CSIRO PUBLISHING

Invertebrate Systematics, 2015, 29, 345–368

http://dx.doi.org/10.1071/IS14063

Phylogenetic reconstruction of scleraxonian octocorals

supports the resurrection of the family Spongiodermidae
(Cnidaria, Alcyonacea)

Stephen D. Cairns A,B and Herman H. Wirshing A

A
Department of Invertebrate Zoology, National Museum of Natural History, Smithsonian Institution,
PO Box 37012, MRC 163, Washington, DC 20013-7012, USA.
B
Corresponding author. Email: [email protected]

Abstract. The Scleraxonia are a group of octocorals that share similarities of their axis morphology. However,
molecular phylogenetic analyses have shown this group to be largely polyphyletic. As a result, there is a significant lack
of understanding of what constitutes distinct evolutionary units among members of this group, particularly at the family
level. Prompted by the discovery of an unknown spongiodermid scleraxonian octocoral (Anthothelidae) from shallow
water off the Pacific coast of lower Baja California, a phylogenetic analysis of the undescribed specimen, together with
members of six scleraxonian families and an additional 29 non-scleraxonian octocorallian families was performed. Two
mitochondrial loci (mtMutS and COI) and one nuclear locus (28S) supported a monophyletic spongiodermid clade
(Homophyton, Callipodium, Diodogorgia, Titanideum and Sclerophyton, gen. nov.) at the family-level. The unknown
scleraxonian was supported as a new genus and species within the spongiodermid clade, sister to the western Atlantic
genus Titanideum. A morphological examination of the taxa within this clade revealed shared morphological
similarities in solenial (boundary) canals, and medullar and cortical sclerites. A revision, with illustrations, of the
Spongiodermidae was performed. Similar to previous studies, this study underscores the importance of combined
morphological and molecular analyses in order to resolve unstable systematic relationships among octocorals.

Received 10 December 2014, accepted 6 February 2015, published online 28 August 2015

Introduction Alcyoniina’ clade; McFadden et al. 2006). In addition,

Octocorals are a taxonomically diverse group of mostly colonial scleraxonians from the families Coralliidae and Paragorgiidae
benthic marine organisms that are found worldwide, from shallow group in a separate ‘Anthomastus–Corallium’ clade, while
water to the deep sea (Daly et al. 2007). They possess polyps with Briareum (Briareidae) and Erythropodium caribaeorum
eight tentacles, and produce skeletal components that are mostly (Anthothelidae) fall outside the three principal octocorallian
comprised of calcitic material. The Scleraxonia is a group of clades (sensu McFadden et al. 2006; see also McFadden and
gorgonian (or branching) octocorals consisting of seven families van Ofwegen 2012, 2013).
that are recognised by the presence of an axis composed of The discovery of an undescribed shallow-water scleraxonian
fused or unfused sclerites (microscopic biomineralised skeletal octocoral from the lower Baja Peninsula (Pacific Ocean),
components) (Bayer 1981; Fabricius and Alderslade 2001; similar in morphology to the western Atlantic spongiodermid
Daly et al. 2007). Within the Scleraxonia, members of the Titanideum (Anthothelidae), prompted the need for a
family Anthothelidae Broch, 1916 belong to three subfamilies morphological and phylogenetic analysis of the subfamily
(Anthothelinae, Semperininae and Spongiodermatinae) (sensu Spongioderminae in the context of a larger comprehensive
Bayer 1981) that are characterised by an axis that contains two octocorallian phylogeny. In order to confirm the identity and
layers, an inner medulla and an outer cortex, separated by phylogenetic placement of the undescribed scleraxonian, a
boundary canals (Daly et al. 2007). Phylogenetic analyses of phylogenetic analysis was performed using representatives of
the Scleraxonia have revealed this group to be largely six of the eight putative scleraxonian families (Briareidae,
polyphyletic (Sanchez et al. 2003; McFadden et al. 2006). For Anthothelidae, Subergorgiidae, Paragorgiidae, Coralliidae and
example, many anthothelid taxa are spread throughout the Melitheidae) together with members of 29 non-scleraxonian
octocorallian tree, and members of the anthothelid subfamilies octocorallian families. In addition, a revision of the resurrected
Semperininae (Iciligorgia, Solenocaulon) and Anthothelinae family Spongiodermidae Wright & Studer, 1889 was completed,
(Anthothela) group within a large clade consisting of many and a new genus and species, Sclerophyton bajaensis, gen.
taxa from the Holaxonia and Alcyoniina (‘Holaxonia– nov., sp. nov., are described.

Journal compilation
CSIRO 2015 www.publish.csiro.au/journals/is

346 Invertebrate Systematics S. D. Cairns and H. H. Wirshing

Materials and methods USA). Cycle-sequenced products were purified using

Samples for genetic analyses Sephadex G-50 Fine (GE Healthcare, Piscataway, NJ, USA),
and DNA sequencing was performed on an ABI 3730 at the
Scleraxonian colonies were sampled from the collections of the Laboratories of Analytical Biology (NMNH). Sequence contigs
Smithsonian National Museum of Natural History (NMNH). were collated and edited using Geneious Pro 7.1.7 (Biomatters;
Representatives from six of the eight putative scleraxonian http://www.geneious.com/).
families (Briareidae, Anthothelidae, Subergorgiidae,
Paragorgiidae, Coralliidae and Melitheidae) were included
with the exception of Parisididae and Dendrobrachiidae Phylogenetic analyses
(Table 1). Members of the latter two families were either Nuclear and mitochondrial loci were analysed separately to
unavailable or were not suitable for genetic analysis. If determine if the phylogenetic signal between the two datasets
members of a particular taxonomic group did not yield were similar. The two mitochondrial loci were analysed together
suitable amounts of genomic DNA for downstream analyses, as a single concatenated dataset (mtMutS+COI), and the nuclear
sequences from GenBank were used, if available. In order locus (28S) was examined alone. Because many of the museum-
to robustly interpret phylogenetic relationships among the extracted scleraxonian samples did not amplify well for 28S,
Scleraxonia, members of 29 octocorallian families representing resulting in a limited dataset for this locus compared with the
the three octocorallian orders (Pennatulacea, Helioporacea mitochondrial dataset, a complete concatenated analysis was
and Alcyonacea) and alcyonacean suborders or ‘grades/’ not performed. Protein coding mitochondrial loci were aligned
(Scleraxonia, Holaxonia, Calcaxonia, Stolonifera and using MUSCLE (multiple sequence comparison by log-
Alcyoniina, with the exception of Protoalcyonaria) (Daly et al. expectation) (Edgar 2004), and rDNA was aligned using
2007) were used for phylogenetic analyses. MAFFT (Katoh and Kuma 2002). MrAIC (Nylander 2004),
using the Akaike information criterion (AIC), was used to
Molecular analyses – DNA extraction, amplification, determine the best model of nucleotide sequence evolution for
sequencing each locus – GTR+G+I for all loci.
Three to four polyps, or ~4–5 mm3 of tissue were used for DNA Phylogenetic reconstruction was performed on two datasets,
extraction using a DNeasy blood and tissue kit (Qiagen, Inc., mtMutS+COI and 28S, using Bayesian inference (BI)
Valencia, CA, USA). Final elution was in 60 mL of manufacturer- performed with MrBayes 3.1.2 (Ronquist and Huelsenbeck
provided AE buffer, and samples were stored at
20C. Dried or 2003), maximum likelihood (ML) with RAxML (Stamatakis
ethanol-preserved colonies (70% to 95% EtOH) were used, 2006), and maximum parsimony (MP) with PAUP* v4.0b10
although higher DNA yields were generally obtained from (Swofford 2004). All analyses were run on the Topaz Cluster,
colonies collected within 10 years and/or stored in 95% which is maintained at the Smithsonian NMNH. Bayesian
EtOH. For colonies collected >10 years ago and stored in 70% inference analyses consisted of two independent runs with four
EtOH, dried colonies of the same taxa tended to be better suited chains, with trees sampled every 100th generation. Markov chain
for DNA extraction. The polymerase chain reaction (PCR) Monte Carlo (MCMC) runs were carried out for 12 million
was used to amplify two mitochondrial loci – mtMutS, generations for the mtMutS+COI dataset, and 10 million
AnthoCorMSH: 50 -AGG AGA ATT ATT CTA AGT ATG G- generations for the 28S dataset. The mtMutS+COI dataset was
30 (Herrera et al. 2010), Mut3458R: 50 -TSG AGC AAA AGC partitioned into character sets for each locus, and the model of
CAC TCC-30 (France and Hoover 2001); and COI, COI-LA- sequence evolution chosen by MrAIC applied to each partition.
8398-F: 50 -GGA ATG GCG GGG ACA GCT TCG AGT ATG All model parameters (tratio, statefreq, shape, pinvar) were
TTA ATA CGG-30 , COIoct-R: 50 -ATC ATA GCA TAG ACC unlinked, and the rate prior (prset ratepr) was set to ‘variable’.
ATA CC-30 (McFadden et al. 2011) – and one nuclear locus – 28S, To increase the efficiency with which the four chains exchanged
Far: 50 -CAC GAG ACC GAT AGC GAA CAA GTA-30 , Rab: information (up to 62% for mtMutS+COI, and 50% for 28S),
50 -TCG CTA CGA GCT TCC ACC AGT GTT T-30 (McFadden the heating parameter (lset temp) was set to 0.04 and 0.08,
and van Ofwegen 2012). respectively. Convergence was determined when the average
Polymerase chain reaction amplifications used final standard deviation of split frequencies was <0.01 and the
concentrations of 3 pmol of each primer, 500 mM dNTPS, potential scale reduction factor (PSRF) was 1.00. Tracer v1.5
3 mM MgCl and 1–5 mL of genomic DNA (varying final (Rambaut and Drummond 2009) was used to confirm the
elution concentrations) using 0.05 U mL–1 of Biolase DNA stationarity of the runs, and to verify that an adequate number
polymerase (Bioline, Inc., Taunton, MA, USA) and of trees were sampled from the posterior distribution (effective
manufacturer-provided buffers. Sterile DNA-grade water was sample size (ESS) > 4500 for all parameters). To calculate
added to make final reaction volumes of 10 or 20 mL. Thermal posterior probabilities, a ‘burn-in’ of 25% adequately removed
cycler reaction parameters included an initial denaturing step of trees before convergence. Maximum parsimony analyses
94C for 3 min, followed by 35 or 40 rounds of 94C for 30 s, using PAUP* v4.0b10 utilised heuristic searches with default
annealing temperatures of either 48C, 50C or 55C for 30 s or options. Maximum likelihood with RAxML was run using rapid
1 min, an extension at 74C for 30 s or 1 min, and a final extension bootstrap analysis and search for best-scoring ML tree (-f m)
step of 72C for 5 min. Polymerase chain reaction products and the GTRGAMMA model (-m). One thousand bootstrap
were neutralised using ExoSAP-IT (Affymetrix, Santa Clara, replicates were performed with MP and ML.
CA, USA), and prepared for sequencing using BigDye Three outgroups were used for each dataset. Montastraea
Terminater v3.1 (Applied Biosystems, Grand Island, NY, franksi (Scleractinia) (GenBank accession # NC007225),
Spongiodermidae and phylogeny of Scleraxonia Invertebrate Systematics 347

