SPIXIANA 37 1 45-59 München, August 2014 ISSN 0341-8391

Sea slugs of Peru:

Peruvian-Chilean faunal elements

(Mollusca, Heterobranchia)

Michael Schrödl & Yuri Hooker

Schrödl, M. & Hooker, Y. 2014. Sea slugs of Peru: Peruvian-Chilean faunal ele-
ments (Mollusca, Heterobranchia). Spixiana 37 (1): 45-59.
The Peruvian marine invertebrate fauna is thus far poorly investigated referring
to sea slugs. From recent surveys along the entire Peruvian coast we present new
distributional data on those 15 benthic opisthobranch gastropod species that were
formerly known from Chilean waters. Our findings include 12 nudibranch, 1 ce-
phalaspidean and 2 sacoglossan species. These are the first records for Peru of 7
species, such as Janolus rebeccae and Hancockia schoeferti. Known distributional
ranges are extended to the north for 9 species, in case of Polycera priva for more than
3000 kilometres; the latter was formerly considered as a Magellanic species en-
demic to southern Chilean fjords. Photographs of living specimens as well as de-
scriptions of habitats and biological observations are given. We also present the
first record of splanchnotrophid copepods from Peru, infesting the aeolid nudi-
branch Phidiana lottini. Some further data and discussions are provided for species
with insufficient or disputed information available.
Michael Schrödl, SNSB – Zoologische Staatssammlung München, Münchhausen-
str. 21, 81247 München, Germany;  e-mail:  [email protected]
Yuri Hooker, Laboratorio de Biología Marina, Facultad de Ciencias y Filosofía,
Universidad Peruana Cayetano Heredia, Lima, Peru

Introduction the Peruvian faunal province (Tarazona et al. 2003,

Hooker 2009). The South American coast south of
Peruvian coastal waters can be roughly divided into roughly 41° S on both the Pacific and Atlantic sides
three sections:  1) the northernmost part north of 4° S, comprises the Magellanic (or Magellan) province, of
which is steadily influenced by tropical waters,  2) a which the fauna of the Chilean fjord area recently has
narrow transitional section (approx. 4-6° S) between received considerable survey activity (see Häusser-
tropical and temperate waters, and 3) a large zone mann & Försterra 2009). There are several reviews
south of 6° S, which is dominated by the temperate on Magellanic opisthobranch gastropods (Marcus
Peru current (Humboldt current). The latter area 1959, Schrödl 1996a, 2003, 2009). Bergh (1898) de-
shows frequent cold water upwellings in normal scribed a couple of new opisthobranch species from
years, but also periodical El Niño events with warm central and northern Chile and limited knowledge
northern waters, poor in nutrients and oxygen, on opisthobranchs from the Chilean part of the
expanding towards the south (Arntz & Fahrbach Atacama Desert coast was summarized by Schrödl
1996). According to these current regimes, the marine (2003). The coastal sea slug fauna of nearby Peru
coastal fauna of northernmost Peru is largely Pana- was even less explored, and still may be among the
manian, while the region from approx. 6° S (Peru) to poorest known on a global scale.
41° S (southern Chile) hosts a fauna that is adapted Older faunal studies include D’Orbigny (1835-
to temperate conditions, and was considered as 47), who described several opisthobranch species

45
from Callao, central Peru, and Dall (1909), who Molecular results of Ornelas-Gatdula et al. (2012)
described Peruvian shells. Faunal lists such as those showed that Peruvian and other Eastern Pacific
by Álamo & Valdivieso (1987) and Paredes et al. Navanax aenigmaticus specifically differ from their
(1999) summarized historical records of Peruvian Atlantic sibling species. Pruning unsubstantiated
opisthobranchs, however, including several unsub- literature records and adding 3 new own records
stantiated records, as noted by Uribe et al. (2013). of nudibranchs in their review, Uribe et al. (2013)
Based on collected specimens, Millen et al. (1994) considered 56 opisthobranch species (2 Acteonoidea
described Okenia luna Millen, Schrödl, Vargas & species, 23 Nudibranchia, 27 Euopisthobranchia, 3
Indacochea, 1994 as a new nudibranch species from Sacoglossa and 1 Acochlidia) as valid for Peru. Uribe
Chile and Peru. Revising the Chilean and Magellanic et al. (2013) emphasized the need of revision of Pe-
nudibranch fauna, Schrödl (2003) reported Flabellina ruvian Aplysia species and thecosomes, the need of
sp. 2 (found by S. Millen) also from Peru; this spe- molecular surveys to recognize potentially cryptic
cies likely refers to Flabellina cerverai Fischer, Van species, and the strong need for further collectings
de Velde & Roubos, 2007, which was established on along the Peruvian coast. In particular, many spe-
Chilean specimens by Fischer et al. (2007). Nakamura cies already known from central or northern Chile
(2006) first reported several Panamanian species from (Schrödl 2003) might be suspected to extend more
northernmost Peru, Martynov et al. (2011) described or less far north into the temperate regions of Peru,
the new species Corambe mancorensis Martynov, thus augmenting the species diversity.
Brenzinger, Hooker & Schrödl, 2011, and Jörger et During our own recent expeditions we collected
al. (2012) reported the morphologically cryptic aco- benthic opisthobranchs along the entire Peruvian
chlidian Pontohedyle sp. The latter was described as coast. On longer terms, together with a multinational
a new species P.  yuri­hookeri Jörger & Schrödl, 2013 team, we attempt to revise the Peruvian opistho-
based mainly on evidence from gene sequences branch fauna and its evolution as a whole. It should
recently (Jörger & Schrödl 2013). In southern Peru, be mentioned that here we use “opisthobranchs” as a
Uribe & Pacheco (2012) discovered specimens ex- descriptive term for euthyneuran marine slugs and
ternally resembling the Atlantic nudibranch Spurilla snails, while decisively abandoning the traditional
neapolitana, a name which rather referred to a species taxon “Opisthobranchia”, which was shown to be
complex (Carmona et al. 2013). Recently, Carmona polyphyletic and reclassified (e. g. Jörger et al. 2010,
et al. (2014) assigned Peruvian Spurilla specimens Schrödl et al. 2011, Brenzinger et al. 2013); for a recent
to Spurilla braziliana MacFarland, 1909, showing review see Wägele et al. (2014).
a wide Western Atlantic and Pacific distribution Herein we focus on reporting those findings of
and occurring in tropical and temperate waters. opisthobranch species in Peru that were already

