AÇOREANA, 2014, Suplemento 10: 149-165
MORPHOLOGY AND SYSTEMATIC RELATIONSHIPS OF BROCCHINIA CLENCHI 
PETIT, 1986 (MOLLUSCA, CAENOGASTROPODA, CANCELLARIIDAE) 
FROM THE AZORES
M.G. Harasewych1 & Richard E. Petit2
1Dept. of Invertebrate Zoology, MRC-163, National Museum of Natural History, Smithsonian Institution, PO Box 
37012, Washington, D.C. 20013-7012 USA e-mail: Harasewych@si.edu
2806 St. Charles Road, North Myrtle Beach, SC 29582 USA
ABSTRACT
Data on the shell morphology, microstructure, anatomy, and DNA sequences of two 
mitochondrial genes (COI and 16S rDNA) are used to infer the phylogenetic relationships of 
Brocchinia clenchi Petit, 1986 within the morphologically diverse and taxonomically complex deep-
water genus Brocchinia. Initial molecular data show that this small, narrow-shelled species from 
the Atlantic is more closely related to a morphologically and ecologically similar species from the 
Indo-Pacifi c than either is to a representative of a broad-shelled group within the genus that is also 
present in the Atlantic and Indo-Pacifi c Oceans. While the thick shell, prominent columellar folds 
and apertural lirae of many Brocchinia species are features shared with members of the subfamily 
Cancellariinae, the absence of a radula is a characteristic of some, though not all Admetinae. 
Sequence data for the COI gene places Brocchinia within the Cancellariinae, while 16S rDNA data 
analyzed separately or together with COI places Brocchinia in a clade with Mirandaphera. This clade 
falls within Admetinae (16S only), or is separate from both Admetinae and Cancellariinae (COI + 
16S), but not with strong support. Neither morphological characters nor molecular data have yet 
been able to unambiguously place the genus Brocchinia within the family Cancellariidae.
RESUMO
Utilizaram-se dados sobre a morfologia da concha, microestrutura, anatomia e sequências de 
ADN de dois genes mitocondriais (COI e 16S rDNA) para inferir das relações fi logenéticas de 
Brocchinia clenchi Petit, 1986 dentro do morfologicamente diverso e taxonomicamente complexo 
género de águas profundas Brocchinia. Dados moleculares preliminares mostram que esta pequena 
espécie de concha estreita do Atlântico é mais relacionada com uma espécie do Indo-Pacífi co 
morfológica e ecologicamente similar do que ambas o são de um representante de um grupo de 
conchas largas dentro do género, também presente nos oceanos Atlântico e Indo-Pacífi co. Enquanto 
que a concha espessa, as dobras columelares proeminentes e as liras aperturais de muitas espécies 
de Brocchinia são características partilhadas com membros da subfamília Cancellariinae, a ausência 
de rádula é uma característica de alguns, embora não todos, Admetinae. Dados de sequência para o 
gene COI colocam Brocchinia com os Cancellariinae, enquanto que dados de 16S rDNA analizados 
separadamente ou juntamente com COI colocam Brocchinia num clado com Mirandaphera. Este clado 
está incluído em Admetinae (apenas 16S), ou é separado de Admetinae e de Cancellariinae (COI + 
16S), mas não com forte suporte. Nem os caracteres morfológicos nem os dados moleculares têm 
até agora sido capazes de colocar sem ambiguidade o género Brocchinia na família Cancellariidae.
INTRODUCTION
The genus Brocchinia was introduced by Jousseaume (1887: 221) to contain 
two small species of cancellariids 
with elongated shells, one, Brocchinia 
mitraeformis (Brocchi, 1814), a fossil 
from the Pliocene of Italy, the other, 
Brocchinia pusilla (H. Adams, 1869), a 
Recent species from bathyal depths off 
the Canary Islands. The fossil species, 
Brocchinia mitraeformis, was subsequently 
designated as the type species of the 
genus by Sacco (1894: 68) (Figures 1-3). 
As both taxa included by Jousseaume are 
junior homonyms, their nomenclature 
was clarifi ed by Petit (1986). 
While the family Cancellariidae 
Paginação_Açoreana_Suplemento 10_prova05.indd   149 05-10-2014   17:44:52
150 2014, Suplemento 10: 149-165A Ç O R E A N A
comprises a morphologically distinctive 
and easily recognized clade, the 
allocation of species among the more 
than 120 described genera (Petit & 
Harasewych, 2005) has often been 
diffi  cult and provisional, particularly 
when based solely on shell morphology, 
as is necessary in cases when the type 
species are fossils. Thirteen Recent species 
are currently aĴ ributed to Brocchinia 
(Table 1), eight from the Atlantic, fi ve 
from the Indo-Western Pacifi c, nearly all 
from bathyal depths. Many had originally 
been described in or subsequently 
assigned to other genera, among them 
Admete Kröyer in Möller, 1842, Anapepta 
Finlay, 1930, Cancellaria Lamarck, 1799, 
Gergovia Cossmann, 1899, Inglisella Finlay, 
1924, Merica H. & A. Adams, 1854, and 
Solutosveltia Habe, 1961.