Table 1. Taxonomic information and GenBank accession numbers for octocoral taxa and outgroups used for phylogenetic analyses
Taxa in bold represent newly sequenced individuals (this study); voucher numbers for these specimens are in parentheses following their genus/species
identification

Family Genus/species GenBank accession

COI mtMutS 28S rDNA
Acanthoaxiidae Acanthoaxis wirtzi JX203816 JX203764 JX203635
Acanthogorgiidae Acanthogorgia breviflora GQ342378 GQ342464 JX203636
Acrossotidae Acrossota amboinensis GQ342379 DQ985956 JX203637
Alcyoniidae Acrophytum claviger JX203823 JX203770 JX203655
Alcyoniidae Alcyonium coralloides GQ342380 GQ342465 JX203640
Alcyoniidae Anthomastus ritteri JX203824 DQ302816 JX203761
Alcyoniidae Cladiella sphaerophora GQ342386 GQ342471 JX203653
Alcyoniidae Discophyton rudyi GQ342387 DQ302808 JX203659
Alcyoniidae Eleutherobia aurea JX203817 JX203766 GQ377454
Alcyoniidae Klyxum utinomii GQ342392 GQ342476 JX203654
Alcyoniidae Lampophyton planiceps GQ342393 GQ342477 JX203656
Alcyoniidae Lobophytum pauciflorum GQ342394 DQ280575 JX203649
Alcyoniidae Malacacanthus capensis GQ342395 DQ302811 JX203660
Alcyoniidae Nephthyigorgia sp. JX203864 JX203804 JX203732
Alcyoniidae Paraminabea aldersladei JX203819 JX203767 JX203763
Alcyoniidae Rhytisma fulvum GQ342396 GQ342478 JX203728
Alcyoniidae Sarcophyton ehrenbergi JX203821 DQ280516 JX203650
Alcyoniidae Sinularia querciformis GQ342399 FJ621469 JX203652
Alcyoniidae Thrombophyton coronatum GQ342400 DQ302814 JX203661
Anthothelidae Alertigorgia sp. JX203825 JX203771 JX203662
Anthothelidae Anthothela cf. grandiflora (USNM1095147) KP714035 KP713985 KP714021
Anthothelidae Anthothela sp. (USNM1139023) KP714036 KP713986 KP714022
Anthothelidae Anthothela sp. (USNM83402) KP714037 KP713987 KP714023
Anthothelidae Erythropodium caribaeorum GQ342401 GQ342480 JX203664
Anthothelidae Erythropodium caribaeorum (USNM1017543) KP714048 KP713998 n/a
Anthothelidae Erythropodium caribaeorum (USNM81337) KP714049 KP713999 n/a
Anthothelidae Iciligorgia brunnea (USNM86848) KP714050 KP714000 KP714024
Anthothelidae Iciligorgia sp. GQ342402 GQ342481 JX203665
Anthothelidae Solenocaulon sp. GQ342404 GQ342483 JX203668
Arulidae Arula petunia JX203827 JX203773 JX203670
Briareidae Briareum asbestinum GQ342405 GQ342484 JX203669
Briareidae Briareum asbestinum (USNM1122631) KP714038 KP713988 n/a
Chrysogorgiidae Radicipes gracilis HM590861 DQ297424 n/a
Chrysogorgiidae Stephanogorgia faulkneri GQ342406 GQ342485 JX203718
Chrysogorgiidae Trichogorgia capensis JX203863 JX203798 JX203719
Clavulariidae Azoriella bayeri GQ342407 GQ342486 JX203672
Clavulariidae Carijoa riisei JX203829 JX203775 JX203673
Clavulariidae Cervera atlantica JN620805 JN620804 JX203677
Clavulariidae Clavularia sp. JX203834 JX203778 JX203678
Clavulariidae Inconstantia exigua JX203870 JX203790 JX203690
Clavulariidae Incrustatus comauensis GQ342391 GQ342475 JX203691
Clavulariidae Knopia octocontacanalis GQ342410 GQ342488 JX203692
Clavulariidae Paratelesto sp. GQ342411 GQ342489 JX203693
Clavulariidae Telestula sp. JX203846 DQ302803 JX203697
Coelogorgiidae Coelogorgia palmosa GQ342413 DQ302805 JX203698
Coralliidae Corallium medea (USNM1110365) KP714040 KP713990 n/a
Coralliidae Corallium rubrum (USNM45614) KP714041 KP713991 n/a
Coralliidae Paracorallium tortuosum (USNM1072445) KP714051 KP714001 n/a
Coralliidae Paracorallium tortuosum (USNM1196760) KP714052 KP714002 KP714025
Cornulariidae Cornularia pabloi JX203847 JX203792 JX203699
Ellisellidae Ellisella sp. JX203850 JX203793 JX203702
Ellisellidae Viminella sp. JX203852 JX203794 JX203703
Gorgoniidae Eugorgia multifida GQ342417 GQ342494 JX203706
Gorgoniidae Eunicella tricoronata JX203853 JX203795 JX203707
Gorgoniidae Gorgonia flabellum GQ342418 GQ342495 JX203708
Gorgoniidae Leptogorgia rigida GQ342420 GQ342496 JX203709
(continued next page )
348 Invertebrate Systematics S. D. Cairns and H. H. Wirshing

Table 1. (continued )

Family Genus/species GenBank accession

COI mtMutS 28S rDNA
Gorgoniidae Pacifigorgia media GQ342421 GQ342497 JX203710
Gorgoniidae Pinnigorgia flava GQ342422 GQ342498 JX203711
Gorgoniidae Pseudopterogorgia bipinnata GQ342423 GQ342499 JX203712
Gorgoniidae Pterogorgia anceps GQ342424 GQ342500 JX203714
Gorgoniidae Rumphella sp. JX203855 JX203797 JX203715
Helioporidae Heliopora coerulea GQ342426 DQ302872 JX203716
Ifalukellidae Ifalukella yanii GQ342427 GQ342501 JX203717
Isididae Acanella eburnea EF672731 EF672731 n/a
Melithaeidae Acabaria erythraea GQ342430 GQ342503 JX203720
Melithaeidae Acabaria sinaica GQ342431 GQ342504 JX203721
Melithaeidae Clathraria rubrinodis GQ342432 GQ342505 JX203722
Melithaeidae Melithaea sp. JX203856 JX203799 JX203723
Melithaeidae Wrightella coccinea JX203858 JX203801 n/a
Nephtheidae Eunephthya thyrsoidea JX124384 JX124364 JX124340
Nephtheidae Gersemia rubiformis GQ342390 GQ342474 JX203648
Nephtheidae Paralemnalia thyrsoides GQ342436 GQ342509 JX203727
Nidaliidae Chironephthya sp. GQ342440 GQ342513 JX203730
Nidaliidae Pieterfaurea khoisanianum GQ342437 GQ342510 JX203657
Paragorgiidae Paragorgia alisonae (USNM1126251) KP714053 KP714003 n/a
Paragorgiidae Paragorgia arborea (USNM1092768) KP714054 KP714004 KP714026
Paragorgiidae Paragorgia arborea (USNM1120404) KP714055 KP714005 KP714027
Paragorgiidae Paragorgia johnsoni (USNM73767) KP714056 KP714006 KP714028
Paragorgiidae Paragorgia regalis (USNM1072340) KP714057 KP714007 n/a
Paragorgiidae Paragorgia sp. (USNM1075751) n/a n/a KP714031
Paragorgiidae Paragorgia sp. (USNM1075751) KP714059 KP714009 n/a
Paragorgiidae Paragorgia sp. (USNM1123838) KP714060 KP714010 n/a
Paragorgiidae Paragorgia sp. (USNM1164628) KP714058 KP714008 n/a
Paragorgiidae Sibogagorgia cauliflora (USNM54831) KP714062 KP714012 n/a
Paragorgiidae Sibogagorgia dennisgordoni (NIWA3328) KP714063 KP714013 n/a
Paralcyoniidae Paralcyonium spinulosum JX124389 DQ302833 JX124347
Paralcyoniidae Studeriotes sp. GQ342443 GQ342515 JX124348
Pennatulidae Gyrophyllum sibogae JX203865 DQ302869 JX203740
Plexauridae Astrogorgia sp. JX203861 JX203805 JX203733
Plexauridae Bebryce sp. JX203862 JX203806 JX203734
Plexauridae Eunicea tourneforti GQ342445 GQ342517 n/a
Plexauridae Menella sp. GQ342447 GQ342519 JX203736
Plexauridae Muricea atlantica GQ342448 GQ342520 JX203747
Plexauridae Muriceopsis flavida GQ342449 GQ342521 JX203744
Plexauridae Plexaura kuna JX203866 JX203807 JX203748
Plexauridae Plexaurella nutans GQ342451 GQ342523 JX203745
Plexauridae Pseudoplexaura wagenaari GQ342452 GQ342524 JX203746
Primnoidae Callogorgia formosa GQ342453 GQ342525 JX203749
Primnoidae Thouarella grasshoffi FJ268636 GQ868334 n/a
Protoptilidae Distichoptilum gracile GQ342454 DQ302866 JX203739
Renillidae Renilla sp. GQ342455 GQ342526 n/a
Spongiodermatidae Callipodium (YPM8594) KP714039 KP713989 n/a
Spongiodermatidae Diodogorgia ceratosa (USNM1147814) KP714042 KP713992 n/a
Spongiodermatidae Diodogorgia nodulifera JX203826 JX203772 JX203663
Spongiodermatidae Diodogorgia nodulifera (USNM59128) KP714043 KP713993 n/a
Spongiodermatidae Diodogorgia nodulifera (USNM78673) KP714044 KP713994 n/a
Spongiodermatidae Diodogorgia nodulifera (USNM1147817) KP714045 KP713995 n/a
Spongiodermatidae Diodogorgia nodulifera (USNM1147822) KP714046 KP713996 n/a
Spongiodermatidae Diodogorgia nodulifera (USNM1147872) KP714047 KP713997 n/a
Spongiodermatidae Homophyton verrucosum GQ342403 GQ342482 JX203666
Spongiodermatidae Sclerophyton bajaensis (colony 1a) (USNM1246058) KP714067 KP714017 KP714030
Spongiodermatidae Sclerophyton bajaensis (colony 1b) (USNM1246058) KP714068 KP714018 n/a
Spongiodermatidae Sclerophyton bajaensis (colony 2) (USNM1246058) KP714069 KP714019 n/a
Spongiodermatidae Titanideum frauenfeldii FJ264916 GU563314 n/a
(continued next page )
Spongiodermidae and phylogeny of Scleraxonia Invertebrate Systematics 349