Table 1.  Collecting data; range extensions and species new for Peru in bold.

Localities San Juan Laguna Paracas Callao Sechura Mancora,

de Grande Bay Punta Sal,
Marcona Tumbes
Nudibranchia: Doridoidea Doris fontainei x
Diaulula variolata x x
Baptodoris peruviana x
Rostanga cf. pulchra x
Corambe lucea x x x x
Okenia luna x
Polycera priva x
“Arminoidea” Janolus rebeccae x x
Dendronotoidea Doto uva x x
Hancockia schoeferti x
Aeolidoidea Phidiana lottini x x x
Flabellina cf. cerverai x x
Cephalaspidea Navanax aenigmaticus x x x
Sacoglossa cf. Ercolania evelinae x
Elysia cf. hedgpethi x x

46
known from Chile. Besides giving new morphologi- temperature 23 °C, 27.11.2006. B) Islas Ballestas
cal, biological and distributional data we also modify (13°43.8' S, 76°23.9' W), steep rock walls, at 8-15 m
and discuss zoogeographic patterns observed for depth, currents and swell, water temperature 15 °C,
temperate southeastern Pacific nudibranchs, which 28.11.2006.  C) Peninsula, sediment bottom off sandy
were biased to information from Chile in the past. beach, at 3-5 m depth, 28.11.2006.  D) Paracas, scallop
farm on sediment bottoms (13°48.7' S, 76°17.6' W), at
0-5 m depth, 28.11.2006. E) Paracas, fishery pier
(15°21'20" S, 75°09'39" W), shallow water macroalgae
Material and methods and epizoans, 25.11.2006.
3) La Punta, Callao, central Peru. Sandy bottom with
In November and December 2006 we performed a pub- stones, and pier of IMARPE (12°04'04" S, 77°09'38" W)
lic transport (bus, mototaxi, taxi) based survey of with macroalgae and epizoans, at 0-3 m depth,
opisthobranch molluscs at 7 major stations along the water temperature 18 °C, 22.11.2006.
Peruvian coast at San Juan de Marcona (several sites), 4) Bahía Sechura, Piura, northern Peru. A) Several
Ica; Reserva Nacional de Paracas (Laguna Grande, Islas collecting stations off Bayovar (5°48' S, 81°02' W),
Ballestas, and 3 sites near Paracas), Ica; La Punta, Callao; rocky and sandy bottoms at 4-27 m depth, 8.12.2011.
Bahía Sechura (several sites), Piura; Máncora (several B) Sediment bottom with scallop farms (5°46' S,
sites), Piura; Punta Sal (several sites), Tumbes; Puerto 80°58' W), at 6-10 m depth, 09.12.2006.
Pizarro, Tumbes (see station list below; Fig. 1). All sta- 5) Máncora, Piura, northern Peru. A) Fishery pier
tions were intertidal and shallow subtidal (collecting via (4°6'36" S, 81°4'2" W) with macroalgae and epizoans,
tide pooling and snorkelling), and all but Callao and at 0-5 m depth, 02.12.2006 and 06.12.2006.  B) Sandy
Puerto Pizarro also were subtidal down to 15-27 m beach with intertidal and shallow subtidal rocks and
depth, with SCUBA dives at several sites. We investi- stones (4°06'20" S, 81°03'31" W), 02.12.2006 and
gated all available benthic habitats, such as soft and 04.12.2006.
rocky bottoms, deeper rock walls, caverns, and macroal- 6) Punta Sal, Tumbes, northern Peru.  A) Several col-
gae; gravel, algae, hydrozoa and other substrate samples lecting stations off coast (3°59' S, 80°59' W), rocky and
were collected and searched for small opisthobranchs sandy bottoms, at 8-15 m depth, 03.12.2006.  B) Pier
in aquaria and under the binocular microscope. Speci- of Cancas (3°56' S, 80°56' W), 05.12.2006.
mens, egg masses and potential prey were collected, 7) Puerto Pizarro, Tumbes, northern Peru. Isla de Amor
members of each species documented in living condi- (3°29'47" S, 80°23'28" W), intertidal sands and man-
tion, and specimens fixed (4 % formalin-seawater; 3.7 % groves, 01.12.2006.
buffered glutaraldehyde, 70 % or 96 % ethanol) for
further taxonomic analysis. Specimens were identified
externally to the level possible using the keys and de-
Results
scriptions given by Schrödl (2003, 2009) and more spe-
cialized literature referred to separately.
Reference specimens were deposited at the Zoolo- Euthyneura, Nudipleura, Nudibranchia,
gische Staatsammlung München (ZSM) and the Colec- Doridoidea, “Cryptobranchia”, Dorididae,
ción de Zoologia Acuática (CZA) of the Universidad Doris Linnaeus, 1758
Peruana Cayetano Heredia. Additional specimens from
other collecting trips along the Peruvian coast (Paracas, Doris fontainei (D’Orbigny, 1837)
Pucusana, Bahía Ferrol, Máncora and Punta Sal) and Fig. 2
photographs of living specimens were provided by one
of the authors (YH). Material.  Many specimens with egg masses, 6 speci-
mens collected (ZSM Moll 20090686), rocky reefs off
Punta de San Juan de Marcona, 24 November 2006, at
Collecting sites 8-15 m depth. Two specimens (ZSM Moll 20100765)
from Pucusana, south of Lima, 30 August 2004, coll. YH.
1) San Juan de Marcona, Ica, southern Peru.  A) Rocky An additional record comes from an islet in the Bahía
reefs (15°22'11" S, 75°12'16" W) off Punta de San Juan Ferrol (9°08'59.58" S, 78°36'48.75" W, Ancash, at 17 m
de Marcona, rocky bottom, at 6-15 m depth, water depth, 12 °C water temperature, found on 25 Septem-
temperature 15 °C, strong currents from swells, ber 2007 during the ESPER 2007 Expedition (Projecto
24.11.2006.  B). Punta San Juan de Marcona (15°21' Esponjas del Peru, YH).
23" S, 75°10'44" W), sandy beach with rocks, at 0-2 m
depth, water temperature 15 °C, 22.11.2006.  C) Fish-
Distribution.  Bahía Ferrol, Ancash, Peru (9ºS; Uribe
ery harbour, pier (15°21'29" S, 75°09'42" W) with et al. 2013) and south along the Peruvian and entire
macroalgae and epizoans, 25.11.2006. Chilean continental coast, as well as Argentinian
2) Reserva Nacional de Paracas, Ica, southern Peru. Patagonia to northern Argentina (see Schrödl 2003,
A) Sediment and hard bottoms around Laguna 2009, Schrödl et al. 2005).
Grande (14°09'S, 76°15'W), at 0-8 m depth, water