The anatomy of Brocchinia remains 
largely unknown. Verhecken (2007: 
Table 1) reported that radulae are absent 
in B. nodosa and B. decapensis. In their 
description of B. canariensis, Rolán & 
Hernández (2009: 105, fig. 7) described the 
animal as being white, with black eyes, a 
fl aĴ ened and elongated penis, a gill of 9 
thick lamellae, and without a radula or 
operculum. Martins et al. (2009: pl. 13, 
fi g. 213) illustrated a living specimen of B. 
clenchi in color. 
In their landmark paper on the 
molecular phylogeny of Cancellariidae, 
Modica et al. (2011) included two species 
of Indo-Pacifi c Brocchinia. These authors 
noted that this genus had traditionally 
been classifi ed in the Cancellariinae 
(their table 1), but their results indicated 
that these two Brocchinia formed a 
TABLE 1. Recent species aĴ ributed to the genus Brocchinia Jousseaume, 1887 in the World Register 
of Marine Species (Bouchet & Gofas, 2013) *. 
Species Locality Depth (m)
Atlantic Species
Brocchinia azorica (Bouchet & Warén, 1985) Azores 1,220-2,900
Brocchinia canariensis Rolán & Hernández, 2009 Canary Islands 350-400
Brocchinia clenchi Petit, 1986 Azores, Josephine Bank, Canary Islands 15-1,520
Brocchinia decapensis (Barnard, 1960) Cape Point, South Africa 2,890-2,963
Brocchinia harasewychi Barros & Lima, 2007 Brazil 452-690
Brocchinia nodosa (Verrill & S. Smith, 1885) Eastern USA 678-1,950
Brocchinia pustulosa Verhecken, 1991 Brasil 637
Brocchinia verheckeni Barros & Lima, 2007 Brasil 365-690
Indo-Pacific Species
Brocchinia exigua (E.A. Smith, 1891) Australia 200-1,463
Brocchinia fi scheri (A. Adams, 1860) Korea to South Australia 40-1,017
Brocchinia kaiensis Verhecken, 1997 Indonesia 400
Brocchinia septentrionalis (Finlay, 1930) New Zealand 137
Brocchinia tanimbarensis Verhecken, 1997 Indonesia 300-500
*two additional, unnamed species from Indonesia have been illustrated and discussed by Verhecken (2011). 
  
Paginação_Açoreana_Suplemento 10_prova05.indd   150 05-10-2014   17:44:52
151HARASEWYCH & PETIT: BROCCHINIA CLENCHI PETIT, 1986 FROM THE AZORES
clade together with two species of 
Mirandaphera (possibly due to long branch 
aĴ raction). Although this clade was 
provisionally included in the subfamily 
Admetinae (their fi gures 3-5), the authors 
acknowledged that the relationship is not 
well supported (Modica et al., 2011: 691; 
Figs. 3, 4), and noted that this group “may 
represent an independent lineage … 
worthy of more detailed investigation.”  
Of the three species of Cancellariidae 
that occur in the Azores (Verhecken, 
2007), two have been referred to the genus 
Brocchinia. Brocchinia azorica (Bouchet & 
Warén, 1985), was originally described 
in the genus Admete, but transferred to 
Brocchinia by Verhecken (1991: 549). It is 
a wide ranging species (off  NW Ireland to 
the Azores, and possibly to South Africa) 
that inhabits lower bathyal to abyssal 
depths (1,220-2,900 m).  
In contrast, the range of Brocchinia 
clenchi Petit, 1986, is narrower, and 
includes the Macaronesian Archipelagos 
(Azores, Madeira, Canary Islands), the 
Josephine Bank and adjacent seamounts, 
and the coast off  Western Sahara, at depths 
of 15-1,520 m. Verhecken, (2007: 313) 
noted that specimens from shallower 
waters off  the Azores and Canary Islands 
tend to be smaller and smoother than the 
more strongly sculptured deeper water 
samples from the Josephine Banks and 
the Canary Islands, but provisionally 
regarded the variants to be conspecifi c in 
the absence of additional material. 
Brocchinia clenchi has a complex 
taxonomic history. In his description of 
Brocchinia clenchi, Petit (1986: 25) stated 
that this species had been previously 
described as Cancellaria pusilla Adams, 
1869, but that name is preoccupied 
by Cancellaria pusilla Sowerby, 1832. 
Although a specimen from the Canary 
Islands in the collection of The Natural 
History Museum (NHMUK 1855.4.4.202, 
ex McAndrew collection) was labeled 
as a “possible type” of Cancellaria pusilla 
Adams, Petit concluded that the type of 
C. pusilla Adams could not be positively 
identifi ed. Instead of designating a nomen 
novum for this species, Petit chose to 
redescribe the species so that a holotype 
could be designated, and included 
the NHMUK specimen as a paratype 
of Brocchinia clenchi. Cancellaria pusilla 
Adams, 1869 (type locality Canary Islands, 
depth unspecifi ed) was considered to 
be a synomym of Brocchinia clenchi (type 
locality Josephine Bank, 622-786 m). 
More recently, Rolán & Hernández 
(2009) described Brocchinia canariensis from 
slightly shallower depths (350-400 m) off  
the Canary Islands, diff erentiating it from 
the type specimens of B. clenchi primarily 
on the basis of it having a proportionally 
larger protoconch and smaller, narrower 
shell with a more pronounced suture 
and a proportionally smaller fi nal whorl. 
