: WOODS HOLE Postil PHIC INSTITUTION
Peabody Museum OS | la Number 205

of Natural History
Yale University
New Haven, CT 06511

(Received Dy September 1988)
Abstract

The synonymy of the genera Pariphinotus and
Heterophlias has been debated many times in
the literature. Historically a distinction has
been maintained between these two phliantid
genera because of morphological differences
reported in the literature by the original
descriptors and subsequent workers. Our
examination of specimens of both genera
demonstrates Pariphinotus and Heterophlias
to be synonymous. Heterophlias has been
regarded as the valid genus by most authors;
Pariphinotus, however, is shown to be the
senior synonym of Heterophiias.

Key Words

Amphipoaa, Pariphinotus, Heterophiias,
synonymy, Phliantidae.

Contribution No. 1200 from the Bermuda Biological
Station for Research.

© Copyright 1989 by the Peabody Museum of
Natural History, Yale University. All rights reserved.
No part of this publication, except brief quotations
for scholarly purposes, may be reproduced without
the written permission of the Director, Peabody
Museum of Natural History.

ISBN No. 0-912532-18-3

RESEARCH .- 4
LIBRARIAN i U +

10 July 1989

ce

1889

Pariphinotus Kunkel, 1910,

the Senior Synonym of
Heterophlias Shoemaker, 1933
(Crustacea: Amphipoda:
Phliantidae)

Eric A. Lazo-Wasem
Adam J. Baldinger
Michael F. Gable

Introduction

The dinosaurian-like phliantid amphipods are
rarely collected (Barnard 1979); two
sporadically reported and morphologically
similar genera, Pariphinotus and Heterophlias,
have caused considerable debate over the
last three decades. Kunkel (1910) erected the
genus Pariphinotus from Bermuda, and
Shoemaker (1933) established the genus
Heterophlias from Dry Tortugas, Florida. For
decades the literature contained no mention
of these monotypic genera other than their
inclusion in Barnard’s 1958 Index. Not until the
next decade, primarily when Barnard (1962)
described a new subspecies of H. sec/usus
and when he first suggested that Pariphinotus
and Heterophlias might be synonymous
(Barnard 1969a), did these genera again
receive scientific attention. Mills (1964)
examined Kunkel’s two type specimens, but
because both were desiccated and the male,
damaged, he concluded only that Kunkel’s
(1910) description for Pariphinotus must stand.
Since the late sixties, two new species of
Heterophlias have been described (Ortiz 1976,
Barnard 1979). Range extensions for H.
seclusus (Wakabara and Pereira Leite 1977;
Nelson 1978, 1979) and H. seclusus
escabrosa (Barnard 1969b, c) have also been
given. The known distribution for the phliantids
under consideration in this paper now includes
semitropical and tropical marine waters of the
western Atlantic and eastern Pacific. In the

di WOODS HOLE Postilia “ INSTITUTION

Peabody Museum

Number 205

of Natural History 10 July 1989
Yale University
New Haven, CT 06511 RESEARCH 4
Nee Q
UBRARIAN (T° Uta 1989

ie
AY seri te a

(Received 27 September 1988)
Abstract

The synonymy of the genera Pariphinotus and
Heterophlias has been debated many times in
the literature. Historically a distinction has
been maintained between these two phliantid
genera because of morphological differences
reported in the literature by the original
descriptors and subsequent workers. Our
examination of specimens of both genera
demonstrates Pariphinotus and Heterophlias
to be synonymous. Heterophlias has been
regarded as the valid genus by most authors;
Pariphinotus, however, is shown to be the
senior synonym of Heterophiias.

Key Words

Amphipoda, Pariphinotus, Heterophlias,
synonymy, Phliantidae.

Contribution No. 1200 from the Bermuda Biological
Station for Research.

© Copyright 1989 by the Peabody Museum of
Natural History, Yale University. All rights reserved.
No part of this publication, except brief quotations
for scholarly purposes, may be reproduced without
the written permission of the Director, Peabody
Museum of Natural History.

ISBN No. 0-912532-18-3

Pariphinotus Kunkel, 1910,

the Senior Synonym of
Heterophlias Shoemaker, 1933
(Crustacea: Amphipoda:
Phliantidae)