47
Observations.  As usual for central and northern in synonymy with the senior Diaulula; thus Schrödl
Chilean specimens (Schrödl 2009), living speci- (2003) briefly discussed this species under Diaulula
mens from Peru show a network of dark pigment variolata. While members of Diaulula are supposed
between the notal tubercles. Egg masses are broad to have special caryophyllidid tubercles, Valdés
translucent ribbons with up to 6 spirals, containing & Muniain (2002) could not find caryophyllidiid
small yellow eggs. tubercles as earlier described from D. variolata by
Marcus (1959) in their material available, and thus
Remarks.  This species was revised in detail under
transferred the species to the caryophyllidiid-lacking
the name Anisodoris fontaini by Schrödl (1997a,
genus Peltodoris. However, having caryophyllidiid
2000a), then called Doris fontainei by Valdés & Mu-
(small) tubercles, a generic placement of D. variolata
niain (2002), and placed tentatively into the genus
within Diaulula may be justified until comprehen-
Archidoris by Schrödl (2003). We feel that comprehen-
sive genetic data is available on dorid nudibranchs
sive molecular phylogenetic analyses are required to
confirming or rejecting current morphology-based
rearrange dorid systematics. Regardless of uncertain
classification.
generic affiliations and different opinions on the
valid species name, this southern American species
is unmistakable because of its large body size up to Baptodoris Bergh, 1884
10 cm, triangular, grooved oral tentacles and large
notal tubercles. For a temperate benthic invertebrate Baptodoris? peruviana (D’Orbigny, 1837)
species it has an extreme latitudinal range of roughly Fig. 4
45 degrees in the Pacific, and still 20 latitudinal
degrees in the southwestern Atlantic. Material.  Three specimens (ZSM Moll 20100802), off
Punta de San Juan de Marcona, 24 November 2006, at
6-12 m depth. Additional specimens were collected by
Discodorididae, Diaulula Bergh, 1878 Vreni Häussermann and Günther Försterra in Chile, dur-
ing December 1997: one specimen (ZSM Moll 20110443),
Diaulula variolata (D’Orbigny, 1837) Playa Corazones (18°31'39" S, 70°19'18" W), Arica, at
Fig. 3 4-5 m depth; one specimen (ZSM Moll 20110460), west
of Punta Lobos (20°59'56" S, 70°09'49" W), Rio Seco,
Material.  Several specimens, 8 collected (ZSM Moll at 15 m depth; two specimens (ZSM Moll 20110465),
20090679, 20100787), at and off Punta de San Juan de Playa Piquero (26°08' S, 70°40' W), Pan de Azucar, at
Marcona, 23-24 November 2006, at 2-15 m depth. Two 8 m depth.
specimens (ZSM Moll 20100760) from Paracas, 3 October
Distribution.  San Lorenzo (off Callao), Peru to
2004, coll. YH; two specimens (ZSM Moll 20100819), 26
Los Molles, Chile (Schrödl 2003, Fischer & Cervera
November 2006.
2005a). Early records from Galápagos and Valparaíso
Distribution.  Paracas, Peru (~14° S; Uribe et al. need confirmation (see Schrödl 2003).
2013) and south along the Peruvian and Chilean
Remarks.  The synonymy of the just externally
continental coast to Concepción (see Schrödl 1997b,
known Doris peruviana D’Orbigny, 1837 from Cal-
2003). More recently, D. variolata was found off
lao with the Chilean Platydoris punctatella Bergh,
Playa Caballito (41°14'48" S, 73°51'45" W; ZSM Mol-
1898 was proposed by Schrödl (2003) based on
20110395) at 2 m depth, (new range extension herein)
characteristic external shape and whitish or yel-
and even reported from the southern part (51° S) of
lowish coloration with the notum scattered with
the Chilean fjord region by Aldea et al. (2011); the
dark dots, except for the margin. This synonymy
latter record, however, needs confirmation (Uribe
was confirmed by Fischer & Cervera (2005a), who
et al. 2013).
designated a neotype for Doris peruviana using a
Observations.  Light microscopic and SEM analyses specimen from Iquique, northern Chile. The latter
herein show that typical caryophyllidiid tubercles, authors mentioned a museum label of one specimen
with a crown of spicules surrounding a ciliary saying that it was uniformly orange coloured in
sensory knob, are present in D. variolata. While live. We cannot confirm this assumed intraspecific
smaller tubercles are slender caryophyllidiids, larger colour variability, since all newly collected Chilean
tubercles apically show a more complex pattern of and Peruvian specimens resemble the originally
spicules and sensory ciliary fields. described colour pattern.
Dorgan et al. (2002) stated that Platydoris punc­
Remarks.  Doris variolata D’Orbigny, 1837 was
tatella (“from Peru”) due to external shape does not
transferred to Anisodoris by Bergh (1898). Accord-
belong to the genus Platydoris, and thus they did not
ing to Valdés & Gosliner (2001), the latter genus is
include it into their phylogenetic analysis. Schrödl