Table 1. (continued )

Family Genus/species GenBank accession

COI mtMutS 28S rDNA
Spongiodermatidae Titanideum frauenfeldii (USNM49587) KP714065 KP714015 n/a
Spongiodermatidae Titanideum frauenfeldii (USNM60410) KP714066 KP714016 n/a
Subergorgiidae Annella mollis (USNM76231) KP714032 KP713982 n/a
Subergorgiidae Annella ornata (USNM75884) KP714033 KP713983 KP714020
Subergorgiidae Annella reticulata (USNM100821) KP714034 KP713984 n/a
Subergorgiidae Annella mollis GQ342456 JX203808 JX203750
Subergorgiidae Rosgorgia inexpetata (USNM1013161) KP714061 KP714011 KP714029
Subergorgiidae Subergorgia suberosa (USNM75340) KP714064 KP714014 n/a
Subergorgiidae Subergorgia suberosa GQ342457 JX203809 JX203762
Tubiporidae Tubipora sp. GQ342458 JX203811 JX203752
Virgulariidae Virgularia schultzei GQ342459 GQ342527 JX203743
Xeniidae Sarcothelia edmondsoni JX203868 JX203814 JX203757
Xeniidae Xenia hicksoni GQ342463 GQ342529 JX203759
Outgroups
Actiniaria Metridium senile AF000023 n/a n/a
Actiniaria Nematostella vectensis n/a n/a AY345871
Antipatharia Leiopathes glaberrima FJ597644 n/a
Antipatharia Leiopathes cf. expansa n/a n/a FJ626241
Scleractinia Montastraea franksi NC007225 n/a AY026375

Leiopathes glaberrima (Antipatharia) (FJ597644) and Metridium members of the families Subergorgiidae (Subergorgia, Annella
senile (Actiniaria) (AF000023) were used for the MutS+COI and Rosgorgia) and Anthothelidae (Erythropodium, Anthothela,
dataset. For 28S, Nematostella vectensis (Actiniaria) Solenocaulon, Iciligorgia and Alertigorgia) were both
(AY345871), Montastraea franksi (Scleractinia) (AY026375) polyphyletic, with the anthothelid genera distributed within
and Leiopathes cf. expansa (Antipatharia) (FJ626241) were used. and outside the HA clade. Members of the Coralliidae and
Paragorgiidae grouped within a well-supported clade that
Morphological analyses showed a close phylogenetic affinity to Anthomastus ritteri
Standard morphological terminology for octocorals is used, (Alcyoniidae) and Paraminabea aldersladei (Alcyoniidae)
based on the glossary of Bayer et al. (1983). In addition, the (sensu AC clade; McFadden et al. 2006) with the mtMutS
abbreviation L : D is used to indicate the ratio of the length to +COI dataset (only Anthomastus ritteri with 28S). With
diameter of a sclerite. mtMutS+COI, the representative paragorgiids were not
monophyletic; both species of Sibogagorgia (S. dennisgordoni
Results and S. cauliflora) did not group with the Paragorgia spp., but
were sister to a clade containing the Paragorgia species and
Genomic DNA (gDNA) yields from NMNH scleractinian
the coralliids Paracorallium tortuosum, Corallium rubrum and
octocorals varied considerably. The greatest yields tended to
C. medea (Fig. 1A).
be from samples stored in 95% EtOH and/or collected within
Members of the anthothelid subfamily Anthothelinae
10 years. Genomic DNA was reliably recovered from dried
(Anthothela, Alertigorgia and Erythropodium) were polyphyletic
specimens >10 years old; however, fungal contamination was
with both datasets. The Anthothela species grouped together
common, and likely contributed to higher total gDNA yields as
within the HA clade but did not group with Alertigorgia sp.,
fungal nuclear rDNA was often preferentially amplified in many
and Erythropodium caribaeorum was an unresolved lineage
dried specimens. As a result, many scleraxonian specimens
outside the HA clade. Members of another anthothelid
were not sequenced for octocoral-specific 28S.
subfamily, Semperininae (Iciligorgia and Solenocaulon),
showed a close mutual phylogenetic relationship, and together
Phylogenetic reconstruction – Scleraxonia
with Azoriella bayeri and Cervera atlantica (both clavulariids)
Analyses of mitochondrial (mtMutS+COI) and nuclear (28S) were sister to a monophyletic melithaeid clade. Briareum
loci each recovered phylogenies congruent with previous asbestinum (family Briareidae) fell outside the HA clade and
studies (e.g. McFadden et al. 2006; McFadden and van grouped as a sister taxon to members the Spongiodermidae
Ofwegen 2012 2013) (Fig. 1). Both phylogenies contained (formerly in the anthothelid subfamily Spongioderminae).
supported clades consistent with the ‘Holaxonia–Alcyoniina’
(HA), ‘Anthomastus–Corallium’ (AC) and ‘Calcaxonia–
Pennatulacea’ (CP) clades discussed in McFadden et al. Spongiodermidae Wright & Studer,
(2006); however, the AC and CP clades were unresolved with 1889 (resurrected family)
28S (Fig. 1B). Neither dataset recovered a monophyletic With both mitochondrial and nuclear datasets, members of
Scleraxonia, but instead scattered many of the families and the family Spongiodermidae (Callipodium, Diodogorgia,
genera throughout the octocorallian phylogeny. For example, Homophyton, Titanideum and Sclerophyton, gen. nov.)
350 Invertebrate Systematics S. D. Cairns and H. H. Wirshing

formed a well-supported clade outside the HA clade that was their boundary canal systems in cross section (Fig. 3), more
sister to Briareum asbestinum. This clade was unresolved likely points to a lack of genetic resolution, or ‘false
among the other octocorallian taxa at deeper nodes with negatives’ (McFadden et al. 2011), among these taxa.
mtMutS+COI (Fig. 1A), but was sister to the HA clade with However, the minimal variation among sclerite morphologies
28S (Fig. 1B). Phylogenetic relationships among the between Sclerophyton and Titanideum makes these taxa
Spongiodermidae with mtMutS+COI were largely consistent difficult to distinguish (see ‘Taxonomy’). Nevertheless,
across all the reconstruction methods, and differed only with keeping S. bajaensis, sp. nov. in a separate genus sister to
the placement of Homophyton verrucosum with maximum Titanideum is likely a better representation of the biodiversity
parsimony (MP) and likelihood (ML and MB) analyses and evolutionary history within the Spongiodermidae.
(Fig. 2). For example, with BI and ML (Fig. 2A), In addition to the spongiodermids, and similar to other
H. verrucosum was sister to all of the spongiodermid studies (e.g. Aguilar-Hurtado et al. 2012; McFadden and van
genera. With MP (Fig. 2B), H. verrucosum was sister to a Ofwegen 2012, 2013; Reijnen et al. 2014), the melithaeids
clade containing Callipodium pacificum and Diodogorgia also formed a monophyletic clade. This clade was part of a
nodulifera. The remaining taxa, Titanideum frauenfeldii and larger well-supported clade containing three anthothelid taxa
Sclerophyton bajaensis, sp. nov., grouped in a separate sister (Iciligorgia, Solenocaulon and Alertigorgia) and two
clade. With all methods, Sclerophyton bajaensis, sp. nov. was alcyoniids (Azoriella and Cervera), and suggests a close
sister to Titanideum frauenfeldii, and Callipodium pacificum phylogenetic relationship among these morphologically
was sister to Diodogorgia nodulifera. disparate taxa. Increased taxon sampling, specifically with the
addition of the monotypic melithaeid genus Asperaxis (sensu
Reijnen et al. 2014), and additional morphological analyses
Discussion need to be performed to clarify the relationships among the
Phylogenetic analysis of taxa representing six scleraxonian taxa found in this clade.
families recovered several polyphyletic clades scattered Like other studies that have examined upper-level
throughout a larger octocorallian phylogeny. However, one of systematic relationships among octocorals (e.g. McFadden
the clades was consistent with a monophyletic Spongioderminae et al. 2006), this study highlights the limitation, yet relative
(subfamily within the Anthothelidae), but was resolved at the usefulness, of standard molecular markers and well-sampled
family level. In addition, this clade was sister to Briareum taxa to resolve enigmatic systematic relationships among
asbestinum, a member of the family Briareidae. This octocorals. Undoubtedly, the addition of novel markers,
relationship could be interpreted as B. asbestinum belonging most likely through the use of next-generation sequencing
to a more inclusive Spongiodermidae (resurrected family). techniques, will help resolve many of the recalcitrant
However, the genetic distance (uncorrected p) of B. asbestinum groups within the octocorallian phylogeny and enhance our
relative to the other spongiodermids was 3.9%, a value that understanding of this ecologically important and biologically
may seem low, but can be interpreted as relatively high for diverse group of marine organisms.
mitochondrial DNA in octocorals (McFadden et al. 2011; see
also Shearer et al. 2002). Additionally, the sclerite morphology
of B. asbestinum is generally distinct from those of the Taxonomy
spongiodermids. Therefore, these two indices suggest that
B. asbestinum most likely belongs to an evolutionary linage Order ALCYONACEA
that is sister to, but separate from, the spongiodermids at the
family level. Subordinal group SCLERAXONIA
The pairwise genetic distance between S. bajaensis, sp. nov.
and its sister taxon, Titanideum, was 0.9%. This value is above Family SPONGIODERMIDAE Wright & Studer, 1889
what is generally found for intraspecific variation in octocorals
(0.5%), but within the range of what can be considered as Spongioderminae Wright & Studer, 1889: xxxi (in part: not Iciligorgia).
separate congeneric species (McFadden et al. 2011). This Briareidae: Kükenthal, 1924: 9 (in part).
Spongiodermatidae Aurivillius, 1931: 16 (in part: not Suberia).
suggests that S. bajaensis, sp. nov. could be a species of
Spongiodermatinae Bayer, 1955: 208; 1956: F196; 1981: 945.—
Titanideum; however, the genetic distance between Callipodium Williams, 1992a: 186.
and Diodogorgia, also inter-oceanic taxa and with greater Type genus: Spongioderma Kölliker, 1871 (=Homophyton).
morphological differentiation, was 0.6%. Thus, although a
strict interpretation of an octocoral ‘barcode’ suggests minimal
taxonomic diversity among these four genera (Titanideum, Diagnosis
Sclerophyton, Callipodium and Diodogorgia), the presence of Monomorphic Scleraxonia with its medulla surrounded by a
discernable contrasting morphologies, most notably among ring of longitudinal solenial (boundary) canals; a secondary

