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Spring 5-2008
A Systematic and Taxonomic Review of the Family
Pseudotanaidae (Crustacea: Peracarida: Tanaidacea) Based
Primarily on Morphometry Cladistic Analyses
Jerry Alan McLelland
University of Southern Mississippi
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Peracarida: Tanaidacea) Based Primarily on Morphometry Cladistic Analyses" (2008). Dissertations.
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The University of Southern Mississippi
A SYSTEMATIC AND TAXONOMIC REVIEW OF THE FAMILY PSEUDOTANAIDAE
(CRUSTACEA: PERACARIDA: TANAIDACEA) BASED PRIMARILY ON
MORPHOMETRY CLADISTIC ANALYSES
by
Jerry Alan McLelland
A Dissertation
Submitted to the Graduate Studies Office
of The University of Southern Mississippi
in Partial Fulfillment of the Requirements
for the Degree of Doctor of Philosophy
May 2008
COPYRIGHT BY
JERRY ALAN McLELLAND
2008
The University of Southern Mississippi
A SYSTEMATIC AND TAXONOMIC REVIEW OF THE FAMILY PSEUDOTANAIDAE
(CRUSTACEA: PERACARIDA: TANAIDACEA) BASED PRIMARILY ON
MORPHOMETRIC CLADISTIC ANALYSES
by
Jerry Alan McLelland
Abstract of a Dissertation
Submitted to the Graduate Studies Office
of The University of Southern Mississippi
in Partial Fulfillment of the Requirements
for the Degree of Doctor of Philosophy
May 2008
ABSTRACT
A SYSTEMATIC AND TAXONOMIC REVIEW OF THE FAMILY PSEUDOTANAIDAE
(CRUSTACEA: PERACARIDA: TANAIDACEA) BASED PRIMARILY ON
MORPHOMETRY CLADISTIC ANALYSES
by
Jerry Alan MeLelland
May 2008
Species of the tanaid family Pseudotanaidae, established by Jurgen Sieg in 1973, have
been reported from all the world's oceans in depths ranging from near the surface to below 6,000
m. The family currently consists of 46 characteristically small tanaidomorph species residing in
two subfamilies, the Cryptocopinae (seven species) and the Pseudotanainae (39 species).
Members of the more ancestral Crytocopinae differ from the Pseudotanainae by having females
with four rather than three antennule articles and a complete separation of the maxilliped endites.
The primary objective of this study was to test the hypothesis that the current structure of the
family Pseudotanaidae should remain intact as a monophyletic group or, alternatively, should be
pared to exclude the members of the current subfamily Cryptocopinae. A heuristic search using
the parsimony optimality criterion was conducted on 69 unordered and unweighted
morphological characters to test the relationships among 53 species of Pseudotanaidae and four
out-groups. Six equally parsimonious trees were obtained that showed substantial support values
at the node dividing Pseudotanainae from Cryptocopinae, thus providing strong evidence for
removing Crytocopinae from the family Pseudotanaidae. A new family, Cryptocopidae is
proposed containing the subfamilies Cryptocopinae (one species), Cryptocopoidinae (three
species), and Iungentitanainae (four species). The remaining 46 species of Pseudotanaidae
demonstrated inconsistent structure in the cladistic analysis due to the large amount of homoplasy
among the characters used, thus suggesting a revised taxonomy based more on observed
ii
groupings of character traits rather than support values. Three subfamilies are recognized: the
Akanthinotanainae (ten species), the Parapseudotanainae (two species) and the Pseudotanainae
(33 species). The more ancestral Akanthinotanainae is represented solely by the new genus
Akanthinotanais, a former subgenus of Pseudotanais. Two new sub-genera within the
Akanthinotanainae are recognized: Akanthinotanais, s.s. with nine species and Guilleitanais
represented by a single species, A. guillei, with unique ancestral characters. Among the
Pseudotanainae, a single genus, Pseudotanais, and two sub-genera, Pseudotanais, s.s. and the
new Mystriocentrus, are recognized. Seven new taxa including four species of Pseudotanais,
s.s., one of Pseudotanais (Mystriocentrus) and one each of the genera Parapseudotanais and
Cryptocopoides were discovered during this research.
in
ACKNOWLEDGEMENTS
I wish to thank first and foremost my advisor, Dr. Richard Heard, whom I consider an
esteemed friend and colleague, for providing the impetus for this dissertation. His boundless
spirit, scientific curiosity and phenomenal grasp of the 'big picture' has inspired many students
over the years and started them on paths toward their academic goals. Likewise, it was he who
piqued my interest in the study of tanaids, as he similarly did when I earlier aspired to study
polychaete annelids, and suggested the problem concerning the family Pseudotanaidae.