48
Fig. 1.  Map of collecting areas. Large inserts show authors and equipment transported via mototaxi in Mancora,
fisherman operating hookah diving compressor, and second author collecting in cold waters off San Juan de Mar-
cona using compressor and garden hose. Small inserts show authors documenting live specimens after collecting,
and Humboldt penguins resting at the Atacama desert coast, southern Peru.

49
(2003) interpreted the cool water species P. punctatella “Phanerobranchia”, Onchidorididae s. l.,
as a potential stem offshoot of Platydoris, still lacking Corambe Bergh, 1869
the characteristic, very flattened shape and wide free
mantle rim of mainly tropical congeners. Fischer Corambe lucea Marcus, 1959
& Cervera (2005a) found denticles on all radular Fig. 6
teeth of Platydoris peruviana and considered this as
an apomorphy of the genus Baptodoris. However, Material.  Many specimens with egg masses, 13 speci-
members of Baptodoris (as listed in their table 1) are mens collected (ZSM Moll 20080512-16, 20100458,
heterogeneous regarding other anatomical features, 20100462), pier of San Juan de Marcona, 25 November
e. g. regarding the absence versus presence of large 2006, at 0-3 m depth, on Macrocystis. Many speci-
mens with egg masses, 1 specimen collected (ZSM
vaginal and/or atrial hooks, and thus the monophyly
Moll 20080523), Ballestas Islands, 28 November 2006,
of such assembly may be doubted. As mentioned by at 8-15 m depth, on red algae with Membranipora.
Fischer & Cervera (2005a), shapes of radula teeth Many specimens, 15 specimens collected (ZSM Moll
also differ within Baptodoris, and radular denticles 20090491-92, 20090663), pier of IMARPE, Punta de
are present on outer radular teeth also in some Callao, 22 November 2006, at 0-3 m depth. Many
Platydoris species. Overall, members of the probably specimens, 6 specimens collected (ZSM Moll 20090743,
closely related genera Baptodoris and Platydoris, both 20090747, 20090767), sandy bottom (05°47' S, 81°03' W)
sister to Gargamella (see Garovoy et al. 1999), show off Bayóvar, Bay of Sechura, northern Peru, 8 December
a mixture of radular and inconsistently defined and 2006, at 27 m depth, on Caulerpa and red algae.
coded reproductive features (see Garovoy et al. Distribution.  Sechura Bay, northern Peru, along
1999, Dorgan et al. 2002, Fischer & Cervera 2005a) the Peruvian and Chilean coast south to Golfo Cor-
that indicates considerable convergence and unclear covado (almost 44° S), southern Chile (see Schrödl
basal relationships. Although listing this species 2009). These are the first records of C. lucea from Peru,
in the genus Baptodoris (see Uribe et al. 2013) for extending its known range by approx. 18 latitudinal
practical reasons, we still emphasize that molecular degrees to the north.
markers are needed to clarify the phylogeny of such
problematic caryophyllidiid-bearing dorids. Observations.  Among specimens showing the nor-
mal mottled, cryptic coloration (Fig. 6), in Sechura
Bay we found a single albinistic specimen lacking
Rostanga Bergh, 1879 any body pigmentation. Albinism in otherwise pig-
mented opisthobranch species is rarely encountered
Rostanga cf. pulchra MacFarland, 1905 (see Marcus 1975, Roginskaya 1990).
Fig. 5
Remarks.  Corambe lucea was known from intertidal
Material.  One specimen (ZSM Moll 20100459), off and shallow subtidal waters, and was usually associ-
Punta de San Juan de Marcona, 24 November 2006, at ated with Macrocystis pyrifera covered with colonies
6-12 m depth. of its prey, the bryozoan Membranipora isabelleana
(see Schrödl & Wägele 2001, Schrödl et al. 2005).
Distribution.  Rostanga pulchra, if a single species, The present record from relatively deep sandy bot-
would occur in temperate waters of South America toms off Sechura Bay with nests of green and red
and north-eastern Pacific (Schrödl & Grau 2006, algae, the latter covered with Membranipora colonies,
Alvim & Pimenta 2013). Earlier reports of R. pul­ suggests that C. lucea has a wider ecological and
chra from Peru were based on range extrapolations bathymetrical range.
(see Uribe et al. 2013). So this is the first record of The anatomy and phylogeny of Peruvian and
Rostanga from Peru that is substantiated by material worldwide Corambidae were addressed in recent
collected. works (Martynov & Schrödl 2011, Martynov et al.
Remarks.  Preliminary molecular data indicates 2011) showing that corambids are an anatomically
that northern and southern hemispherical Rostanga derived part of the family Onchidorididae, with
pulchra may belong to different species (unpublished evolutionary adaptations to new habitats and shorter
data); the identity of the Peruvian specimens remains life cycles driven by heterochronic shifts in develop-
to be investigated. ment.