Fig. 1. Molecular phylogenies: (A) mtMutS+COI; (B) 28S. Scleraxonian taxa are colour coded by family (see legend), and newly sequenced individuals
contain their USNM ID. Each phylogeny was generated using Bayesian inference (BI), maximum likelihood (ML) and maximum parsimony (MP). Node
support for BI, ML and MP is shown from left to right. Clades with support <50 for maximum likelihood and parsimony, and <0.80 for Bayesian posterior
probabilities were collapsed to the next well-supported clade; ‘–’ represents less than the threshold for that method or not following the branching topology for
that method.
Spongiodermidae and phylogeny of Scleraxonia Invertebrate Systematics 351

(A)

Fig. 1. (continued)
352 Invertebrate Systematics S. D. Cairns and H. H. Wirshing

(B)

Fig. 1. (continued)
Spongiodermidae and phylogeny of Scleraxonia Invertebrate Systematics 353

(A) Homophyton verrucosum (B) Homophyton verrucosum

Callipodium pacificum (YPM8594)
70
Callipodium pacificum (YPM8594)
Diodogorgia nodulifera (USNM78673)
1.00/99 1.00/100 Diodogorgia nodulifera (USNM78673)
Diodogorgia nodulifera (USNM59128)
100
Diodogorgia nodulifera (USNM59128)
Diodogorgia nodulifera (USNM1147817)
Diodogorgia nodulifera (USNM1147817)
Diodogorgia nodulifera (USNM1147872)
1.00/99 97
Diodogorgia nodulifera (USNM1147822) Diodogorgia nodulifera (USNM1147872)
.95/66 99
Diodogorgia nodulifera (USNM1147822)
Diodogorgia ceratosa (USNM1147814)
Diodogorgia ceratosa (USNM1147814)
Diodogorgia nodulifera
Sclerophyton bajaensis (colony 1a) (USNM1246058) Diodogorgia nodulifera
1.00/100 Sclerophyton bajaensis (colony 1a) (USNM1246058)
Sclerophyton bajaensis (colony 1b) (USNM1246058) 96
Sclerophyton bajaensis (colony 1b) (USNM1246058)
Sclerophyton bajaensis (colony 2) (USNM1246058)
1.00/98
Sclerophyton bajaensis (colony 2) (USNM1246058)
Titanideum frauenfeldii 97
1.00/72
63 Titanideum frauenfeldii
Titanideum frauenfeldii (USNM49587)
1.00/100 100 Titanideum frauenfeldii (USNM49587)
Titanideum frauenfeldii (USNM60410)
Titanideum frauenfeldii (USNM60410)
0.01

Fig. 2. Phylogenies depicting phylogenetic relationships among the Spongiodermidae with the mtMutS+COI dataset. (A) Phylogram generated by Bayesian
inference (BI) and maximum likelihood (ML). (B) Cladogram generated by maximum parsimony (MP). With BI and ML, Homophyton verrucosum was sister
to all of the spongiodermid genera. With MP, H. verrucosum was sister to a clade containing Callipodium pacificum and Diodogorgia nodulifera. See Fig. 1
for node support explanation.

ring canal system may also separate inner from outer cortex. scleraxonian families being the unconsolidated axis (medulla)
Medullar sclerites stout rods, often bifurcate, branched, and/or of its genera.
spiny; cortical sclerites always include radiates, and may include The first modern account of the scleraxonian families was
spindles, ovals and double heads. that of Aurivillius (1931), who divided the suborder into eight
families or subfamilies, a classification that is currently used.
He also provided useful keys to the scleraxonian families and
Remarks genera within each family. He elevated the spongiodermids to
Three of the five previously known spongiodermid genera the family rank for the first time and considered it to be a new
were described between 1864 and 1869, another in 1919, and family, but according to the International Code of Zoological
the fifth in 1955, thus the generic complement of the family Nomenclature, this must be considered as an elevation of
has been fairly stable for a long time and subject to the evaluation Wright and Studer’s (1889) subfamily Spongioderminae and
of the four comprehensive revisions of the Octocorallia, i.e. thus take Wright and Studer’s name and date. The correct
Wright and Studer (1889), Kükenthal (1924) and Bayer spelling of the elevation to family status would then be
(1956, 1981). Nonetheless, Kölliker (1871) was the first to Spongiodermidae, not Spongiodermatidae, as later advocated
notice the resemblance of Titanideum and Spongioderma by Aurivillius (1931) and Bayer (1956). Aurivillius (1931)
(=Homophyton), but placed them in the associated defined the family as having monomorphic polyps, an axis
scleraxonian family Briareidae. (medulla) that was non-jointed, not cemented together and not
In the first comprehensive classification of the Octocorallia, penetrated by solenial canals (instead the canals form rings in
Wright and Studer (1889) divided the scleraxonian section of the cortex), and with at least some cortical radiate sclerites.
the Gorgonacea into four families, one of these four families, Despite Aurivillius’ reasoning, Bayer (1955, 1956, 1981),
the Briareidae, with two subfamilies, the Briareinae and without explanation, downgraded the spongiodermids to the
the Spongioderminae, the latter containing Titanideum, subfamily grade, sharing the family Anthothelidae with the
Spongioderma (=Homophyton) and Iciligorgia. Thus, the subfamilies Anthothelinae and Semperininae. Essentially he
proper authorship of the family-group name Spongiodermidae distinguished the Spongioderminae from the other two
should be attributed to Wright and Studer (1889), not Aurivillius subfamilies by its cortical radiates and branching medullar
(1931), as is common. Subfamily Briareinae was distinguished rods. Contemporary classifications (e.g. Williams and Cairns
from the Spongioderminae by having solenial canals in the 2014) have followed that classification.
medulla, the latter not. A similarity to Callipodium was However, as discussed above, genetic analysis of mitochondrial
mentioned, but that encrusting genus was placed by Wright and nuclear loci support the spongiodermid genera (with the
and Studer (1889) in the rather distant family Cornulariidae in exception of Tripalea, not used in this study) as a monophyletic
the order Alcyonacea. clade at the family level, that is sister to the family Briareidae.
Kükenthal’s (1924) revision of the Gorgonaria was somewhat Thus, the family level taxon Spongiodermidae is resurrected, as
of a step backwards in that he lumped 14 scleraxonian genera first suggested by Wright and Studer (1889). There are now
into the Briareidae, including those now assigned to the nine families considered to fall within the scleraxonian group:
Spongiodermidae, Briareidae, Anthothelidae and Paragorgiidae, six families with an unconsolidated axis (Briareidae,
the character distinguishing this family from the other three Anthothelidae, Spongiodermidae, Subergorgiidae, Paragorgiidae
354 Invertebrate Systematics S. D. Cairns and H. H. Wirshing

(A) Sclerophyton (B) Titanideum

Medulla
Medulla

Cortex

Cortex

(C) Diodogorgia (D) Homophyton

Medulla Medulla

Cortex Cortex

(E) Tripalea

Medulla

Cortex

Fig. 3. (A–E) Diagrammatic representations of the relative sizes and positions of the
medulla, cortex, and inner and outer circles of ring canals.

and Sibogagorgiidae (resurrected family; Figueroa and Baco Diagnosis

2015)) and three with consolidated axes (Coralliidae, Colonies unbranched or sparsely branched in dichotomous
Melithaeidae and Parisididae). fashion. Outer cortex composed of 3- to 8-radiates; inner
cortex composed of larger 4-, 5-, 6-, 7- and 8-radiates, often
with an elongate neck region connecting sets of tubercles.
Medulla composed of closely packed longitudinally arranged
Genus Sclerophyton, gen. nov. rods, often with bifurcating tips and occasional side branches;
Type species: Sclerophyton bajaensis, here designated. contiguous medullar sclerites often fused. Two longitudinal rings
urn:lsid:zoobank.org:act:15B67DCE-7ED8-47E2-B565-87F030 of boundary canals present, one delimiting outer from inner
CC37A4 cortex, the other delimiting the inner cortex from the medulla.
Spongiodermidae and phylogeny of Scleraxonia Invertebrate Systematics 355