They did not include in their discussion 
or comparisons the 93 specimens taken off  
the Canary Islands at depths ranging from 
65-1,520 m that were listed by Verhecken 
(2007: 312). Rolán & Hernández incorrectly 
referred to the NHMUK specimen as the 
lectotype of Cancellaria pusilla Adams 
and acknowledged that it could be an 
abnormal form of B. canariensis. However, 
they (Rolán & Hernández, 2009: 105-106) 
chose not dispute its status as a paratype 
of B. clenchi. This ambiguity may lead to 
one of the following interpretations:
1) That two distinct upper 
bathyal species of Brocchinia (B. 
clenchi and B. canariensis) both occur 
in the Canary Islands. 
2) That these two species do not 
co-occur in the Canary Islands, in 
which case the NHMUK specimen 
is mis-aĴ ributed to B. clenchi, and 
Cancellaria pusilla Adams is actually 
a synonym of B. canariensis rather 
than of B. clenchi. 
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152 2014, Suplemento 10: 149-165A Ç O R E A N A
3) That only a single, 
morphologically variable, upper 
bathyal species of Brocchinia occurs 
throughout the broad geographic 
range aĴ ributed to B. clenchi by 
Verhecken (2007: 313), in which 
case, both Cancellaria pusilla and 
Brocchinia canariensis become 
synonyms of B. clenchi. 
FIGURES 1-6. Brochinia. 1-3, Brocchinia mitraeformis (Brocchi, 1814). Petit collection (#2126), Rio 
Torsero, Savona, Italy (Lower Pliocene). 1, Apertural, lateral and dorsal views of shell. 2, Apical 
and lateral views of the protoconch. 3, Detail of shell, showing piĴ ed surface (intriticalx?). 4-6. 
Brocchinia clenchi Petit, 1986. 4, Apertural, dorsal and apical views of the holotype, USNM 849002. 
5, Apertural, and 6, apical views of paratype, USNM 189694, both from Josephine Bank, due 
W of Cape St. Vincent, Portugal (~ 37°N, 14°W), in 622-786 m. Arrows indicate transition from 
protoconch to teleoconch.
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153HARASEWYCH & PETIT: BROCCHINIA CLENCHI PETIT, 1986 FROM THE AZORES
The present study documents the 
shell morphology, anatomy, biology 
and investigates the phylogenetic 
relationships (based on morphology 
and DNA sequences) of Brocchinia 
clenchi from the Sabrina Bank, off  São 
Miguel, Azores, in order to provide a 
basis for more comprehensive studies 
of the biogeography and evolutionary 
relationships within the genus Brocchinia. 
MATERIALS AND METHODS
Specimens of Brocchinia clenchi were 
collected from dredged sediment samples 
take off  Portinho Velho das Capelas, São 
Miguel, Azores. A sample from 50-100 m 
(Sta. SAB2011.DR2, 37°51’N, 25°40’W, 
July 7, 2011) (USNM 1197432) contained 
a single living specimen and four dead 
shells inhabited by sipunculans, as well 
as several shell fragments. A sample from 
200 m in the same area (Sta. SAB2011.
DR3, 37°51.280’N, 25°41.971’W, July 7, 
2011)(USNM 1197434) contained ten 
dead shells inhabited by sipunculans 
and several fragments. A third sample, 
from 200 m on the Sabrina Bank, off 
São Miguel, Azores (Sta. SAB2011.DR6, 
37°51.547’N, 25°41.706’W, July 8, 2011) 
(USNM 1197435) contained four living 
specimens as well as eight dead shells 
and fragments. Living animals and dead 
shells populated by sipunculans were 
preserved in 95% ethanol. Empty shells 
and fragments were preserved dry. 
Measurements and whorl counts for 
the holotype (USNM 849002) and paratype 
(USNM 189694) of Brocchinia clenchi as 
well as for ten Azorean shells (including 
two juveniles) were obtained from 
enlarged scanning electron micrographs 
of the apertural and apical views of the 
specimens using a digital caliper (Table 
2). Data for the holotype and paratype of 
Brocchinia canariensis and for Cancellaria 
pusilla Adams were obtained from 
published fi gures (Rolán & Hernández, 
2009: pls. 1-2). Data for Brocchinia 
mitraeformis was obtained from scanning 
electron micrographs of a specimen from 
the Petit collection (PC 2126). The number 
of protoconch whorls was counted using 
the method of Jablonski & Luĵ  (1980: 332) 
in order to be consistent with a prior study 
of cancellariid development (Pawlik et 
al., 1988). This method of measurement 
results in values that are ½ whorl greater 
than those produced by the method used 
by Verhecken (2007: fi g. 4). A dendrogram 
based on these measurements was 
produced using Dendro-UPGMA 
(Garcia-Vallve et al., 1999; hĴ p://genomes.
urv.cat/UPGMA/) using the Pearson (r) 
coeffi  cient.