50
 Okenia Menke, 1830 “Arminoidea”, Zephyrinidae, Janolus Bergh, 1884

Okenia luna Millen, Schrödl, Vargas Janolus rebeccae Schrödl, 1996

& Indacochea, 1994 Fig. 9
Fig. 7
Material.  One specimen (ZSM Moll 20080529) col-
Material.  Many specimens, 6 specimens collected (ZSM lected at Paracas, Industrial harbour, southern Peru, 28
Moll 20090669, 20090675), pier of IMARPE, Punta de November 2006, at 2-5 m depth, on epibionts on nets
Callao, 22 November 2006, at 0-3 m depth, on algae of scallop aquacultures; three specimens (ZSM Moll
and epibionts. 20100761) collected at Paracas, 3 October 2004, by YH.
Six specimens (ZSM Moll 20090726, 20090727), collected
Distribution.  Ancón, Peru to Bahía de Coliumo, from nets of scallop aquacultures off Bayóvar, Bay of
central Chile (Schrödl 2003). Sechura, northern Peru, 09 December 2006, at 6-10 m
depth.
Observations.  No endoparasitic splanchnotrophid
copepods were found in the herein investigated spec- Distribution.  Bay of Sechura, northern Peru (first
imens, but Ismaila androphila Haumayr & Schrödl, record for Peru), to Bahía Inglesa, northern Chile
2003 is often associated to Chilean populations of (Schrödl 2003).
O. luna (see Haumayr & Schrödl 2003, Schrödl 2003).
Remarks.  Specimens found in Peru externally agree
with descriptions of northern Chilean J. rebeccae by
Polyceridae, Polycera Cuvier, 1817 Schrödl (1996b, 2003). Digestive gland branches
are brownish, ramified within the cerata and vis-
Polycera priva Marcus, 1959 ible through the integument, contrasting J. rebeccae
Fig. 8 from the formal description of J. chilensis Fischer,
Cervera & Ortea, 1997 by Fischer et al. (1997). Most
Material. Two specimens were photographed at Islas of the Peruvian specimens were associated with
Ballestas (13°43'46" S, 76°23'18.33" W) at a depth of ap- nets of scallop aquacultures, obviously feeding
prox. 15 m, by YH on 13 August 2002. on epibiontic bryozoans. We cannot rule out the
possibility of artificial introduction of J. rebeccae via
Distribution.  Paracas, Peru (herein), and Chilean aquaculture activities. More likely, however, we
Patagonia (see Schrödl et al. 2005, Schrödl 2009). just sampled additional populations from a natural,
This is the first record of P. priva from Peru and the continuous geographic range from northern Chile
Peruvian faunal province. The geographical range of to northern Peru.
P. priva was previously thought to be very narrow,
i. e. limited to the northern part of the Chilean fjord
region (41-44° S), but is herein extended by almost Dendronotoidea, Dotidae, Doto Oken, 1815
30 latitudinal degrees to the north.
Observations.  Apart from a single occasion, to our Doto uva Marcus, 1955
knowledge Polycera priva has never again been found Fig. 10
in Paracas or in Peru since.
Material.  Three specimens collected (ZSM Moll
Remarks.  Polycera priva was redescribed in anatomi- 20100455, 20100461), off Punta and pier of San Juan
cal detail by Schrödl et al. (2005). Because of sharing de Marcona, Ica, southern Peru, 23 and 25 Novem-
the characteristic body shape and coloration, there is ber 2006, at 0-4 m depth. Many specimens, 2 speci-
little doubt that our Peruvian photographic record mens and oviposture collected (ZSM Moll 20080524,
refers to P. priva. Future studies will explore whether 20080525, 20020100839), Ballestas Islands (13°43'53" S,
76°23'59" W), off Paracas, Ica, southern Peru, 28 Novem-
this species is a regularly overlooked Peruvian spe-
ber 2006, at 8-15 m depth, on algae with hydrozoans.
cies extending south to the southern Chilean fjords,
Many specimens, 51 specimens collected (ZSM Moll
or basically Magellanic, as numerous recent findings 20000665, 20090674), pier of IMARPE, Punta de Callao,
in fjords and channels had indicated (Schrödl 2009), 22 November 2006, at 0-3 m depth, on algae.
with just sporadic occurrences in far northern but
still Humboldt current influenced areas. The latter Distribution.  Callao, Peru (herein), to southern
scenario could be explained by northwards transport Chilean fjord region (48°S); southern Brazil (see
of larvae with currents, plus triggered metamorpho- Fischer et al. 2006).
sis in the presence of adequate adult prey. Remarks.  Fischer et al. (2006) redescribed Chilean
and Brazilian specimens of Doto uva in morphologi-

51
Figs 2-9.  Peruvian sea slug species, living specimens.  2. Doris fontainei (ca 5 cm), San Juan de Marcona.  3. Diaulula
variolata (ca 5 cm), San Juan de Marcona.  4. Baptodoris peruviana (ca 3 cm), San Juan de Marcona.  5. Rostanga cf.
pulchra (ca 1 cm), San Juan de Marcona.  6. Corambe lucea (ca 1 cm), Callao.  7. Okenia luna (ca 1 cm), with egg masses,
Callao.  8. Polycera priva (ca 2.5 cm), Paracas.  9. Janolus rebeccae (ca 2 cm), Paracas.

