventricaria, 22

Verdigellas, 2, 17, 18, 23, 24, 30

fimbriata, 17, 23, 30

peltata, 17, 18, 23, 24, 30

Wrangelia, 10, 11, 13, 29

bicuspidata, 10, 11, 29

gordoniae, 11, 29

Wrightiella, 10, 29

tumanowiczi, 10, 29

Wurdemannia, 11, 20, 29

miniata, 11, 20, 29
SUMMARY OF REQUIREMENTS FOR SMITHSONIAN CONTRIBUTIONS SERIES

For comprehensive guidelines and specifications, visit www.scholarlypress.si.edu. 

ABSTRACTS must not exceed 300 words. 

TEXT must be prepared in a recent version of Microsoft Word; use a Times font in 12 point for regular text; be double 

spaced; and have 1” margins. 

REQUIRED ELEMENTS are title page, abstract, table of contents, main text, and references.

FIGURES must be numbered sequentially (1, 2, 3, etc.) in the order called out; have components lettered consistently (in 

size, font, and style) and described in captions; include a scale bar or scale description, if appropriate; include any legends 

in or on figures rather than in captions. Figures must be original and must be submitted as individual TIF or EPS files.

FIGURE FILES must meet all required specifications in the Digital Art Preparation Guide. Color images should be 

requested only if required. 

TAXONOMIC KEYS in natural history manuscripts should use the aligned-couplet form for zoology. If cross referencing 

is required between key and text, do not include page references within the key, but number the keyed-out taxa, using the 

same numbers with their corresponding heads in the text.

SYNONYMY IN ZOOLOGY must use the short form (taxon, author, year:page), with full reference at the end of the 

manuscript under “References.”

REFERENCES should be in alphabetical order, and in chronological order for same-author entries. Each reference should 

be cited at least once in main text. Complete bibliographic information must be included in all citations. Examples of the 

most common types of citations can be found at SISP’s website under Resources/Guidelines.