Shells were cracked for four of the 
living animals. One male and one female 
were dissected. Following dissection, the 
uppermost whorls of the male and female 
specimens, comprising portions of the 
digestive gland and gonad, were used for 
DNA extraction, as were portions of the 
introvert of two specimens of sipunculans 
inhabiting empty Brocchinia clenchi 
shells. Using protocols identical to those 
documented for Cerion in Harasewych et 
al. (2011), portions of the mitochondrial 
cytochrome c oxidase I and 16S rDNA 
genes were sequenced for all four samples. 
A search of the GenBank nucleotide 
database using the Basic Local Alignment 
Search Tool (BLAST) identifi ed both 
sipunculan samples as Aspidosiphon 
(Aspidosiphon) muelleri Diesing, 1851, 
based on a 97% match of the COI 
sequences and a 99% match of the 16S 
sequences.
Sequences for the cytochrome c 
oxidase I and 16S rDNA genes from two 
specimens of Brocchinia clenchi (1♂ + 
1♀) and a representative selection of 14 
cancellariid taxa (Table 3) spanning the 
cancellariid phylogenetic tree (Modica et 
al., 2011: fi g. 5) were aligned against those 
of Bivetiella cancellata (Linnaeus, 1767) 
derived from its complete mitochondrial 
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154 2014, Suplemento 10: 149-165A Ç O R E A N A
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Paginação_Açoreana_Suplemento 10_prova05.indd   154 05-10-2014   17:44:53
155HARASEWYCH & PETIT: BROCCHINIA CLENCHI PETIT, 1986 FROM THE AZORES
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Paginação_Açoreana_Suplemento 10_prova05.indd   155 05-10-2014   17:44:53
156 2014, Suplemento 10: 149-165A Ç O R E A N A
genome (16,648 bp) (Cunha et al., 2009) 
using Clustal X 2.1 (Larkin et al., 2007). 
For the cytochrome c oxidase I gene, an 
alignment of 658 bases, corresponding 
to positions 39 to 696 of the COI gene in 
Bivetiella cancellata, was produced after 
primers were trimmed.  After trimming 
primers, the 16S rDNA sequences 
corresponded to a 514 bp region (positions 
806 to 1319) of this gene in Bivetiella 
cancellata, derived from its complete 
mitochondrial genome. However, due to 
length variation in this portion of the gene 
among the studied taxa, the alignment 
spanned 588 positions.  
Using default seĴ ings and the Akaike 
Information Criterion, jModelTest v. 2.1.3 
(Daribba et al., 2012) selected the HKY+I+G 
nucleotide substitution model for the COI 
data, and the GTR+I+G model for the 16S 
data.  The COI, 16S, and concatenated 
(COI+16S) data sets were analysed using 
Mr. Bayes v. 3.2.1 (Ronquist et al., 2012). 
The concatenated data set was analyzed 
keeping the two partitions unlinked, 
using the corresponding model for each 
partition. Metropolis-coupled Markov 
Chain Monte Carlo simulations were 
performed with four chains for 1,000,000 
generations, with trees sampled every 
500 generations, after a burn-in of 2500 
generations, for each of the data sets. The 
standard deviation of split frequencies 
was below 0.01 and the potential scale 
reduction factor was 1.00 for each of the 
analyses. As only 16S sequences were 
available for Brocchinia tanimbarensis, this 
species appears only in the analysis based 
on 16S sequences.
SYSTEMATICS
Class GASTROPODA
Order Neogastropoda Wenz, 1943
Family Cancellariidae Forbes & Hanley, 
1851
Genus Brocchinia Jousseaume, 1887
Brocchinia clenchi Petit, 1986
Synonymy
Cancellaria pusilla H. Adams, 1869: 274, pl. 19, 
fi g. 12 (non Cancellaria pusilla G.B. Sowerby 
I, 1832: 7, fi g. 34).
Cancellaria mitraeformis Brocchi − Jeff reys, 
1885: 49.
Cancellaria pusilla H. Adams − Sykes, 1911: 
332.
Cancellaria (Brocchinia) pusilla H. Adams − 
Dauĵ enberg 1927: 73, 377.
Narona (Brocchinia) pusilla (H. Adams) − 
Nordsieck, 1968: 151, pl. 25, fi g 87.30.
Narona (Sveltia) pusilla (H. Adams) – 
Nordsieck & Garcia-Talavera, 1979: 152.
Narona pusilla (H. Adams) − Rolan Mosquera 
1983: 236.
Brocchinia clenchi Petit, 1986: 24, fi gs. 1-4; 
Verhecken 2002: 510; Verhecken 2007: 311, 
fi gs. 28-30.
?Brocchinia canariensis Rolán & Hernández, 
2009: 104,fi gs.1-7; Hernández et al. 2011: 
221-222, fi g. 72 J-M. 
Descџiption
Shell morphology: Shell (Figures 4-9, Table 
2) small (to 5.2 mm), thick, with tall, 
conical, spire, and rounded anterior. 