52
Figs 10-15.  Peruvian sea slug species, living specimens.  10. Doto cf. uva (ca 5 mm), Callao.  11. Hancockia schoeferti
(ca 1.5 cm), San Juan de Marcona.  12. Phidiana lottini (ca 2 cm), infested with splanchnotrophid endoparasites, 2 pairs
of egg-sacs protruding from body; San Juan de Marcona.  13. Phidiana lottini (ca 3 cm), non-infested; Paracas.  14. Fla­
bellina cf. cerverai (ca 7 mm), Sechura Bay.  15. Elysia cf. hedgpethi (ca 5 mm), Paracas.

cal and histological detail; these authors suspected have more elongated ones. Comparative molecular
that a complex of cryptic species may explain the studies are required to resolve the status of this ap-
considerable external and colour variation observed. parently widespread species, under current concepts
Our specimens from San Juan de Marcona resemble showing a disjunct amphi-South American range
Chilean specimens by having quite stout cerata, (Schrödl 2003).
while specimens from Paracas and Callao may

53
 Hancockiidae, Hancockia Gosse, 1877 Remarks.  The taxonomy of Phidiana lottini and its
junior synonym Phidiana inca was revised by Schrödl
Hancockia schoeferti Schrödl, 1999 (1996a, 2003). In the past, there were several records
Fig. 11 of P. lottini from Callao, and Uribe et al. (2013) re-
ported it from Isla Santa, Ancash, and, with reserva-
Material.  Eight specimens collected (ZSM Moll tions, from tropical waters of Tumbes, which should
20100460, 20100463), pier of San Juan de Marcona, Ica, be confirmed by future surveys.
southern Peru, 25 November 2006, at 0-3 m depth, on
algae with hydrozoans.

Distribution.  San Juan de Marcona, Peru (first Flabellinidae, Flabellina Voigt, 1834
record from Peru), and central to southern Chile
(37-43° S) (Schrödl 2009). Flabellina cf. cerverai Fischer, Van der Velde
& Roubos, 2007
Remarks.  Hancockia schoeferti is an opportunistic Fig. 14
species, in central and southern Chile occurring in
locally high abundances on seasonally hydrozoan Material.  Many specimens with egg masses, 42 speci-
covered algae such as kelp (Schrödl 1999a). Our mens collected (ZSM Moll 20090668), pier of IMARPE,
finding in hydrographically similar, rather cold Punta de Calloa, 22 November 2006, at 0-3 m depth.
Peruvian waters suggests it will ultimately be found One specimen, Sechura Bay, at 6-10 m depth, 9 De-
in localities in between as well. cember 2006.

Distribution.  Bay of Coliumo, central Chile, to Se-

chura Bay, northern Peru (range extended herein).
Aeolidoidea, Glaucidae, Phidiana Gray, 1850
Remarks. As Flabellina sp. 2, this species was briefly
Phidiana lottini (Lesson, 1831) described by Schrödl (1996a, 2003) as a small
Figs 12, 13 slender aeolid with white-tipped, light reddish or
brownish cerata in groups, wrinkled rhinophores
Material.  Many specimens with egg masses, 1 speci- and propodial tentacles, occurring in central Chile
men collected (ZSM Moll 20100457), rocky reefs off and in Ancón, Pucusana, Peru (found by S. Millen).
Punta de San Juan de Marcona, 24 November 2006, The herein reported material from Peru is highly
at 8-15 m depth. Several specimens, pier of San Juan similar and thus likely to be conspecific. Flabellina
de Marcona, Ica, southern Peru, 23 and 25 November
sp. 2 externally resembles Flabellina cerverai Fischer,
2006. Many specimens, 2 specimens collected (ZSM Moll
Van der Velde & Roubos, 2007 that was described
20100762), off Paracas (13°48'50" S, 76°17'36" W), 28 No-
vember 2006, at 2-4 m depth, on algae, nets and buoys. based on material from Coquimbo, Chile (Fischer
One specimen (ZSM Moll 20100728) collected at Pucu- et al. 2007). Comparative anatomical and molecular
sana, south of Lima, Peru, 19 July 2005, by YH. Several studies may confirm conspecifity.
specimens, 2 specimens collected (ZSM Moll 20090670),
pier of IMARPE, Punta de Callao, 22 November 2006, at
0-3 m depth, on algae. Euthyneura, Tectipleura, Euopisthobranchia,
Cephalaspidea, Aglajidae,
Distribution. Callao, central Peru, to Melinka,
Navanax Pilsbry, 1895
Guaitecas Islands (44° S), northern part of the Chilean
fjord region. Navanax aenigmaticus (Bergh, 1893)
Observations.  One specimen collected off San Juan
de Marcona is infested by at least two female Ismaila Material.  Five specimens (ZSM Moll 20100035,
(Copepoda: Poecilostomatoida: Splanchnotrophi- 20100777), Laguna Grande, on sandy bottom, at 2-8 m
depth, 27.11.2006. One specimen (ZSM Moll 20100773),
dae), possibly I. robusta Haumayr & Schrödl, 2003.
Sechura Bay, sediment bottom with scallop farms, at
These highly specialized endoparasites (see Anton 6-10 m depth, 09.12.2006. One specimen (ZSM Moll
& Schrödl 2013a,b) hide their externally protrud- 20100730), Punta Sal, collected by YH, 11.05.2005. Four
ing paired egg sacs between the cerata of the host specimens (ZSM Moll 20100753, -54), Máncora, collected
(Fig. 12). While central Chile hosts an especially rich by YH, 10.04.2005.
splanchnotrophid fauna (Schrödl 2002, Haumayr
& Schrödl 2003), apparently representing a rather Distribution.  Eastern Pacific (Ornelas-Gatdula et al.
recent radiation switching towards different hosts 2012), i. e. Southern California to Panama, Galapagos.
(Anton & Schrödl 2013b), this is the first record of a Northern Peru (Uribe et al. 2013) to central Chile (see
splanchnotrophid copepod for Peru. Schrödl 2007a,b).