Remarks diameter of the lower stem reaches up to 12 mm. Calyces flush

The morphological differences between Titanideum and with branch. Colonies pinkish-orange in colour, but a thin
Sclerophyton are small, primarily a slight difference in the encrusting sponge (Fig. 6C) gives most of colony a white
sclerite component of the cortex (i.e. Titanideum has double patina. Thin outer cortex (Fig. 6B, C, E) densely packed with
heads and Sclerophyton does not). Also, Sclerophyton often 3- to 8-radiates, but primarily 3- and 4-radiates that are
has fused medullar sclerites (Titanideum does not), and the 0.068–0.110 mm in length; inner cortex (Fig. 6F) composed of
longitudinal canals of the two ring systems of Sclerophyton larger (0.106–0.172 mm long) 4- to 8-radiates, but somewhat
are uniform in diameter, whereas those of the inner ring of elongate, sets of tubercles separated by a short neck region; L : D
Titanideum appear to be slightly larger. Morphologically, it is ratio up to 1.6. Medullar region composed of densely packed,
a subjective opinion whether the new taxon should be a new longitudinally arranged rods (Fig. 6D, G) 0.18–0.27 mm in
species of Titanideum or a separate genus. There is no quantitative length. These rods usually distally bifurcate and may also have
morphological measure to support either option – just a feeling short lateral branches; they are relatively smooth, with few
borne of experience working with octocorals and shared by tubercles. Contiguous medullar sclerites often laterally fused
taxonomists such as Deichmann, Bayer, Verrill, Kükenthal, (Fig. 6D) conferring a greater strength to branch axis. Cortical
Stiasny, Kölliker and Verseveldt. However, the genetic distance sclerites orange, whereas medullar sclerites colourless. Outer
for mtMutS+COI between Sclerophyton and Titanideum (0.9%) ring canal system (Figs 3A, 6A–C) lies just beneath (0.18 mm)
is greater than that between Callipodium and Diodogorgia branch surface and is fairly well defined. Inner ring system
(0.6%) (see ‘Discussion’), the latter two taxa with more (Figs 3A, 6C), which encircles medulla, sometimes indistinct
discernable morphological differences. Therefore, the molecular but always present. Canals of both systems of same relatively
evidence suggests that Sclerophyton and Titanideum are most small size: 0.10–0.14 mm in diameter.
likely two separate genera.
Distribution
Other species As for the genus.
Monotypic.
Etymology
Distribution Named for the region from which it was collected.
Pacific side of southern Baja California, 15–18 m.
Genus Titanideum Verrill, 1864
Etymology (Figs 3B, 4D, 7A–H)
From the Greek root scleros (meaning hard, and also the basis for Titanideum Verrill, 1864a: 10; 1864b: 39; Kölliker, 1871: 16; Wright &
the suborder Scleraxonia) + phyton (meaning plant, a common Studer, 1889: xxxiii; Kükenthal, 1919: 81, 82; 1924: 30; Deichmann,
suffix in scleraxonian genera). A fungus has the same name 1936: 83; Bayer, 1956: F196; 1961: 77; 1981: 945.
Sclerophyton Eschw., 1824, but the name is still available for Type species: Gorgonia suberosa Ellis & Solander, 1786, by monotypy
(=Solanderia frauenfeldii Kölliker, 1865). The name of G. suberosa
an animal taxon.
Ellis & Solander, 1786 is not nomenclaturally available because it is
a junior homonym of Gorgonia suberosa Pallas, 1766 (=Subergorgia
Sclerophyton bajaensis, sp. nov. suberosa). The next available junior synonym is Solanderia
(Figs 3A, 4B, C, 6A–G) frauenfeldii Kölliker, 1865.

urn:lsid:zoobank.org:act:195BE748-591F-4EEB-A1E5-9238 Diagnosis
8BB26AD1 Colonies unbranched or sparsely branched from basal region,
with long ascending branches. Outer cortex composed of 3-
Material examined and 4-radiates and double heads; inner cortex composed of 4-,
Holotype: Turtle Bay, Baja California (Pacific side, approx. 27380 N, 5-, 6-, 7- and 8-radiates, these sclerites often elongate with a
114540 W), MEXICO, 15–18 m, xi.1959, USNM 1246057, coll. James neck region connecting sets of tubercles. Medulla composed of
Stewart. Paratypes: from type locality, six colonies and 23 branches, closely packed longitudinally arranged rods, often with bifurcate
USNM 1246058; Punta Hughes, Baja California (Pacific side, near tips and occasionally short medial branches. Two longitudinal
Magdalena Bay), MEXICO, 18 m, viii.1957, three colonies, USNM rings of boundary canals present, one delimiting outer from inner
1246059, coll. James Stewart. cortex, the other delimiting the inner cortex from the medulla.
Type locality Description of type species
Turtle Bay, Baja California (Pacific side), Mexico, 15–18 m. Colonies unbranched or sparsely branched, most long ascending
branches originating near colony base (Fig. 4D); colonies reach
Description up to 31 cm in height and 15 mm in basal branch diameter.
Colonies unbranched (Fig. 4B) or sparsely, dichotomously Calyces usually flush with branch surface. Colonies usually
branched (Fig. 4C), the latter with U-shaped axils, and in both uniformly yellow or orange, the latter sometimes with pale
cases the colony reaches up to 28 cm in height. Branch tips blunt yellow calyces, or pinkish-red, or rarely white. Cortex sclerites
to slightly clavate. Colonies have a thin encrusting base, the pink, medullar sclerites colourless. Thin outer cortex densely
356 Invertebrate Systematics S. D. Cairns and H. H. Wirshing

(A) (B)

(C) (D)

Fig. 4. (A) Diodogorgia nodulifera, USNM 49705, Palm Beach, Florida, 73 m. (B) Sclerophyton bajaensis, USNM 1246059,
paratype. (C) Sclerophyton bajaensis, USNM 1246057, holotype. (D) Titanideum frauenfeldii, USNM 60410, Cape Hatteras,
beach.

packed with 3- and 4-radiates (Fig. 7C, F) and double heads (Figs 3B, 7B), consisting of 10–12 contiguous, relatively large
(Fig. 7G), the former 0.038–0.09 mm in length, the latter (0.15 mm in diameter) canals that border the medulla. Outer ring
0.044–0.068 mm in length. Inner cortex composed of larger 4- canal system somewhat indistinct (Figs 3B, 7A), consisting of
to 8-radiates (0.11–0.19 mm in length), the tubercles often more numerous, smaller (0.075–0.10 mm in diameter), non-
separated by a short neck region and thus approximating a contiguous canals lying only ~0.15 mm beneath branch
tuberculate spindle (Fig. 7H); these sclerites have an L : D ratio surface, distinguishing outer from inner cortex.
of up to 1.6. Relatively small medullar region composed of
densely packed longitudinally arranged rods (Fig. 7B, D, E)
0.22–0.36 mm in length. These rods usually distally bifurcate Remarks
and may also have short lateral branches; rods relatively smooth Among the four spongiodermid genera with two longitudinal
and have few tubercles. Inner ring canal system well defined ring canal systems (Table 2), Titanideum is unique in having
Spongiodermidae and phylogeny of Scleraxonia Invertebrate Systematics 357

(A) (B)

(C)

(F )
(D)

(E)

Fig. 5. (A) Diodogorgia nodulifera, USNM 1147814, Discovery Bay, Jamaica, 82 m. (B) Callipodium pacificum, YPM 588,
Panama, syntype. (C) Tripalea clavaria, USNM 43421, 36420 S, 56230 W, 21 m. (D) Homophyton verrucosum, USNM 1091355,
29270 S, 31310 E, 68 m. (E) Callipodium rubens, USNM 50523, off Freeport, Texas, 9.1 m. (F) Tripalea clavaria, MNRJ 5478,
–2358’06.0000'', -04529’26.4000'' 2358.10 S, 4529.440 W, 45 m.

double heads in its outer cortex. It is further compared to The genus Paratitanideum Kükenthal, 1919: 84 (in part,
Sclerophyton in the account of that genus. type species Titanideum friabilis Nutting, 1911, by subsequent
Useful descriptions, comments and/or illustrations of the designation (Kükenthal 1924)) was placed in the Semperinidae
type species can be found in: Verrill (1864a), Kükenthal by Aurivillius (1931).
(1919, 1924), Deichmann (1936), Bayer (1961), Cairns and
Bayer (2009), and DeVictor and Morton (2010). Its Other species
distribution is: North Carolina to Cuba, and south-east Gulf of ?Titanideum obscurum Thomson, 1927: Azores, 1250 m.
Mexico, 13–293 m. Although not examined, the illustrations in the original
358 Invertebrate Systematics S. D. Cairns and H. H. Wirshing

(A) (B)

(C) (D)

(E) (F )

(G)

Fig. 6. (A–G) Sclerophyton bajaensis, paratype, USNM 1246058: (A) stereo view of branch cross section showing inner and outer ring canals; (B) stereo view
of polyp cross section in the outer cortex; (C) stereo view of branch cross section showing cortex, medulla, ring canals and sponge sclerites encrusting the
branch; (D) stereo view of interlocking branched medullar rods; (E) 3- to 8-radiates from outer cortex; (F) 4- to 8-radiates with a short neck region, from inner
cortex; (G) medullar sclerite.
Spongiodermidae and phylogeny of Scleraxonia Invertebrate Systematics 359

(A) (B)

(C) (D) (E )

(F ) (H)

(G)

Fig. 7. (A–G) Titanideum frauenfeldii, USNM 49587, off Little River Reef, Florida, 14 m; USNM 8348, 35120 3000 N, 75250 W (Cape Hatteras), 88 m: (A) stereo
view of branch cross section showing inner and outer ring canals and longitudinal section of five polyps; (B) stereo view of medulla surrounded by inner ring
canal; (C) stereo view of radiates in outer cortex; (D) branched medullar rods in situ; (E) branched medullar rods; (F) 3- and 4-radiates from outer cortex;
(G) double heads from outer cortex; (H) 4- to 8-radiates with neck region from inner cortex.
360 Invertebrate Systematics S. D. Cairns and H. H. Wirshing

radiates (no distinction

Girdled and branched
spindles (L : W up to
description show elongate spindles in the cortex and no radiates,

3.8); ovals; 7-, 8-

between layers)
which would not be consistent with this genus. It is also quite

No ring canals
deep for this family.