Protoconch (Figures 6, 10) paucispiral, 
increasing in diameter from 251 μm to 
731 μm in 1.4 smooth, evenly rounded, 
conical, glassy whorls (mean values, see 
Table 2). Protoconch initially deviated 
from teleoconch axis by ~ 15°. Transition 
to teleoconch (Figure 10, arrow) distinct, 
marked by an abrupt change from smooth, 
glassy surface to maĴ e texture and onset 
of numerous, very fi ne prosocline growth 
lines. Transition may be less conspicuous 
on dead or eroded specimens. Teleoconch 
of up to 3.5 very thick, evenly convex 
to weakly shouldered whorls. Axial 
sculpture of fi ne prosocline growth striae 
and weak, irregularly spaced nodes 
most evident above shell periphery in 
whorls 1−2.5, becoming less prominent 
as shell increases in thickness. Spiral 
sculpture of 3−5 low cords, one running 
along shell periphery, 1−2 above and 1−2 
below periphery. Suture distinct, weakly 
impressed, abuĴ ing peripheral cord 
Paginação_Açoreana_Suplemento 10_prova05.indd   156 05-10-2014   17:44:53
157HARASEWYCH & PETIT: BROCCHINIA CLENCHI PETIT, 1986 FROM THE AZORES
in early whorls (Figure 8), descending 
below peripheral cord and becoming 
more strongly impressed after second 
whorl (Figures 7, 9). Aperture D-shaped, 
defl ected from coiling axis by 20-22°. 
Outer lip smooth, rounded anteriorly and 
posteriorly. Inner surface with 5-8 short 
(≤ ¼ whorl) lirae that expand to form 
pronounced denticles recessed within the 
lip, beneath thickened axial node (Figure 
12). Parietal callus thin to moderately 
thick, translucent. Columella thick, 
straight, axial, forming an angle of 122-
136° with parietal region (angle decreases 
as whorl number increases), with two 
oblique columellar folds (Figure 11, cf), 
and a weaker siphonal fold (Figure 11, 
sf). Posterior columellar fold longer and 
thicker. Siphonal canal short, broad, not 
well diff erentiated. Shell color uniform 
white. Operculum absent.
Shell ultrastructure: (Figure 13) Shell thick 
(≈220 μm), composed of four distinct 
crystal layers. The innermost layer 
(Figure 13, in) (≈10 μm) comprises the 
glaze that lines the inner surfaces of the 
shell. The adjacent layer (Figure 13, per) 
(≈60 μm) is composed of cross-lamellar 
aragonite, with crystal faces oriented 
roughly perpendicular to the growing 
edge of the shell. The next layer (Figure 
13, in) (≈110 μm) is also comprised of 
cross-lamellar aragonite, with crystal 
faces orthogonally oriented parallel to 
the growing edge. The outermost layer 
(Figure 13, in) (≈40 μm) is prismatic. It 
lacks the piĴ ed surface (intriticalx?) that 
is present in the Pliocene type species 
Brocchinia mitraeformis (Figures 1, 3), 
and in some Recent Indo-Pacifi c species 
[e.g., B. fi scheri and B. kaeiensis but not B. 
tanimbarensis (see Verhecken, 1997: 300-
307)].
External anatomy: Extended animal 
(Figure 16) spans 3½ whorls, with the 
mantle cavity covering nearly 1 whorl, 
the kidney occupying ¼ whorl along its 
right posterior limit, and the digestive 
gland and gonad spanning 2¼ whorls. 
The terminal 1-1¼ whorls appears to 
be primarily gonadal tissue. The foot is 
twice as long as wide, with a pronounced 
propodial groove. The head is broad 
with a pair of long tentacles (contracted 
when preserved) and large black eyes 
along their outer margins. Mantle edge 
thickened, smooth, without a conspicuous 
siphon. When alive, the animal is white, 
with brighter white speckles. Preserved 
animals are a uniform pale ivory. 
Mantle cavity: The mantle cavity is 
broad, with organs situated as in most 
neogastropods. The osphradium is 
greatly reduced, the ctenidium consisting 
of about a dozen lamellae. A thin 
hypobranchial gland extends across the 
roof of the mantle cavity. The pericardium 
is situated at the left posterior margin of 
the mantle cavity and contains a minute 
auricle.
Alimentary system: The proboscis is 
short, broad and pleurembolic (L/W 
~3), occupying most of the cephalic 
haemocoel. Mouth small, forming a 
dorsoventral slit at proboscis tip, leads 
to a small buccal cavity. Buccal mass 
minute (~½ proboscis width), ovate, 
situated posterior to proboscis tip. 
Radula absent. Neither a cuticularized 
oral tube nor a jaw were observed in 
this species. A minute valve of Leiblein 
is situated posterior to the buccal mass. 
Salivary glands were large and fl anked 
the nerve ring, with narrow salivary 
ducts leading to the buccal mass. The 
mid-esophagus passes through the 
nerve ring then expands as it runs 
posteriorly to rear of cephalic hemocoel 
and to the stomach, which is thin, 
narrow and runs along the anterior 
portion of the digestive gland. Intestine 
leads to rectum that runs along the right 
Paginação_Açoreana_Suplemento 10_prova05.indd   157 05-10-2014   17:44:53
158 2014, Suplemento 10: 149-165A Ç O R E A N A
FIGURES 7-15. Brocchinia clenchi Petit, 1986. USNM 1197435, Sabrina Bank, São Miguel, Azores, 
Portugal (37°51.547’N, 25°41.706’W), dredged in 200 m, July 8, 2011 (Sta. SAB2011.DR6). 7, 
Apertural, lateral, dorsal, and apical views of adult specimen. 8, Apertural, lateral, dorsal, and apical 
views of juvenile specimen. 9, Apertural and apical views of juvenile specimen. 10, Lateral view 
of protoconch, showing transition to teleoconch (arrow). 11, Shell fractured to reveal columella. 