I have had the help of several outside experts in the field of tanaid zoology. First among
these is Dr. Kim Larsen who, in the mold of the late Jurgen Sieg, has become a legend in the
tanaid world for his grasp of the systematics of the entire order. In his role as external advisor to
my dissertation, he corrected mistakes and misconceptions made early in this study and critiqued
an early draft of this dissertation. During his postdoctoral work at Gulf Coast Research
Laboratory (GCRL, Ocean Springs, Mississippi), he conducted fine sorting and identification of
the Texas A&M Gulf of Mexico samples which provided me with most of the material for this
research, and also proved to be a daunting Starcraft opponent. 1 wish to thank others of the tanaid
community including Drs. Graham Bird, Jurgen Guerro-Komritz, Roger Bamber and Magdalena
Blazewicz-Paszkowycz for providing literature or commenting on early drafts of species
descriptions.
1 thank Drs. Chet Rakocinski, Jeffrey Lotz, Bruce Comyns and Jinx Campbell for
serving on my graduate committee as well as teaching courses and offering their advice during
my years as a doctoral student in Coastal Sciences. Dr. Campbell was especially helpful in
teaching me the phylogenetic methods used in this research, providing the computer programs
required and giving advice when needed on the proper procedures for conducting a cladistic
analysis with morphological data.
Others 1 wish to thank include Drs. David Knott and Rachel King of the Southeastern
Regional Taxonomic Center (SERTC, Charleston, South Carolina) for providing specimens of
iv
Cryptocopoides collected off the east coast of the United States; Drs. Irvin Otvos and Alex
Pozhitkov (GCRL) for translating Russian literature; Cathy Schloss (GCRL Gunter library) for
procuring interlibrary loan material; Sara LeCroy (GCRL museum curator ) for making her
scanner available and for helping obtain loaned specimens; Dr. Stephen (Ash) Bullard for advice
on formatting and other matters involved in putting this dissertation together; Dr. William
Hawkins, GCRL director, for his encouragement of my research and providing working space;
and the numerous other faculty, students and staff of GCRL I have worked with over the last 33
years.
I am grateful to my fellow students, especially John Foster, Guillermo Sanchez, Jana
Thoma, Brent Thoma, and David Drumm for their advice and encouragement in the day to day
tedium of conducting this research. Brent Thoma provided invaluable assistance in teaching me
the techniques of the DELTA, Photoshop, and PAUP computer programs. David Drumm taught
me to use the computer program, TreeRot, a valuable tool for computing decay indices on
phylogenetic trees.
Most importantly, I wish to acknowledge the love, support and encouragement of my
family including my parents, Charner and Doris McLelland, my sons Marcus and Gregory
McLelland and my three brothers and numerous in-laws. Finally, my wife Margaret deserves my
utmost gratitude for her perseverance, love and unflagging support throughout the years of my
career but especially during the trying times of long hours and late nights during my doctoral
research; it is to her I dedicate this dissertation.
v
TABLE OF CONTENTS
ABSTRACT
ii
ACKNOWLEDGEMENTS
iv
LIST OF ILLUSTRATIONS
vii
LIST OF TABLES
xi
CHAPTER
I. INTRODUCTION
Systematics and General Information about the Study Group
Problem Statement
Significance of Study
1
II. REVIEW OF RELATED LITERATURE
Foundational literature
Geo-distribution
Oculate species and vertical distribution
5
III. METHODOLOGY
Material studied
Laboratory Techniques
Data Management and Character Selection
Data Analysis
12
IV. RESULTS OF DATA ANALYSIS
Parsimony Analysis
Summary of Results
18
V. SYSTEMATICS AND TAXONOMY
Proposed Systematic Revision
Taxonomy of Pseudotanaidae
Taxonomy of Cryptocopidae
Key to Species of Cryptocopidae and Pseudotanaidae
29
APPENDIXES
A. Gulf of Mexico Station Data
186
B. Description and diagnostics of most parsimonious tree from parsimony analysis .190
C.
Character Matrix for Phylogenetic Analysis of Pseudotanaidae
192
D.