54
Remarks.  Recent molecular analyses have shown pers. comm.). Molecular studies are needed to test
that eastern Pacific N. aenigmaticus differ from ex- the assumed conspecifity of northeastern Pacific
ternally similar Atlantic species (Ornelas-Gatdula E. hedgpethi with southeastern Pacific ones, and with
et al. 2012). Navanax aenigmaticus is common in externally similar but larger growing southwestern
Panamanian and warm northern Peruvian waters, Atlantic E. patagonica.
and was very abundant in southern Peruvian Laguna
Grande; this is an almost closed shallow lagoon
showing much higher water temperatures than Discussion
nearby open waters (23 °C versus 15 °C during our
visit). This suggests Laguna Grande populations are There can hardly be any greater contrast in climate
either permanent or temporary relics of warm water and geography as that between cold-humid and
fauna extensions during El Niño years. In Laguna glacier-influenced southern Patagonian fjords and
Grande, Navanax aenigmaticus apparently preys upon the northern Chilean and Peruvian Atacama desert
Bulla punctulata A. Adams in Sowerby, 1850, another coasts. Nevertheless, we found several marine
tropical species (see Uribe et al. 2013), which was also euthyneuran species occurring in both areas. Doris
just locally abundant in sandy subtidal sediments. fontainei (or Archidoris fontaini) is remarkable in show-
ing considerable colour variation (Schrödl 2003, 2009)
and having an enormous latitudinal extension, from
Euthyneura, Tectipleura, Panpulmonata, Mar del Plata in warm-temperate northern Argentina
Sacoglossa, Plakobranchacea, Limapontiidae, throughout the colder Magellan region to northern
Ercolania Trinchese, 1872 (non-tropical) Peru, as found herein. Molecular
studies are needed to explore the connectivity and
cf. Ercolania evelinae (Marcus, 1959) phylogeography of Pacific and Atlantic populations.
Prior to this study, Hancockia schoeferti was reported
Material.  Single specimen (ZSM Moll resin block 9k1), from Chile at 43-37° S; this area was sometimes re-
pier, Paracas, on algae, 25 November 2006. garded as transitional between the Magellanic and
Distribution.  Magellan Strait, Chile to Paracas, Peru Peruvian regions, or regarded as an own central
(first record of limapontiid sacoglossans for Peru). Chilean faunal entity characterized by having own
endemics (e. g. Brattström & Johanssen 1983, Schrödl
Remarks. The single tiny specimen externally 2003). Similarly, Corambe lucea appeared to show
resembled specimens collected in Chile (Schrödl an extended central Chilean distribution pattern
1996a). Secure generic placement requires histologi- (see Schrödl 2003), occurring from roughly 22° to
cal examination of the reproductive system. almost 44° S. Reporting H. schoeferti from southern
Peru and C. lucea even from northern (non-tropical)
Peru herein shows that increasing inventory may
Plakobranchidae, Elysia Risso, 1818 reject these and other hypotheses of endemism.
Considering the present records from the Peruvian
Elysia cf. hedgpethi Marcus, 1961 coast north to Bay of Sechura (approx. 6° N), C. lucea
Fig. 15 thus covers the entire range of the Peruvian faunal
province and, as several other Peruvian faunal
Material. Four specimens (ZSM Moll 20080519-22), elements, it extends a bit south into Magellanic
pier, Paracas, on algae, 26 November 2006. waters. This confirms earlier suggestions (Schrödl
Distribution.  Magellan Strait and Chilean coast to 1999b, 2003, 2009, Schrödl et al. 2005) that, at least
Paracas (Schrödl 2009, as E. patagonica) and Sechura for opisthobranchs, Chiloé Island represented an
herein; this appears to be the first record of a small- artificial, collection effort biased rather than a strict
sized green Elysia species for Peru that is backed by distributional border. The next candidate potentially
collected material. acting as a faunal barrier is the Peninsula Taitao (ca
46° S), separating the better explored northern part of
Remarks. Small, green specimens with small red the Chilean fjord region (Northern Patagonian Zone)
and blue dots collected from temperate Peruvian from the south (Häussermann & Försterra 2009).
waters herein externally resemble those from the Even more surprising was the record of the
Magellan Strait (Schrödl, 1996a) and from the central supposedly Magellanic Polycera priva, a previous
Chilean Bay of Coliumo (unpublished). Fischer & endemic to Patagonian fjords north of the Peninsula
Cervera (2005b) reported on (preserved) E. hedg­ Taitao (Schrödl 1996a, 1999b, 2003, 2009), from Islas
pethi from central Chile, and further green Elysia Ballestas in southern central Peru. Similarly, a few
specimens were found near Coquimbo (Martin Thiel, specimens of the common Magellanic chromodoridid