Eric A. Lazo-Wasem
Adam J. Baldinger
Michael F. Gable

Introduction

The dinosaurian-like phliantid amphipods are
rarely collected (Barnard 1979); two
sporadically reported and morphologically
similar genera, Pariphinotus and Heterophiias,
have caused considerable debate over the
last three decades. Kunkel (1910) erected the
genus Pariphinotus from Bermuda, and
Shoemaker (1933) established the genus
Heterophlias from Dry Tortugas, Florida. For
decades the literature contained no mention
of these monotypic genera other than their
inclusion in Barnard’s 1958 Index. Not until the
next decade, primarily when Barnard (1962)
described a new subspecies of H. sec/usus
and when he first suggested that Pariphinotus
and Heterophlias might be synonymous
(Barnard 1969a), did these genera again
receive scientific attention. Mills (1964)
examined Kunkel’s two type specimens, but
because both were desiccated and the male,
damaged, he concluded only that Kunkel’s
(1910) description for Pariphinotus must stand.
Since the late sixties, two new species of
Heterophlias have been described (Ortiz 1976,
Barnard 1979). Range extensions for H.
seclusus (Wakabara and Pereira Leite 1977;
Nelson 1978, 1979) and H. seclusus
escabrosa (Barnard 1969b, c) have also been
given. The known distribution for the phliantids
under consideration in this paper now includes
semitropical and tropical marine waters of the
western Atlantic and eastern Pacific. In the

2 Pariphinotus, Senior Synonym
of Heterophlias

Atlantic, species extend from North Carolina
(Nelson 1978, 1979) and Bermuda (Kunkel
1910 and recent collections) to approximately
20° S off the coast of Brazil (Wakabara and
Pereira Leite 1977). In the Pacific, species can
be found from Cayucos, California, to the
Galapagos Islands (Barnard 1979).

In addition to the description of new
species and the extension of species’ ranges,
debate on the synonymy of Pariphinotus and
Heterophlias has continued (Barnard 1972,
1979, 1981; Wakabara and Pereira Leite 1977;
Ledoyer 1982). This debate has remained
speculative because specimens of
Pariphinotus from Bermuda were unavailable
or assumed to be so. However, recent
collections of Pariphinotus in Bermuda and a
re-examination of the female paralectotype of
P. tucker, deposited in the Peabody Museum
of Natural History, Yale University (YPM), have
allowed us to clarify definitively the questions,
problems and ambiguities presented in and
raised by the literature regarding the status of
Pariphinotus and Heterophiias.

Morphology and Systematics

The focus of the debate on the possible
synonymy of Pariphinotus and Heterophlias
has centered on three morphological features:
the third uropod, the mandibular molar, and
the inner lobes of the lower lip.

Kunkel (1910) described P. tuckeri as
lacking a third uropod; Shoemaker (1933)
described H. seclusus as possessing a third
uropod consisting of a single thick joint
(=peduncle, no rami). This difference has
been the main diagnostic feature separating
the two genera (Barnard 1969a; Wakabara
and Pereira Leite 1977). Wakabara and Pereira
Leite (1977) pointed out that the third uropod
of Heterophlias is difficult to discern because
it lies hidden dorsally by the large telson. They
believed that Kunkel (1910) might have
overlooked it. For the same reason, Barnard
(personal communication) urged our
examination of Bermuda material to determine
the presence or absence of the third uropod.
Despite Mills’ (1964) dismissal of the

Postilla 205

Pariphinotus type specimens, our re-
examination of the desiccated female, a
relative giant at 5.5 mm, revealed that Kunkel
did indeed overlook uropod 3. Also, recently
collected specimens from Bermuda
[collections of the YPM and the United States
National Museum (USNM)] all possess a third
uropod, without rami, and therefore
Pariphinotus and Heterophlias are
indistinguishable genera in this important
respect.

One recently collected Bermuda specimen
possesses third uropods that
uncharacteristically extend beyond the telson
(Fig. 1). The specimen is medium-sized and
the condition therefore is not allometric.
Wakabara and Pereira Leite’s (1977) figure of
a female Heterophlias exhibits a similar
variation of the telson/uropod three complex.

The mandibular molar is lacking in
Pariphinotus according to Kunkel’s (1910)
description (no figure provided); in
Heterophlias, the mandibular molar is present
as a conical projection terminating in a large
spine. Although for Heterophlias the
mandibular molar is variably described in
prose, in figures it always appears similar with
one minor exception, the figure of Wakabara
and Pereira Leite (1977) in which there is no
distinction between projection and spine. In
1979 Barnard believed that a generic
distinction should be maintained because of
the presence or absence of a molar. Kunkel
(1910) indeed stated that there is no molar in
Pariphinotus, but careful reading of his
description suggests he actually did see one,
for he went on in the same sentence to
describe a “‘spine row with a single spine,” a
phrase which we interpret as a case of
mistaken identity, |.e., he was actually
describing the molar. Whatever the case
historically, our examination of the mandibles
of recently collected P. tuckeri from Bermuda
(Fig. 1) reveals a molar matching precisely
that described by Shoemaker (1933) for H.
seciusus and by Barnard (1979) for H.
galapagoanus. Pariphinotus and Heterophlias
are therefore indistinguishable genera with
respect to their mandibular molars.