12, Inner surface of outer lip, showing depth of denticles and lirae. 13, Shell ultrastructure, surface 
parallel to growing edge of shell, ¼ whorl behind the lip. 14, Naticid borehole, indicative of 
drilling predation. 15, Repaired shell break, indicative of shell peeling predation by a crustacean. 
cf, columellar fold; in, innermost shell layer; ou, outer, prismatic shell layer; par, crossed-lamellar 
aragonite, crystal faces parallel to growing edge; per, crossed-lamellar aragonite, crystal faces 
perpendicular to growing edge; sf, siphonal fold.
Paginação_Açoreana_Suplemento 10_prova05.indd   158 05-10-2014   17:44:53
159HARASEWYCH & PETIT: BROCCHINIA CLENCHI PETIT, 1986 FROM THE AZORES
side of the mantle cavity. Anus situated 
posterior to thickened mantle edge. 
Male reproductive system: The testis 
spans the apical whorl of tissue, lining 
the digestive gland tissue and giving 
rise to a narrow testicular duct that 
leads anteriorly to the right wall of the 
mantle cavity, where it expands to form 
a slightly convoluted prostate gland 
and then constricts to form a narrow 
vas deferens that descends to the fl oor 
of the mantle cavity before entering the 
base of the penis, situated just behind 
the right cephalic tentacle. Penis long, 
dorsoventrally fl aĴ ened, parallel sided, 
tapering distally but lacking a distinct 
papilla. 
Female reproductive system: The ovary is 
large, acinous, occupying most of the 
terminal whorl of the visceral mass. 
A thin-walled oviduct runs anteriorly 
along the columellar side of the digestive 
FIGURES 16-17. Anatomical features of Brocchinia clenchi Petit, 1986. 16, Male specimen removed from 
shell, partially uncoiled and mantle cavity opened mid-dorsally. 17, Diagrammatic representation 
of female pallial gonoduct. a, anus; ag, albumen gland; bc, bursa copulatrix; cg, capsule gland; ct, 
ctenidum; dg, digestive gland; e, eye; fo, female opening; hg, hypobranchial gland; ig, ingesting 
gland; k, kidney; ko, kidney opening; od, oviduct; os, osphradium; pc, pericardium; p, penis; pg, 
prostate gland; pr, proboscis(extended); r, rectum; s, siphon; sto, stomach; t, tentacle; te, testes.
Paginação_Açoreana_Suplemento 10_prova05.indd   159 05-10-2014   17:44:55
160 2014, Suplemento 10: 149-165A Ç O R E A N A
gland, passing alongside the kidney and 
pericardium before giving rise to the 
pallial gonoduct (Figure 17), consisting 
of an albumen gland, ingesting gland, 
large capsule gland, and a small bursa 
copulatrix, with the female opening at its 
tapered, distal end.  
Ecology: Published records of Brocchinia 
clenchi in the Azores range in depth from 
15-1,250 m (Petit, 1986: 26; Verhecken, 
2007: 312), with the deepest record 
(1,250 m) from between São Jorge and 
Pico. Martins et al. (2009: 64, 98-99, text 
fi gure 1) reported this species off  the 
south central coast of São Miguel, living 
at depths of 58-81 m. In this study, the 
species was collected at depths ranging 
from 50-200 m, on sand and rubble 
boĴ oms in the vicinity of the Sabrina 
Bank, off  western São Miguel. Most 
(80%) of the intact shells were dead 
collected and inhabited by sipunculans 
[Aspidosiphon (Aspidosiphon) muelleri 
Diesing, 1851]. Of these, most (63%) 
had been drilled by naticids (Figures 10, 
14). Two specimens had repaired breaks 
to the outer lip (Figure 15), indicating 
that they had survived an aĴ ack by a 
crustacean. 
Molecular phylogeny: Bayesian analyses 
of the COI and 16S gene sequences 
analyzed separately and together each 
support the monophyly of the genera 
Brocchinia, Mirandaphera, Cancellaria 
and the subfamily Cancellariinae 
(based on very limited taxon sampling). 
However, inference of the relationships 
of Brocchinia diff er depending on the 
gene. COI sequences place Brocchinia, but 
not Mirandaphera, within Cancellariinae 
(Figure 20). 16S sequence data place 
Brocchinia as sister taxon to Mirandaphera 
in a well-supported clade within the 
Admetinae (Figure 21), while the 
concatenated data set (COI + 16S) retain 
the sister group relationship between 
Brocchinia and Mirandaphera, but place 
this clade outside of both Cancellariinae 
and Admetinae (Figure 23). 