Bifurcate rods
Callipodium
Encrusting

Distribution
Western Atlantic (North Carolina to Cuba, Gulf of Mexico),
?Azores, 13–293-?1250 m.
up to 5.5), vacuous
Spiny spindles (L : W
Tuberculate spindles,

Genus Diodogorgia Kükenthal, 1919

Unbranched, clavate

(Figs 3C, 4A, 5A, 8A–H)

One inner ring
Bifurcate rods
spheroids

Diodogorgia Kükenthal, 1919: 96, 97; 1924: 36; Aurivillius, 1931: 16, 18,
Tripalea

19; Deichmann, 1936: 85, 86; Stiasny, 1937: 67; Bayer, 1956: F196;
Bayer, 1961: 71, 72; Bayer, 1981: 945; Williams, 1992a: 193.
Type species: Diodogorgia ceratosa Kükenthal, 1919, by subsequent
designation (Kükenthal, 1924) (=Solandaria nodulifera Hargitt in
Hargitt & Rogers, 1901 (senior synonym); =Solanderia cristata
Hargitt in Hargitt & Rogers, 1901; =Diodogorgia cervicornis
indistinct with small
Two rings: inner ring
Short, bifurcate rods
sparsely branched

outer ring distinct

Kükenthal, 1919; =Corallium vanderbilti Boone, 1933).

and canals much
diameter canals,
Ovals (L : W ~2)
3- to 8-radiates,
Unbranched or

Diagnosis
Homophyton

spheroids
Table 2. Comparison of the spongiodermatid genera

Colonies regularly and dichotomously branched in a flabellate

larger

fashion, or unbranched (i.e. D. capensis). Outer cortex composed

of small 3-, 4-, 5- and 6-radiates, tuberculate spheroids, and short
tuberculate spindles; inner cortex composed of longer tuberculate
spindles. Medulla composed of closely packed, longitudinally
spindles (L : W up to

arranged rods, only rarely bifurcate at tips. Two well-defined

Rods, rarely bifurcate
spindles; spheroids
Dichotomous, planar

3-, 4-, 5-, 6-radiates;

longitudinal rings of solenial boundary canals present in

Two discrete rings
Long tuberculate

branches, one ring between outer and inner cortex (termed the
tuberculate
Diodogorgia

‘lateral canals’ by Verseveldt 1940), the other between the inner

cortex and medulla (termed the ‘median canals’ by Verseveldt
5.6)

1940).

Description of type species

Colonies abundantly and dichotomously branched (Figs 4A, 5A);
3-, 4-radiates; double

with a neck region

sparsely branched

(L : W up to 1.55)

branches often slightly clavate. Individual colonies up to 25 cm in

Two discrete rings,
inner ring canals

height, with a basal branch diameter up to 15 mm. Calyces form

4- to 8-radiates
Unbranched or

Bifurcate rods

small protuberances even when contracted, thus the name

from base
Titanideum

nodulifera. Most common colour of corallum yellow with red

heads

large

or purplish-red polyps, but other variations include uniformly

yellow, red or pink. Most sclerites red, those of calyces usually
yellow. Thin outer cortex densely packed with 4-, 5- and 6-
radiates 0.065–0.110 in diameter (Fig. 8D), even smaller
tuberculate spheroids (Fig. 8F) ~0.050 mm in diameter, and
Two rings, outermost
Sparse dichotomous

with neck region

sometimes fused
(L : W up to 1.6)

short tuberculate girdled spindles (Fig. 8E) up to 0.15 mm in

(2- or 4-radiates
most common)

4- to 7-radiates,

length. Rather abruptly at outer (lateral) boundary ring canal, the

3- to 8-radiates

Bifurcate rods,
Sclerophyton

sclerite composition changes to larger tuberculate spindles

indistinct

(Fig. 8G) some of which are up to 0.60 mm in length and with

an L : D ratio of up to 5.5. Inner medullar region composed of
densely packed, longitudinally oriented rods (Fig. 8A–C, H)
0.23–0.57 mm in length and rarely if ever bifurcate. Medullar
rods smooth compared with cortical spindles, but do occasionally
Longitudinal canals

have superficial tubercles. Medullar rods do not fuse, except

within holdfast, where they form a somewhat continuous
Colony shape

Outer cortex

foundation. Polyp sclerites similar to those of outer cortex.

Inner cortex

Two rings of longitudinal solenial canals well defined and

Medulla

their adjacent canals within a ring are closely spaced (Figs 3C,
8A–C), canals of both rings circular to elliptical in outline and
Spongiodermidae and phylogeny of Scleraxonia Invertebrate Systematics 361

(A) (B)

(C) (G)

(D)

(H)

(E)

(F)

Fig. 8. (A, B, D, G, H) Diodogorgia nodulifera, USNM 49705 (see above – C, E, F, USNM 59128) off Little Cayman, 21.3 m: (A) stereo view of branch cross
section showing inner and outer ring canals, cortex and medulla; (B) stereo view of enlargement of cortex and part of inner and outer ring canal system; (C) stereo
view of branch longitudinal section showing cortex and medulla; (D) 3- to 6-radiates from outer cortex; (E) tuberculate spindles from outer cortex; (F) two
tuberculate spheroids from outer cortex; (G) elongate tuberculate spindles from inner cortex; (H) medullar rods.
362 Invertebrate Systematics S. D. Cairns and H. H. Wirshing

about the same size in both rings, i.e. 0.18–0.20 mm in greater cylinders up to 3 mm in height and ~1.7 mm in diameter, and
diameter. often of a different colour from branch, but they may also be
flush with branch surface when retracted. Colonies uniformly
Remarks yellow, pinkish-white, reddish-orange or dark red; calyces
Among the four spongiodermid genera with two longitudinal either white, yellow or red (Williams 1992a). Thin outer
ring canal systems (Table 2), Diodogorgia differs in having cortex (Fig. 9A, B) composed of 3- to 8-radiates (Fig. 9D)
well-developed tuberculate spindles in its cortex, as well as 0.048–0.12 mm in length or diameter. Inner cortex composed
various classes of radiates. of much larger and thicker sclerites called ovals (Fig. 9C, E) up
Useful descriptions, comments and/or illustrations of the to 0.28 mm in length and with an L : D ratio of only 2. Large
type species can be found in: Kükenthal (1919), Deichmann medullar region (Fig. 9A) composed of relatively short
(1936), Stiasny (1937), Verseveldt (1940), Bayer (1959, 1961), (0.12–0.29 mm in length), densely packed, longitudinally
Cairns and Bayer (2009), and DeVictor and Morton (2010). The arranged rods (Fig. 9F) that are relatively smooth and often
account by Verseveldt (1940) is by far the most exhaustive. distally bifurcate. Inner ring canal system that occurs at
Indeed, this is one of the better known and discussed octocoral boundary of medulla and inner cortex is indistinct, the small
species in the western Atlantic. DeVictor and Morton (2010) diameter canals measuring only ~0.16 mm in diameter. The
were the first and only to illustrate sclerites with SEM, which outer ring canal system, which separates the inner from the
we add to with stereo views of the ring canal, sclerites in situ, outer cortex, quite distinct (Figs 3D, 9A, B), composed of
and the various sclerite types. Its distribution includes, from large (up to 0.5 mm in diameter) canals.
north to south: Jacksonville, Florida, Bahamas, Greater and
Lesser Antilles, Panama and the northern coast of South Remarks
America to Surinam (Bayer 1961) at depths of 36–183 m. Among the four spongiodermid genera with two longitudinal
ring canal systems (Table 2), Homophyton is distinguished by
Other species having ovals in its inner cortex, and much larger diameter
Diodogorgia capensis (Thomson, 1911): off South Africa, canals in its outer ring than its inner ring.
80–90 m (see Williams 1992b: 397, fig. 23A). Homophyton was resurrected by Bayer (1981) to replace
?Diodogorgia sibogae Stiasny (1941: 67): South Africa, Spongioderma, based on priority of description.
Indonesia, 73 m. Useful descriptions, comments and/or illustrations of the
type species can be found in: Möbius (1861), Gray (1866),
Distribution Kükenthal (1919, 1924) and Williams (1992a, 1993), the most
complete accounts provided by Kükenthal (1919) and Williams
Western Atlantic (from Florida throughout Greater and Lesser
(1992a).
Antilles, eastern Gulf of Mexico, Panama, northern coast of
S. America to Surinam), South Africa, ?Indonesia; 36–183 m. Other species
Genus Homophyton Gray, 1866 The genus is monotypic. Juncella elongata var. capensis
Hickson, 1904, once placed in Spongioderma, is now placed
(Figs 3D, 5D, 9A–F)
in Ideogorgia (see Williams 1992a), and Spongioderma chunii
Solanderia: Möbius, 1861: 3. Kükenthal, 1908 is considered to be a junior synonym of
Homophyton Gray, 1866: 27, fig. 2; Bayer, 1981: 945; Williams, 1992a: H. verrucosum.
186, 187.
Spongioderma Kölliker, 1871: 14 (type species: Solanderia verrucosa Distribution
Möbius, 1861, by monotypy); Wright & Studer, 1889: xxxiv;
Kükenthal, 1919: 89–91; 1924: 34; Aurivillius, 1931: 18; Bayer, South Africa, southern Mozambique, 0–168 m (Williams 1992b:
1956: F196. 397, fig. 24b).
Type species: Homophyton gattyiae Gray, 1866, by monotypy
(=Solanderia verrucosa Möbius, 1861, the senior synonym). Genus Tripalea Bayer, 1955
(Figs 3E, 5C, F, 10A–F)
Diagnosis
Suberia Studer, 1878: 666 (in part: S. clavaria); Kükenthal, 1919: 85
Colonies unbranched or sparsely dichotomously branched. Outer (in part); 1924 (in part).
cortex composed of small 3- to 8-radiates and tuberculate Tripalea Bayer, 1955: 208, 209; Bayer, 1956: F196; 1961: 70; 1981: 911,
spheroids; inner cortex composed of much larger ovals. 945.
Medulla composed of closely packed longitudinally arranged Type species: Suberia clavaria Studer, 1878, by original designation.
rods, often bifurcate at their ends. Two longitudinal rings of
boundary canals present, the inner ring indistinct. Diagnosis
Colonies unbranched, branch tips clavate. Outer cortex composed
Description of type species of tuberculate spheroids and tuberculate and girdled spindles;
Colonies unbranched or sparsely branched in a dichotomous inner cortex spongy, composed of spiny spindles. Medulla
fashion (Fig. 5D), colonies attaining a height of 42 cm with a composed of closely packed longitudinally arranged rods,
basal branch diameter of 15 mm. Branch tips attenuate to blunt often bifurcate at their ends. Only one longitudinal ring canal
(not clavate). Calyces often protuberant (Fig. 5D), expressed as system, located at the boundary of medulla and inner cortex.
Spongiodermidae and phylogeny of Scleraxonia Invertebrate Systematics 363

(A) (B)

(D) (C)

(F )

(E)

Fig. 9. (A–E) Homophyton verrucosum, USNM 1091355 (see above), (F) USNM 53882, 31380 4800 S, 29340 2400 E, 48 m. (A) Stereo view of branch cross
section showing large canals of outer ring and much smaller canals of inner ring, and central medulla; (B) Ssereo view of branch cross section showing
boundary between outer and inner cortex and two large longitudinal ring canals of outer ring; (C) stereo view of an inner cortex oval; (D) 3- to 8-radiates from
outer cortex; (E) ovals from inner cortex; (F) medullar rods.
364 Invertebrate Systematics S. D. Cairns and H. H. Wirshing