Kunkel (1910) described Pariphinotus as not

3 Pariphinotus, Senior Synonym
of Heterophlias

Postilla 205

tele

»

AE
Fig. 1

Pariphinotus tuckeri Kunkel. YPM No. 8720: U3,
uropod 3. USNM Acc. No. 346847: Md, mandible;
LL, lower lip. Scale bar = 0.1 mm.

having inner lobes on the lower lip and there
was no figure. Wakabara and Pereira Leite
(1977) noted that Heterophiias differs from
Pariphinotus in possessing inner lobes on the
lower lip; inner lobes were also described by
Shoemaker (1933). Wakabara and Pereira
Leite (1977) suggested that Kunkel may have
overlooked the inner lobes. Recently collected
specimens from Bermuda do have inner lobes
on the lower lip (Fig. 1); they are, however,
extremely difficult to discern, particularly if the
lip is viewed from the opposite side.
(Barnard’s (1979) figure of H. galapagoanus
does not show inner lobes on the lower lip,
but he obviously figured the lip from the
outside.) The presence or absence of inner
lobes on the lower lip is, then, also not a

difference between Pariphinotus and
Heterophlias.

Recently, another character was mentioned
in the literature that differs from our
observations for the Pariphinotus/Heterophlias
complex. Ledoyer (1986) enigmatically
suggested the presence of a vestigial
mandibular palp for Heterophlias. A diagnostic
feature of the Phliantidae is the lack of a
mandibular palp, and we can find nothing in
the literature on Heterophlias to which such a
statement could be ascribed.

As already mentioned, the possibility of a
synonymy for Pariphinotus and Heterophlias
has been raised several times in the last few
decades. Although Barnard (1979) stated that
Pariphinotus is probably a senior synonym of

4 Pariphinotus, Senior Synonym
of Heterophlias

Heterophlias, all other references to a
synonymy, including a later one by Barnard
(1981), implied Heterophlias should be the
valid generic name, in the event a synonymy
could be demonstrated. Although Kunkel’s
(1910) description of Pariphinotus lacked a
detailed analysis of the mouthparts and was
incorrect with respect to uropod 3,
Shoemaker’s (1933) Heterophlias, as
demonstrated in this paper, undoubtedly
belongs to the same genus. As Kunkel’s
description of Pariphinotus preceded the
description of Heterophlias, the latter must be
regarded as a junior synonym of Pariphinotus.
A synonymy and emended description for
Pariphinotus Kunkel, 1910, are provided.

Pariphinotus Kunkel 1910 (emended)

Pariphinotus: Kunkel, 1910:19. Barnard, 1958:
111. Barnard, 1964:67. Barnard, 1969a:411,
Barnard, 1981:1214, 1216.

Heterophlias: Shoemaker, 1933:250. Barnard,
1958:110. Barnard, 1964:67. Barnard, 1969a:
410. Barnard, 1981:1214, 1216. Ledoyer, 1982:
14.

Pariphinotus tuckeri: Kunkel, 1910:19-21, fig.
6. Barnard, 1958:111. Mills, 1964:2-3. Barnard,
1969a:411. Johnson, 1986:378-79, fig. 125.
Heterophlias seclusus: Shoemaker, 1933:250-
52, figs. 4-5. Barnard, 1958:110. Barnard,
1969a:410, figs. 145-47. Barnard, 1972:193.
Wakabara and Pereira Leite, 1977:90-96, figs.
1-4. Nelson, 1978:103. Nelson, 1979:66.
Heterophlias seclusus escabrosa: Barnard,
1962:79-80, fig. 5. Barnard, 1969b:195-96.

Barnard, 1969c:219. Barnard, 1979:131, fig. 40.
Heterophlias seticoxae: Ortiz, 1976:21-35, figs.

l=3:

Heterophlias galapagoanus: Barnard, 1979:
IGUSSS) else HOS

Type Species

Pariphinotus tuckeri Kunkel.

Postilla 205

Type Locality

Bermuda.

Diagnosis

Mandibular molar conical, terminating in a
large spine, lower lip possessing inner lobes,
maxilla 1 lacking palp, maxillipedal palp
4-articulate, gnathopods simple, inner ramus
of pleopod 3 one-half length of outer ramus,
uropod 3 lacking rami.