DISCUSSION
Species presently allocated to the 
genus Brocchinia conform to one of three 
basic shell morphotypes. Fossil species, 
including the type species of the genus 
(Figures 1-3), tend to have comparatively 
large (to 13 mm), narrow shells with tall, 
conical spires and rounded anteriors, 
with the fi nal whorl occupying about ½ 
the shell length. The protoconch (Figure 
2) spans ~3 whorls, the teleoconch 5-6 
whorls. Sculpture consists of thin but 
distinct spiral cords and low axial nodes 
that are most conspicuous along the 
shoulder. The aperture is ovate, with 
two columellar folds and a prominent 
siphonal fold that demarcates a short but 
distinct siphonal canal. The outer lip is 
thin, with prominent lirae. The shell has 
a distinctively piĴ ed surface (Figure 3), 
especially between spiral cords, which is 
thought to correspond to the intriticalx 
(see Verhecken, 1997: 301 for discussion 
of this piĴ ed sculpture). 
Two diff erent morphotypes occur 
among the Recent species. One is 
characterized by a much broader shell, 
usually <12 mm in length, but may reach 
27 mm in one species. These shells have 
a short, wide spire and a last whorl that 
spans ≥¾ of the shell length. In this group, 
protoconchs range from 1¼ - 3¼ whorls, 
while teleoconchs span 3-4½ whorls. 
Sculpture consists of strong spiral cords 
and prominent axial ribs that produce a 
cancellate to nodulose shell surface. The 
aperture is large, broadest adapically, 
generally with a single columellar fold 
and a prominent siphonal fold. The outer 
lip may be undulating, but lacks lirae. 
A piĴ ed shell surface is present in some 
species. Species with this morphotype 
include: Brocchinia azorica, B. decapensis, B. 
Paginação_Açoreana_Suplemento 10_prova05.indd   160 05-10-2014   17:44:56
161HARASEWYCH & PETIT: BROCCHINIA CLENCHI PETIT, 1986 FROM THE AZORES
nodosa and B. pustulosa in the Atlantic and 
B. kaiensis, as well as B. tanimbarensis in the 
Indo-west Pacifi c. 
The remaining Recent species 
generally have a small (≤6 mm, although 
B. fi scheri is reported to reach 12.8 mm), 
narrow shells, with conical spires and 
rounded anteriors. The last whorl spans 
≤½ the shell length. Protoconchs span 
1¼ - 2 whorls, while teleoconchs usually 
comprise 3¼-4 whorls (up to 6 in B. fi scheri). 
Shell sculpture ranges from smooth, 
with barely discernible spiral cords, to 
prominent axial ribs and spiral cords that 
produce a cancellate to weakly nodulose 
surface. The aperture is D-shaped, with 
an axial columella bearing two columellar 
folds and a weak siphonal fold. The outer 
lip is thick, with prominent lirae in most 
species. PiĴ ed shell surface (intriticalx?) 
is present in one species. Species that 
share this morphotype include: Brocchinia 
FIGURE 18. Inference of larval development type based on relationship between maximum protoconch 
diameter and number of protoconch whorls following the model proposed by Shuto (1974) and 
applied to cancellariids by Pawlik et al. (1988). 
◆ = Brocchinia mitraeformis; ■ = “B. pusilla” NHMUK specimen;  ● = B. clenchi, and ▼ = B. canariensis 
(all based on data in Table 2).  □  = Brocchinia azorica (data from Verhecken, 1997: 310)*1; ⦿ = B. nodosa, 
Verhecken (1997: 317) reported this data but questioned the identifi cation of the juvenile specimen 
on which it was based*1; ✠ = B. pustulosa (data from Verhecken, 1991: 548)*1; ❖ = B. verheckeni; △ = B. 
harasewychi (data from de Barros & de Lima, 2007); ▲ = B. fi scheri; ♦ = B. tanimbarensis; ☼ = B. kaiensis 
(data from Verhecken, 1997: 310)*1
*1  The number of protoconch whorls reported in this plot is based on a method of measuring whorls 
(Jablonski & Luĵ , 1980: 332) that results in values ½ whorl greater than those produced by the method 
shown by Verhecken (2007: fi g. 4). Values reported by Verhecken have been increased by ½ whorl.
Paginação_Açoreana_Suplemento 10_prova05.indd   161 05-10-2014   17:44:56
162 2014, Suplemento 10: 149-165A Ç O R E A N A
clenchi in the eastern and mid-Atlantic, 
B. harasewychi and B. verheckeni in the 
western Atlantic and B. exigua, B. fi scheri 
and B. septentrionalis in the Indo-west 
Pacifi c. 
Although these three morphotypes 
are easily distinguished, several of the 
Recent species have features that span 
two of the morphological groupings. The 
Indo-Pacifi c species Brocchinia fi scheri, 
for example, has the shell shape and 
proportions as well as protoconch and 
aperture morphology that unite it with 
the small, narrow shelled group of Recent 
species, yet shares the large shell size, high 
number of whorls and piĴ ed shell surface 
of the fossil group. Both Brocchinia azorica 
and B. nodosa have the characteristic 
teleoconch features of the broad-shelled 
group of Recent species, yet their 
multispiral protoconch is more similar 
to that of the fossil group. This may also 
be true of B. decapensis, but its protoconch 
morphology remains unknown.
In the absence of direct observations, 
larval development is often inferred 
from the morphology of the protoconch. 