Synopsis of Characters
199
REFERENCES
278
VI
LIST OF ILLUSTRATIONS
Strict consensus of six equally parsimonious trees of the family
Pseudotanaidae
22
Jackknife consensus of six equally parsimonious trees of the family Pseudotanaidae ....23
Bootstrap consensus of six equally parsimonious trees of the family Pseudotanaidae
24
Strict consensus of six equally parsimonious trees of the family Pseudotanaidae omitting
the character for marsupium development
25
Proposed phylogeny of Pseudotanaidae and Crytocopidae
30
Parapseudotanais sp. A-holotype dorsal and lateral views
55
Parapseudotanais sp. A - paratype cheliped, pleotelson, uropod, pleopod, antennule,
and antenna
56
Parapseudotanais sp. A - paratype mouthparts - mandibles, labrum, labium, maxillule,
and maxillipeds
57
Parapseudotanais sp. A - paratype pereopods 1 to 6
Pseudotanais {Mystriocentrus) sp. A - holotype dorsa! and lateral views
58
68
Pseudotanais {Mystriocentrus) sp. A - paratype cheliped, antennule, antenna, pleopod,
and uropod
69
Pseudotanais {Mystriocentrus) sp. A - paratype mandibles, labrum, labium, maxillule,
and maxillipeds
70
Pseudotanais {Mystriocentrus) sp. A - paratype pereopods 1 to 6
Pseudotanais {Pseudotanais) sp. A - holotype dorsal and lateral views
71
124
Pseudotanais {Pseudotanais) sp. A - paratype cheliped, antennule, antenna, pleopod, and
uropod
125
Pseudotanais {Pseudotanais) sp. A - paratype mandibles, labrum, maxillipeds, maxillule
and epignath
126
Pseudotanais {Pseudotanais) sp. A - paratype pereopods 1 to 6
127
Pseudotanais {Pseudotanais) sp. C - holotype dorsal and lateral views
134
Pseudotanais {Pseudotanais) sp. C - paratype cheliped, antennule, antenna, pleopod and
uropod
135
vii
20.
Pseudotanais {Pseudotanais) sp. C - paratype mandibles, labrum, labium, maxillipeds,
epignath and maxillule
136
21.
Pseudotanais {Pseudotanais) sp. C - paratype pereopods 1 to 6
137
22.
Pseudotanais {Pseudotanais) sp. O - holotype dorsal and lateral views
144
23.
Pseudotanais {Pseudotanais) sp. O - paratype cheliped, antennule, antenna, uropod and
pleopod
145
Pseudotanais {Pseudotanais) sp. O - paratype mandibles, maxillule, maxilliped, labium
and labrum
146
24.
25.
Pseudotanais {Pseudotanais) sp. O - paratype pereopods 1 to 6
26.
Pseudotanais {Pseudotanais) sp. P - holotype dorsal and lateral views
27.
Pseudotanais {Pseudotanais) sp. P - paratype antennule, antenna, pleopod, cheliped,
pleotelson and uropods
155
28.
Pseudotanais {Pseudotanais) sp. P - paratype epignath, labrum, labium, mandibles,
maxilliped and maxillule
147
154
156
29.
Pseudotanais {Pseudotanais) sp. P - paratype pereopods 1 to 6
30.
Cryptocopoides sp. A - holotype dorsal and lateral views; paratype cheliped, antennule,
antenna, pleotelson, uropod, and pleopod
157
170
31.
Cryptocopoides sp. A - paratype mandibles,maxilliped, maxillule endite and labrum ..171
32.
Cryptocopoides sp. A-paratype pereopods 1 to 6
172
33.
Character 1 trace - cephalothorax, length vs. pereonites 1 to 3 length
218
34.
Character 2 trace - eyelobe, presence or absence
219
35.
Character 3 trace - pereonite abbreviation
220
36.
Character 5 trace - antennule, number of articles
221
37.
Character 6 trace - antennule, article 1 percent of total length of antennule
222
38.
Character 7 trace - antennule, article 1 length vs. width
223
39.
Character 8 trace - antennule, article 3 length vs. width
224
40.
Character 9 trace - antennule, distal article length vs. width
225
41.
Character 11 trace - antenna, length vs. antennule length
226
42.
Character 12 trace - antenna, article 2 length vs. article 3 length
viii
227
43.
Character 13 trace - antenna, articles 2 and 3 setal type
228
44.
Character 14 trace - antenna, article 4 length vs. width
229
45.
Character 15 trace - antenna, article 5 length vs. width
230
46.
Character 17 trace -mandible, pars molaris width
231
47.
Character 18 trace - mandible, pars molaris complexity
232
48.
Character 21 trace - maxillule, number of endite terminal spiniform setae
233
49.
Character 23 trace - maxilliped, basis degree of fusion
234
50.
Character 24 trace - maxilliped, endite degree of fusion
235
51.
Character 25 trace - maxilliped, endite setation
236
52.