Material Examined

YPM 5613. 5.5 mm 8. Paralectotype.
Bermuda. W. G. Van Name. 1901. YPM 8720.
4.24 mm @. Ferry Reach, St. George's,
Bermuda. M. F. Gable. 25 May 1985. YPM
8739. 2.7 mm &. North side of Shelly Bay,
Hamilton, Bermuda. E. A. Lazo-Wasem. 28
May 1987. Depth: 0.6m. Within pieces of
limestone on sandy bottom. YPM 8740. 1.6mm
immature. North side of Shelly Bay, Hamilton,
Bermuda. E. A. Lazo-Wasem. 28 May 1987.
Depth: 0.6m. Within pieces of limestone on
sandy bottom. YPM 8741. 3.9 mm 6.
Whalebone Bay, St. George’s, Bermuda. A. J.
Baldinger. 22 May 1987. Depth: 1m. From
Thalassia. USNM 346847. Bermuda. Ferry
Reach, St. George's, Bermuda. M. L. Jones. 2
September 1981.

Acknowledgments

This study was supported in part by a Samuel
Riker Fellowship from the Bermuda Biological
Station to the first author. Funding to the first
author was also provided by the Peabody
Museum of Natural History, Yale University,
and the Eastern Connecticut State University
Foundation. A Connecticut State University
Research Grant provided partial support for
the third author.

5 Pariphinotus, Senior Synonym Postilla 205
of Heterophlias

Literature Cited

Barnard, J. L. 1958. Index to the families, genera, and species of the gammaridean Amphipoda
(Crustacea). Allan Hancock Foundation Publ., Occ. Pap. 19, 145 p.

1962. Benthic marine Amphipoda of southern California 2. Families Tironidae to Gammaridae. Pac.
Naturalist 3:73-115.

1964. Revision of some families, genera and species of gammaridean Amphipoda. Crustaceana 7:

49-74.
1969a. The families and genera of marine gammaridean Amphipoda. Bull. U.S. Nat. Mus. 271: 1-

535.

1969b. Gammaridean Amphipoda of the rocky intertidal of California: Monterey Bay to La Jolla.
Bull. U.S. Nat. Mus. 258: 1-230.

1969c. A biological survey of Bahia de Los Angeles, Gulf of California, Mexico, IV: Benthic
Amphipoda (Crustacea) Trans. San Diego Soc. Nat. Hist. 15:175-228, figs. 1-30.

1972. The marine fauna of New Zealand: Algae-living littoral Gammaridea (Crustacea Amphipoda).
N. Z. Oceanogr. Inst. Mem. 62:7-216, figs. 1-109.

1979. Littoral gammaridean Amphipoda from the Gulf of California and Galapagos Islands.
Smithson. Contrib. Zool. 271:1-149, figs. 1-74.

1981. Redescription of /phiplateia whiteleggei, a New Guinea marine amphipod (Crustacea). Proc.
Biol. Soc. Washington 94:1211-18.

Johnson, S. E. 1986. Order Amphipoda, P. 372-81. In W. Sterrer (ed.), Marine fauna and flora of
Bermuda. New York, John Wiley & Sons.

Kunkel, B. W. 1910. The Amphipoda of Bermuda. Transactions of the Connecticut Academy of Arts and
Sciences 16:1-116.

Ledoyer, M. 1982. Crustaces Amphipodes Gammariens, Familles des Acanthonotozomatidae a
Gammaridae. Faune de Madagascar 59(1):1-598.

1986. Crustaces Amphipodes Gammariens, Familles des Haustoriidae a Vitjazianidae. Faune de
Madagascar 59(2):599-1112.

Mills, E. L. 1964. Noteworthy Amphipoda (Crustacea) in the collection of the Yale Peabody Museum.
Postilla (Yale Peabody Mus. Nat. Hist.) 79:1-41.

Nelson, W. G. 1978. An occurrence of Heterophlias seclusus Shoemaker, 1933 (Amphipoda, Phliantidae)
at Beaufort, North Carolina, U.S.A. Crustaceana 35:103.

1979. Additions to the amphipod crustaceans of North Carolina. Estuaries 2:66.

Ortiz, M. 1976. Un neuvo anfipodo de aguas Cubanas (Amphipoda, Gammaridea, Phliantidae). Ciencias,
La Habana (Investigaciones Marinas) 25:21-35.

Shoemaker, C. R. 1933. Two new genera and six new species of Amphipoda from Tortugas. Carnegie
Inst. Washington Publ. 435:245-56.

Wakabara, Y. and F. P. Pereira Leite. 1977. Heterophlias seclusus Shoemaker, 1933 (Amphipoda,
Phliantidae) from the Brazilian coast. Crustaceana 33:90-96.

The Authors

Eric A. Lazo-Wasem. Division of
Invertebrate Zoology, Peabody Museum of
Natural History, Yale University, New Haven,
CT 06511.

Adam J. Baldinger. Department of Biology,
Eastern Connecticut State University,
Willimantic, CT 06226.

Michael F. Gable. Department of Biology,
Eastern Connecticut State University,
Willimantic, CT 06226.

vs waS

=
=
5 .