The multispiral protoconchs of the fossil 
Brocchinia species have been interpreted 
as being indicative of a planktonic larval 
stage, while the paucispiral protoconchs 
of most of the Recent species were 
inferred to have lecithotrophic, likely 
direct, development (Verhecken, 2007; 
Rolán & Hernández, 2009). The model 
proposed by Shuto (1974), as applied 
by Pawlik et al., (1988: fi g. 23), supports 
the interpretation that most Recent 
Brocchinia with paucispiral protoconchs 
have lecithotrophic larvae (Figure 18). 
However, the type of larval development 
of the fossil and Recent species with 
multispiral protoconchs cannot be 
inferred unambigously as planktotrophic 
using these criteria. The only Recent 
species with multispiral protoconchs 
are those members of the broad-shelled 
group that occur at abyssal depths. 
The species Brocchinia canariaensis 
was distinguished from B. clenchi based 
on comparisons of measurements of the 
protoconchs and adult shells of the type 
material of these two taxa. A somewhat 
broader analysis (cluster analysis, 
UPGMA, Pearson (r) coeffi  cient) that 
included the same type specimens as well 
as eight adult and 2 juvenile Azorean 
specimens and used similar measurements 
(Table 2) did not distinguish B. canariensis 
from B. clenchi (Figure 19). Rather, it 
grouped the holotype of B. clenchi together 
with the paratype of B. canariensis, 
and the holotype of B. canariensis with 
the two paratypes of B. clenchi, of 
which one is the possible syntype of B. 
pusilla from the Canary Islands. This 
analysis is not suffi  ciently rigorous to 
show conclusively that B. canariensis 
is a synonym of B. clenchi. However, 
it does favor Verhecken’s (2007: 313) 
provisional interpretation that only a 
single, highly-variable, bathyal species of 
FIGURE 19. UPGMA cluster analysis [Pearson 
(r) coeffi  cient] based on measurements of 
protoconch and shell (Table 2). HT= holotype, 
PT= paratype . 
Paginação_Açoreana_Suplemento 10_prova05.indd   162 05-10-2014   17:44:56
163HARASEWYCH & PETIT: BROCCHINIA CLENCHI PETIT, 1986 FROM THE AZORES
Brocchinia with a broad geographic and 
bathymetric range inhabits the eastern 
Atlantic. Clearly, additional analyses, 
possibly involving the much larger 
suite of available specimens listed by 
Verhecken (2007: 312), and/or molecular 
comparisons are needed to resolve the 
relationship between these two taxa.
The limited molecular data indicate 
that the species conforming to the 
small, thin-shelled morphotype from 
the Atlantic and Indo-Pacifi c oceans (B. 
clenchi and B. fi scheri) are more closely 
related to each other than either is to the 
broad-shelled group (B. tanimbarensis) 
(Figure 21). As recognized by Modica et 
al. (2011), the relationships of Brocchinia 
within the Cancellariidae are as yet not 
well resolved by molecular data. Analyses 
using COI sequence data place Brocchinia 
within Cancellariinae, not as sister taxon 
of Mirandaphera (Figure 20). However, 
sequence data from the 16S rDNA 
gene analyzed separately (Figure 21) or 
FIGURES 20-23. Relationships of Brocchinia clenchi within Cancellarioidea based on Bayesian 
analyses of:  20, A 658 bp portion of the cytochrome c oxidase I (COI) gene, 21, a portion of the 
16S rDNA gene (alignment length = 588 positions), 22, concatenated COI and 16S DNA sequences. 
Posterior probabilities > 0.85 are indicated at nodes. 
Paginação_Açoreana_Suplemento 10_prova05.indd   163 05-10-2014   17:44:57
164 2014, Suplemento 10: 149-165A Ç O R E A N A
concatenated with the COI sequences 
(Figure 22), supports the sister relationship 
between Brocchinia and Mirandaphera 
and groups these genera with Admete 
(16S only) or in a group separate from 
Admetinae and Cancellariinae (COI + 
16S), although without strong support.
While the thick shell, prominent 
columellar folds and apertural lirae of 
many Brocchinia species are features shared 
with members of the Cancellariinae, the 
absence of a radula is a characteristic of 
some, though not all Admetinae. Thus 
far, neither morphological characters 
nor molecular data have been able 
to determine unambiguously the 
relationship of the genus Brocchinia within 
the family Cancellariidae. As suggested 
by Modica et al. (2011: 691), Brocchinia 
and Mirandaphera may both be part of 
an independent lineage that is certainly 
“worthy of more detailed investigation.”
ACKNOWLEDGEMENTS
We are grateful to Prof. António Frias 
Martins, Dr. Joana R. Xavier and their staff  
and students for organizing the Fourth 
International Workshop of Malacology to 
study the fauna of Sabrina Bank, in São 
Miguel, Azores, which the senior author 
aĴ ended.
Support from FLAD (Portuguese-
American Foundation for Development) 
is gratefully acknowledged.
Thanks also to Elizabeth Diamond 
for assistance with histology, to Yolanda 
Villacampa for taking the Scanning 
Electron Micrographs, and to Eléonore 
Dixon-Roche for preparing the anatomical 
illustrations. 
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