(A) (B)

(C)
(E)

(D)

(F )

Fig. 10. (A, B, F) Tripalea clavaria, USNM 43421 (see above), (C–E) USNM 84390, Mar de Plata, depth unknown. (A) Stereo longitudinal view of distal
branch showing several polyps and a longitudinal canal; (B) stereo view of branch cross section showing single ring canal system, medulla and cortex;
(C) girdled spindles from outer cortex; (D) tuberculate spheroid from outer cortex; (E) spiny spindles from inner cortex; (F) medullar rods.
Spongiodermidae and phylogeny of Scleraxonia Invertebrate Systematics 365

(A) (C)

(B)

(D)

(E )

(F )

Fig. 11. (A, C–F) Callipodium rubens, USNM 50523 (see above), (B) Callipodium pacificum, YPM 588, syntype. (A) Stereo view of girdled tuberculate
spindles from cortex; (B) 7- to 8-radiates from cortex; (C) tuberculate girdled spindles; (D) branched spindles from cortex; (E) ovals from cortex; (F)
medullar rod?
366 Invertebrate Systematics S. D. Cairns and H. H. Wirshing

Description of type species unilinearly arranged on stoloniferous colonies. Uppermost

Colonies unbranched and up to 26 cm in length, with an encrusting layer of cortex composed of small (0.07–0.16 mm in diameter)
base, a basal branch diameter up to 8 mm, and a terminal clavate 7- or 8-radiates, below which are short (0.13–0.32 mm in length),
branch tip up to 12 mm in diameter (Fig. 5C, F). Calyces flush with straight (Fig. 11C) and branched girdled spindles, as well as
branch surface. Colony and sclerites white. Outermost cortex ovals up to 0.29 mm in length, with an L : D ratio of ~1.6. Medulla
composed of small (0.038–0.056 mm in diameter) tuberculate and ringed solenial canals not detectable.
spheroids (Fig. 10D) and short (0.069–0.120 mm in length)
girdled or belted spindles (Fig. 10C). The inner cortex, which is Remarks
thick and spongy (Fig. 10A), consists of slender spiny spindles Callipodium is unique among the spongiodermid genera in
(Fig. 10E) 0.15–0.26 mm in length, with an L : D ratio of up to 5.5. having an encrusting growth form, branched spindles, and
Relatively small medullar region composed of densely packed, lacking longitudinal ring canals (Table 2).
longitudinally arranged rods 0.14–0.22 mm in length and often Useful descriptions, comments and/or illustrations of the
bifurcate at their tips (Fig. 10F). Only one ring of longitudinal type species only can be found in Verrill (1869). Callipodium
boundary canals present (Figs 3E, 10B), located between medulla pacificum is known from the eastern Pacific from the Pearl
and cortex, composed of relatively few (8–10) large (up to 0.35 mm Islands to Peru at intertidal depths.
in diameter) tubes that are circular to elliptical in cross section. Useful references to Anthopodium rubens can be found in:
Smaller (0.12 mm diameter) longitudinal canals occur throughout Verrill (1872), Deichmann (1936) and Bayer (1961). It is known
the cortex but are not arranged in rings (Fig. 3E). from off North Carolina and Texas, at 9–92 m (Cairns and Bayer
2009). Bayer (1961) described a ‘basement’ or medullar zone
Remarks for this species sometimes composed of fused rods.
Tripalea is distinctive among the subergorgiids in having only Verrill (1872: 434) stated that Anthopodium and Callipodium
one longitudinal ring canal system, clavate branches, and spiny had ‘very different spicula’ and thus kept them as separate
spindles in its inner cortex (Table 2). genera. Oddly, Kükenthal (1924) did not mention either genus.
Useful descriptions, comments and/or illustrations of the In Bayer’s (1956) classification, Callipodium is included but
type species can be found in: Studer (1878), Kükenthal (1919) Anthopodium not even mentioned as a junior synonym. Then
and Bayer (1961). five years later Bayer (1961) treated Anthopodium rubens and
stated that ‘the spicules of Anthopodium bear a closer
Other species resemblance to those of Callipodium than Verrill indicated,
although they are not identical’. Finally, in his classification of
The genus is monotypic. key to the genera, Bayer (1981) included in his key Anthopodium
(but not Callipodium) but in his summary classification (Bayer
Distribution 1981: 945) listed: ‘Callipodium (=Anthopodium?)’. Finally,
East coast of South America from southernmost Brazil (Rio unexplainably Anthopodium rubens was not included in the
Grande do Sul) to 40S off Argentina, 20–55 m. revision of DeVictor and Morton (2010). Thus, the treatment
of these two encrusting genera is quite inconsistent and
Genus Callipodium Verrill, 1869 sometimes contradictory. Nonetheless, Anthopodium rubens
(Figs 5B, E, 11A–F) has the same sclerite complement (Fig. 11A, C–F) as
Callipodium pacificum and thus we suggest accepting the
Sympodium: Verrill, 1866: 329 (in part: S. pacificum). synonymy of these two genera.
Callipodium Verrill, 1869: 455; Wright & Studer, 1889: xvi; Bayer, 1956:
F196; 1981: 945.
Other species
Anthopodium Verrill, 1872: 434 (type species: A. rubens Verrill, 1872 by
monotypy); Wright & Studer, 1889: xv; Bayer, 1961: 78, 79; 1981: 909, Callipodium aureum Verrill, 1869: Pacific Panama, depth not
945. stated.
Type species: Sympodium pacifica Verrill, 1866 by original designation. Callipodium rubens (Verrill, 1872): off North Carolina and
Texas, at 9–14 m.
Diagnosis Callipodium australe Verrill, 1876: Bluff Harbour, New
Colonies encrusting (Fig. 5B, E), forming a mat or a reticulate Zealand, depth not stated.
stoloniferous expansion. Sclerite complement consists of short
girdled spindles (Fig. 11A, C), branched spindles (Fig. 11D), Distribution
ovals (Fig. 11E), and 7- and 8-radiates (Fig. 11B). Medullar or North Carolina, northern Gulf of Mexico, eastern Pacific from
basement layer (Bayer 1961) may be present. Ring canals absent. Pearl Islands Panama to Peru, New Zealand, 0–92 m.
Description of type species Acknowledgements
Colonies usually encrust rocks and dead shells, forming a We wish to acknowledge Bob Ford, who photographed and constructed
continuous thin mat (up to 8 cm in width) or a stoloniferous the colour figures, Tim Coffer, who put together the SEM figures, and Taryn
expansion, which is sometimes reticulate. Colonies and Wolf, who assisted with the design of the cross sections. We also thank Eric
sclerites bright red, purplish-red or orange. Calyces often Lazo-Wasem, Yale Peabody Museum, for the loan of type specimens of
project from the surface as much as 0.10 mm, sometimes Callipodium and Anthopodium; Kareen Schnabel, National Institute of
Spongiodermidae and phylogeny of Scleraxonia Invertebrate Systematics 367