Character 26 trace - cheliped, degree of forcipation
237
53.
Character 27 trace - cheliped, carpus length vs. width
238
54.
Character 28 trace - cheliped, carpus number inferior setae
239
55.
Character 29 trace - cheliped, propodus length vs. width
240
56.
Character 30 trace-cheliped, propodus length vs. length of carpus
241
57.
Character 31 trace - cheliped, fixed finger number inferior setae
242
58.
Character 32 trace - cheliped, dactylus percent of propodus length
243
59.
Character 33 trace - cheliped, dactylus width vs. fixed finger width
244
60.
Character 34 trace - cheliped, dactylus setation
245
61.
Character 35 trace - marsupium formation
246
62.
Character 37 trace - pereopod 1, basis length vs. width
247
63.
Character 38 trace - pereopod 1, basis setation
248
64.
Character 39 trace- pereopod 1, ischium setation
249
65.
Character 40 trace - pereopod 1, merus setation
250
66.
Character 41 trace - pereopod 1, carpus setation
251
67.
Character 42 trace - pereopod 1, propodus length vs. width
252
ix
68.
Character 43 trace - pereopod 1, propodus setation
253
69.
Character 44 trace - pereopod 1, dactylus setation
254
70.
Character 45 trace - pereopod 1, dactylus setal type
255
71.
Character 46 trace - pereopod 1, length of dactylus + unguis
256
72.
Character 47 trace - pereopods 2 and 3, basis setation
257
73.
Character 48 trace - pereopods 2 and 3, ischium setation
258
74.
Character 49 trace - pereopods 2 and 3, merus setation
259
75.
Character 50 trace-pereopods 2 and 3, merus spatulate setae
260
76.
Character 51 trace -pereopods 2 and 3, carpus blade-like setae
261
77.
Character 52 trace - pereopods 2 and 3, carpus blade-like setae length
262
78.
Character 53 trace - pereopods 2 and 3, dactylus + unguis length
263
79.
Character 55 trace - pereopod 4, basis length vs. width
264
80.
Character 56 trace - pereopods 4 to 6, basis setation
265
81.
Character 57 trace - pereopods 4 to 6, ischium setation
266
82.
Character 58 trace - pereopods 4 to 6, merus setation
267
83.
Character 59 trace - pereopods 4 to 6, carpus blade-like setae
268
84.
Character 60 trace - pereopods 4 to 6, carpus additional specialized setae
269
85.
Character 61 trace - pereopods 4 to 6, propodus broom setae
270
86.
Character 62 trace - pereopods 4 to 6, dactylus fusion with unguis
271
87.
Character 63 trace - pleopod complexity
272
88.
Character 65 trace - uropod, number of exopod articles
273
89.
Character 66 trace — uropod, exopod length vs. endopod length
90.
Character 67 trace - uropod, exopod proximal article proportion of ramus
275
91.
Character 68 trace - uropod, number of endopod articles
276
92.
Character 69 trace - uropod, endopod proximal article proportion of ramus
277
x
274
LIST OF TABLES
Table
1.
Scores of most parsimonious trees from heuristic searches on datasets with and without
the character for marsupium development
19
2.
Score comparison of unweighted and re-weighted trees
XI
20
1
CHAPTER I
INTRODUCTION
Systeraatics and General Information about the Study Group
Order Tanaidacea.
The order Tanaidacea belongs to the crustacean superorder Peracarida (Malacostraca:
Peracarida) which also contains the common orders Amphipoda, Isopoda, Mysida, and Cumacea
(see Martin and Davis 2001:32, for an updated classification of the Peracarida). Peracarids are
distinguished from other crustaceans by the presence of a marsupium (brood pouch) in which the
females carry eggs until they develop and emerge as "mancas" or juveniles. This degree of
parental care and lack of a planktonic larval stage typifies the niche to which this group belongs
opposite to that of the more fecund crustacean groups such as decapods and may explain the
success of the peracarids in diverse habitats including those in marine, freshwater and even
terrestrial environments; the group represents a wide variety of trophic lifestyles (Brusca and
Brusca 2003). Tanaidaceans are usually considered to be a minor taxonomic group of
peracarids, although they may constitute one of the most bionomically significant groups in a
wide range of marine habitats from intertidal (Holdich and Jones 1983) to hadal depths (Wolff
1956). There are currently over 800 species of Tanaidacea known, most of which live at depths
greater than 200 m with some species occurring as deep as 9000 m (Sieg 1983, Heard 2002,
Larsen 2005, Anderson et al. 2007). Deep water tanaids are among the most abundant and
diverse assemblages present in abyssal zones of the world. At depths of 5000 m they comprise as
much as one-fifth of the macrobenthic biomass (Wolff 1977) and have thus been regarded as a
highly important component in the deep water benthic food web (Gardiner 1977, Ogle et al. 1982,
Sieg and Heard 1989, Kneib 1992, Escobar-Briones and Spears 1995, Gutu and Sieg 1999).