55
Tyrinna nobilis were found in northern Chile (Schrödl Peruvian warm water refuge Laguna Grande. At
& Millen 2001) and recently in Peru (Uribe et al. least sporadically it spreads far south to northern
2013). Overall, increasing collecting activities have and even central Chile (Schrödl 2007a), possibly with
shown that Magellanic opisthobranchs do not have warm, southward directed coastal currents during
Chiloé Island as a northern distributional border, but El Niño events. In summary, South-eastern Pacific
extend more or less far to the north with regularly opisthobranchs appear to be more widespread and
established, stable populations (Schrödl 1997b, 2003). flexible than expected from limited distributional
Southern South American species at least sporadi- data, extending their main ranges with more or less
cally extending such far to the north as suspected sporadic satellite populations. This may depend
for P. priva may be an overlooked but quite common on changing current regimes for larval dispersal
phenomenon that deserves further study in Peru. The and on suitable local environmental conditions, in
farther north established the greater may be the risk particular with regard to available potential prey
for populations of cool-water adapted species (and (sessile invertebrates in most cases).
their prey) to become locally and regionally extinct Even more strictly, splanchnotrophid parasites
latest with the next El Niño event. of the genus Ismaila depend on the presence of their
Hydrographic conditions are highly similar in specific opisthobranch hosts (Haumayr & Schrödl
the entire Humboldt current system (Brattström 2003, Anton & Schrödl 2013a). Interestingly, splanch-
& Johanssen 1983), and water temperatures are notrophids seem to be concentrated or limited to
frequently cooled down to 14 or 17 °C along the central and southern Chilean host populations
entire coast between central Chile and northern (Schrödl 2002), even in host species showing a wider
(non-tropical) Peru due to upwellings, especially Chilean-Peruvian distribution and dense Peruvian
during the summer with prevailing southern winds, populations, such as Okenia luna. Herein we found
and north of large peninsulas, forcing the Humboldt the first exception, referring to a single infested Phidi­
current away from the coast. Water masses and usu- ana lottini specimen among abundant conspecifics in
ally pelagic nudibranch larvae transported by the Callao, central Peru. On the one hand, the pelagic
Humboldt current may be responsible for short to nauplius larva of Ismaila implies a certain dispersal
long distance dispersal. This explains the extremes, potential necessary for initial infestation of remote
i. e. some Magellanic species inhabiting icy waters populations of host species. On the other hand, such
of southern Patagonia but also occurring in Peru, dispersive ability coupled with a short life cycle
at least locally and sporadically. High densities of would also allow for quick and massive infestation
C. lucea at several localities all along the Peruvian of entire host populations in Peru, as seen in central
coast indicate this species has reproductive local Chile (Schrödl 2003), and for following northern host
populations. We found H. schoeferti in a single, but populations during range shifts induced by El Niño
reproducing population in San Juan de Marcona, events. Because this is apparently not the case, i. e.
thousands of kilometres away from its northern- massive infestations by splanchnotrophids have yet
most previous record in central Chile, indicating to be observed in Peru, we may suspect on other
sporadic long distance dispersal and opportunistic potential constrains such as physiology or special
exploration of locally available resources. The larger population dynamics. Even a yet unknown, poten-
species Polycera priva has been found only once in tially sessile intermediate host may be considered,
Peru thus far, but with two specimens at the same limiting splanchnotrophid migration to and infesta-
site; for hermaphrodites enough to produce local tion of ephemeral host populations.
offspring. More data has to be collected in Peru to In total, herein we add 7 benthic sea slug spe-
see whether or not there are continuous and stable cies previously reported from Chile to the coastal
populations. Range extensions to Peru of other marine fauna of Peru. We may have found some of
species already known from the Chilean part of the species that were previously suspected to occur
the Peruvian faunal region, such as Janolus rebeccae in Peru, i. e. Rostanga cf. pulchra and Doto uva, but
and a Chilean morphotype of Doto uva, or known with conspecifity to their Californian and Chilean
ranges extended northwards within Peru are eas- look-alikes to be confirmed or rejected by genetic
ily explained by limited previous collecting efforts markers. However, we could not yet confirm the
along the Peruvian coast. In contrast to all these presence of Cadlina sparsa (see Schrödl 2000b) in Peru,
Magellanic or Peruvian-Chilean faunal elements, predicted by Álamo & Valdivieso (1987). In particu-
Navanax aenigmaticus appears to be a warm water lar, the common and charismatic Chilean nudibranch
adapted eastern Pacific, i. e. mainly Panamanian Thecacera darwini has not been found yet in Peru.
species (Ornelas-Gatdula et al. 2012), which, how- Further records augmenting the documented sea
ever, may have a stable population in the southern slug diversity of Peru will be published successively.

56
 Acknowledgements D’Orbigny, A. D. 1835-1846. Voyage dans l’Ameri-
que méridionale. Volúmen 5, parte 3, Mollusques.
Our thanks go to the friendly bus, taxi and mototaxi 758 pp., Paris (Bertrand).
drivers for transporting us and our collecting equipment Dall, W. 1909. Report on a collection of shells from Peru,
up and down along the Peruvian coast. We are very with a summary of the littoral marine mollusks of
grateful to all the fishermen and friends for diving the Peruvian zoological province. Proceedings of
equipment and boats. Room owners are thanked for the United States National Museum 37: 147-294.
tolerating our wet lab activities. IMARPE, the adminis- Dorgan, K. M., Valdés, Á. & Gosliner, T. M. 2002. Phy-
tration of the Reserva Nacional de Paracas, and the logenetic systematics of the genus Platydoris (Mol-
INRENA (SERNANP) are acknowledged for collecting lusca, Nudibranchia, Doridoidea) with descriptions
permits and logistic support. The German Research of six new species. Zoologica Scripta 31: 271-319.
Foundation (SCHR 667/4) and the GeoBioCenterLMU Fischer, M. A. & Cervera, J. L. 2005a. Baptodoris peruviana
supported field work. (D’Orbigny, 1837) comb. nov., an alternative taxo-
nomic placement for Doris peruviana (Gastropoda:
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