Water and Atmospheric Research, Wellington, for the loan of Sibogagorgia rearrangements, order reversals, and cnidarian phylogenetics. Genome
dennisgordoni; Clovis Castro, for the loan of Tripalea clavaria; and Biology and Evolution 7, 391–409. doi:10.1093/gbe/evu286
Francesca Leasi for technical advice. Finally we acknowledge Odalisca France, S. C., and Hoover, L. L. (2001). Analysis of variation in mitochondrial
Breedy, for help in the preliminary identification of the new species. DNA sequences (ND3, ND4L, MSH) among Octocorallia (=Alcyonaria)
(Cnidaria: Anthozoa). Bulletin of the Biological Society of Washington 10,
110–118.
References
Gray, J. E. (1866). Description of two new forms of gorgonioid corals.
Aguilar-Hurtado, C., Nonaka, M., and Reimer, J. D. (2012). The Melithaeidae Proceedings of the Zoological Society of London 1866, 24–27.
(Cnidaria: Octocorallia) of the Ryukyu Archipelago: molecular and Hargitt, C. W., and Rogers, C. G. (1901). The Alcyonaria of Porto Rico.
morphological examinations. Molecular Phylogenetics and Evolution Bulletin of the U. S. Fish Commission 20, 265–287.
64, 56–65. doi:10.1016/j.ympev.2012.03.005 Herrera, S., Baco, A., and Sánchez, J. A. (2010). Molecular systematics
Aurivillius, M. (1931). ‘The Gorgonarians from Dr. Sixten Bock’s Expedition of the bubblegum coral genera (Paragorgiidae, Octocorallia) and
to Japan and Bonin Islands 1914. Kungliga Svenska Vetenskaps- description of a new deep-sea species. Molecular Phylogenetics and
Akademiens Handlingar, Tredje Serien, Band 9.’ (Almqvist & Wiksell: Evolution 55(1), 123–135. doi:10.1016/j.ympev.2009.12.007
Stockholm.) Hickson, S. J. (1904). The Alcyonaria of the Cape of Good Hope. Part 2.
Bayer, F. M. (1955). Contributions to the nomenclature, systematics, and Marine Investigations of South Africa 3, 211–239.
morphology of the Octocorallia. Proceedings of the United States Katoh, M., and Kuma, M. (2002). MAFFT: a novel method for rapid multiple
National Museum 105, 207–220. doi:10.5479/si.00963801.105- sequence alignment based on fast Fourier transform. Nucleic Acids
3357.207 Research 30, 3059–3066. doi:10.1093/nar/gkf436
Bayer, F. M. (1956). Octocorallia. In ‘Treatise on Invertebrate Paleontology’. Kölliker, A. (1871). Beiträge zur Kenntniss der Polypen. Verhandlungen der
(Ed. R. C. Moore.) pp. F166–189, 192–231. (University of Kansas Press: Physikal-Medicin. Gesellschaft in Würzburg 2, 11–30.
Lawrence, KS.) Kükenthal, W. (1919). Gorgonaria. Wissenschaftliche Ergebnisse der
Bayer, F. M. (1959). Octocorals from Surinam and the adjacent coasts of Deutschen Tiefsee-Expedition auf dem Dampfer “Valdivia”, 1898–1899
South America. Studies on the Fauna of Suriname and Other Guyanas 6, 13, 1–946.
1–43. Kükenthal, W. (1924). ‘Coelenterata: Gorgonaria.’ Das Tierreich 47. (Walter
Bayer, F. M. (1961). The shallow-water Octocorallia of the West Indian de Gruyter & Co.: Berlin.) [In German]
region. Studies on the Fauna of Curaçao and Other Caribbean Islands McFadden, C. S., and van Ofwegen, L. P. (2012). Stoloniferous octocorals
12, 1–373. (Anthozoa, Octocorallia) from South Africa, with descriptions of a new
Bayer, F. M. (1981). Key to the genera of Octocorallia exclusive of family of Alcyonacea, a new genus of Clavulariidae, and a new species
Pennatulacea (Coelenterata: Anthozoa), with diagnoses of new taxa. of Cornularia (Cornulariidae). Invertebrate Systematics 26(4), 331–356.
Proceedings of the Biological Society of Washington 94, 902–947. doi:10.1071/IS12035
Bayer, F. M., Grasshoff, M., and Verseveldt, J. (1983). ‘Illustrated Trilingual McFadden, C. S., and van Ofwegen, L. P. (2013). Molecular phylogenetic
Glossary of Morphological and Anatomical Terms Applied to evidence supports a new family of octocorals and a new genus of
Octocorallia.’ (E. J. Brill: Leiden.) Alcyoniidae (Octocorallia, Alcyonacea). ZooKeys 346, 59–83.
Boone, L. (1933). Scientific results of the cruise of the yachts “Eagle” and doi:10.3897/zookeys.346.6270
“Ara” 1921–1928. Coelenterata, Echinodermata and Mollusca. Bulletin McFadden, C. S., France, S. C., Sánchez, J. A., and Alderslade, P. (2006).
of the Vanderbilt Marine Museum 4, 1–133. A molecular phylogenetic analysis of the Octocorallia (Cnidaria: Anthozoa)
Cairns, S. D., and Bayer, F. M. (2009). Octocorallia (Cnidaria) of the Gulf based on mitochondrial protein-coding sequences. Molecular Phylogenetics
of Mexico. In ‘Gulf of Mexico: Origin, Waters, and Biota’. (Eds and Evolution 41(3), 513–527. doi:10.1016/j.ympev.2006.06.010
D. L. Felder and D. K. Camp.) pp. 321–331. (Texas A&M Press: McFadden, C. S., Benayahu, Y., Pante, E., Thoma, J. N., Nevarez, P. A., and
College Station, TX.) France, S. C. (2011). Limitations of mitochondrial gene barcoding in
Daly, M., Brugler, M. R., Cartwright, P., Collins, A. G., Dawson, M. N., Octocorallia. Molecular Ecology Resources 11, 19–31. doi:10.1111/
Fautin, D. G., France, S. C., McFadden, C. S., Opresko, D. M., Rodriguez, j.1755-0998.2010.02875.x
E., Romano, S., and Stake, J. (2007). The phylum Cnidaria: a review of Möbius, K. (1861). Neue Gorgoniden des naturhistorischen Museums zu
phylogenetic patterns and diversity 300 years after Linnaeus. Zootaxa Hamburg. Nova Acta Academiae Caesareae Leopoldino Carolinae
1668, 1–766. Germanicae Naturae Curiosorum 29, 1–12.
Deichmann, E. (1936). The Alcyonaria of the western part of the Atlantic Nutting, C. C. (1911). The Gorgonacea of the Siboga Expedition. VIII. The
Ocean. Memoirs of the Museum of Comparative Zoology at Harvard Scleraxonia. Siboga-Expeditie, Monographie 13b5, 1–62.
College 53, 1–317. Nylander, J. A. A. (2004). ‘MrAIC.pl.’ Program distributed by the author.
DeVictor, S. T., and Morton, S. L. (2010). Identification guide to the (Evolutionary Biology Centre: Uppsala University.)
shallow water (0–200 m) octocorals of the South Atlantic Bight. Pallas, P. S. (1766). ‘Elenchus Zoophytorum.’ (Comitum, P. van Cleef:
Zootaxa 2599, 1–62. Hague.)
Edgar, R. C. (2004). MUSCLE: multiple sequence alignment with high Rambaut, A., and Drummond, A. J. (2009). ‘Tracer version 1.5.0.’ Available
accuracy and high throughput. Nucleic Acids Research 32(5), at http://tree.bio.ed.ac.uk/software/tracer/.
1792–1797. doi:10.1093/nar/gkh340 Reijnen, B. T., McFadden, C. S., Hermanlimianto, Y. T., and van Ofwegen,
Ellis, J., and Solander, D. (1786). ‘The Natural History of Many Curious L. P. (2014). A molecular and morphological exploration of the generic
and Uncommon Zoophytes ... Systematically Arranged and Described boundaries in the family Melithaeidae (Coelenterata: Octocorallia) and
by the late Daniel Solander.’ (Benjamin White and Son: London.) its taxonomic consequences. Molecular Phylogenetics and Evolution 70,
Fabricius, K., and Alderslade, P. (2001). ‘Soft Corals and Sea Fans – A 383–401. doi:10.1016/j.ympev.2013.09.028
Comprehensive Guide to the Tropical Shallow-water Genera of the Ronquist, F., and Huelsenbeck, J. P. (2003). MrBayes 3: Bayesian
Central-West Pacific, the Indian Ocean and the Red Sea.’ (Australian phylogenetic inference under mixed models. Bioinformatics 19,
Institute of Marine Science: Townsville.) 1572–1574. doi:10.1093/bioinformatics/btg180
Figueroa, D. F., and Baco, A. R. (2015). Octocoral mitochondrial genomes Sanchez, J. A., Lasker, H. R., and Taylor, D. J. (2003). Phylogenetic analyses
provide insights into the phylogenetic history of gene order among octocorals (Cnidaria): mitochondrial and nuclear DNA sequences
368 Invertebrate Systematics S. D. Cairns and H. H. Wirshing

(lsu-rRNA, 16S and ssu-rRNA, 18S) support two convergent clades Verrill, A. E. (1866). On the polyps and corals of Panama with descriptions
of branching gorgonians. Molecular Phylogenetics and Evolution 29, of new species. Proceedings of the Boston Society of Natural History 10,
31–42. doi:10.1016/S1055-7903(03)00090-3 323–333.
Shearer, T. L., Van Oppen, M. J. H., Romano, S. L., and Wörheide, G. (2002). Verrill, A. E. (1869). Review of the polyps and corals of the west coast of
Slow mitochondrial DNA sequence evolution in the Anthozoa (Cnidaria). America. Transactions of the Connecticut Academy of Arts and Sciences
Molecular Ecology 11, 2475–2487. doi:10.1046/j.1365-294X.2002. 1, 423–502.
01652.x Verrill, A. E. (1872). On Radiata from the coast of North Carolina.
Stamatakis, A. (2006). RAxML-VI-HPC maximum likelihood-based American Journal of Science and Arts 3(18), 432–438. doi:10.2475/
phylogenetic analyses with thousands of taxa and mixed models. ajs.s3-3.18.432
Bioinformatics 22(21), 2688–2690. doi:10.1093/bioinformatics/btl446 Verrill, A. E. (1876). Anthozoa. In ‘Contributions to Natural History of
Stiasny, G. (1937). Die Gorgonacea der Siboga-Expedition. Supplement II, Kerguelen Island’. (Ed. J. H. Kidder.) Bulletin of the U. S. National
Revision der Scleraxonia mit ausschuluss der Melitodidae und Museum 3, 76–77.
Coralliidae. Siboga-Expeditie, Monographie 13b8, 1–138. [In German] Verseveldt, J. (1940). Studies on Octocorallia of the families Briareidae,
Stiasny, G. (1941). Studien uber Alcyonaria und Gorgonaria (Parerga und Paragorgiidae and Anthothelidae. Temminkia 5, 1–142.
Paralipomena). II. Zoologischer Anzeiger 134, 53–71. Williams, G. C. (1992a). The Alcyonaria of Southern Africa. Gorgonian
Studer, T. (1878). Ubersicht der Steinkorallen aus der Familie der Octocorals (Coelenterata, Anthozoa). Annals of the South African
Madreporaria aporosa, Eupsammina, und Turbinaria, welche auf der Museum 101, 181–296.
Reise S. M. S. Gazelle um die Erde gesammelt wurden. Williams, G. C. (1992b). Biogeography of the octocorallian coelenterate
Monatsberichte der Königlich Preussischen Akademie der fauna of southern Africa. Biological Journal of the Linnean Society.
Wissenschaften zu Berlin 1877, 625–654. Linnean Society of London 46, 351–401. doi:10.1111/j.1095-8312.1992.
Swofford, D. L. (2004). ‘PAUP: Phylogenetic Analysis Using Parsimony, tb00869.x
version 4.0b10.’ (Sinauer Associates: Sunderland, MA.) Williams, G. C. (1993). ‘Coral Reef Octocorals: An Illustrated Guide to
Thomson, J. A. (1927). Alcyonaires provenant des campagnes scientifiques the Soft Corals, Sea Fans, and Sea Pens Inhabiting the Coral Reefs of
du Prince Albert 1er de Monaco. Résultats des Campagnes Scientifiques Northern Natal.’ (Natural Science Museum: Durban.)
accomplies sur son yacht par Albert 1er, Monaco 73, 1–77. Williams, G. C., and Cairns, S. D. (2014). ‘Systematic list of the valid
Thomson, J. A., and Mackinnon, D. L. (1911). The Alcyonarians of the octocoral genera.’ Available at http://researcharchive.calacademy.org/
“Thetis” expedition. Australian Museum Memoirs 4, 661–695. research/izg/OCTOCLASS.htm
doi:10.3853/j.0067-1967.4.1911.1509 Wright, E. P., and Studer, T. (1889). Report on the Alcyonaria collected by H.
Verrill, A. E. (1864a). List of the polyps and corals sent by the Museum of M.S. Challenger during the years 1873–76. Report on the Scientific
Comparative Zoology to other institutions in exchange, with annotations. Results of the Voyage of H.M.S. Challenger during the years 1873–76.
Bulletin of the Museum of Comparative Zoology 1, 29–60. Zoology 31, 1–314.
Verrill, A. E. (1864b). Revision of the polypi of the eastern coast of the
United States. Memoirs of the Boston Society of Natural History 1, 1–45.

www.publish.csiro.au/journals/is

View publication stats