Most tanaidaceans are small (2-5 mm long) but adults can range in size from 0.5 to 120
mm (Heard 2002). They were originally separated into two families Apseudidae (Leach 1814)
2
and Tanaidae (Dana 1849) by Sars (1882) and then reorganized into the suborders
Monokonophora and Dikonophora by Lang (1956) based on characteristics of the 'genital cones'
in males. Sieg (1980), in his revision of the order, designated the extant suborders
Apseudomorpha, Tanaidomorpha, Neotanaidomorpha, and the extinct Anthracocaridomorpha
based on general morphology of the body and characteristics of the mouthparts; this construction,
which generally reflects differences seen in females, is the standard currently accepted by
tanaidacean workers. In their synopsis of the order, Gutu and Sieg (1999) recognized the above
four suborders consisting of 25 families, 163 genera, and nearly 670 described species. The suite
of characters that are considered (Heard 2002) to characterize the order Tanaidacea include (1) a
carapace formed via fusion of the first two thoracic somites, (2) chelate first pereopods, (3)
usually six free thoracic somites, (4) five abdominal somites usually bearing pleopods and (5) a
pleotelson with a pair of terminal or subterminal uropods. It should be pointed out that none of
these characteristics individually are unique to the Tanaidacea; each can be applied in some
degree to other peracaridean groups.
Suborder Tanaidomorpha
Members of the suborder Tanaidomorpha Sieg, 1980, referred to herein as
'tanaidomorphs', are distinct from the other major suborder, the Apseudomorpha, by generally
being more cylindrical in shape, having a reduction in body parts such as the loss of mandibular
palps and antenular accessory flagella, and demonstrating strong sexual dimorphism whereby the
males are more eccentric looking than females and are usually smaller and adapted for
swimming. There are 13 families within the suborder Tanaidomorpha and all but one (Tanaidae)
fall within the superfamily Paratanaoidea Lang, 1949.
Family Pseudotanaidae
The family Pseudotanaidae was established by Sieg (1973) to include those tanaidomorph
species with the marsupium of females formed by a single pair of sheet-like oostegites emerging
at the basis of the fourth pereopod. Furthermore, Sieg's original diagnosis of females included
3
(1) reduced anterior pereonites, (2) fused maxilliped bases and complete or partial fusion of the
maxilliped endites, (3) an antennule of three or four articles, and (4) the reduction of the mandible
pars molaris, in most cases to a fine point. As Bird and Holdich (1989) observed, none of these
characteristics are limited to the Pseudotanaidae, with the possible exception of the marsupium
formation, but can be found among other tanaidomorph species. Most members of the family are
characteristically small, averaging 1.5 mm in length (Sieg 1986), which is advantageous in deepsea environments with a limited food supply (Jumars et al. 1990).
In his 1977 monograph of the family, Sieg grouped the 21 known species of
Pseudotanaidae into two subfamilies, Cryptocopinae Sieg, 1977 and Pseudotanainae Sieg, 1977,
based largely on the number of antenular articles (four and three respectively). The
Cryptocopinae was represented by four genera: Cryptocope G.O. Sars, 1882; Cryptocopoides
Sieg, 1977; Iungentitanais Sieg, 1977; and Paraiungentitanais Sieg, 1977. The subfamily
Pseudotanainae contained most of the family's described species but had only the type genus
Pseudotanais. Sieg further divided Pseudotanais into the subgenera Pseudotanais sensu stricto
and Akanthinotanais based largely on the presence in the former of blade-like carpal spines on the
posterior three pereopods. Bird and Holdich (1989), in an important work on tanaid fauna of the
northeast Atlantic, recorded 14 species, nine of which were new. They added descriptions of two
new genera {Mystriocentrus and Parapseudotanais) and six new species of Pseudotanais which
represented nearly a 20% increase in the number of taxa. Additionally, Bird and Holdich (1989)
reported some specimens of Cryptocopoides arcticus (Hansen, 1887) as having four pairs of
oostegites rather than the normal single pair, thus casting uncertainty on the placement of the
Cryptocopinae within the family Pseudotanaidae.
Statement of Problem
The primary objective of this study is to test the hypothesis that the current structure of
the family Pseudotanaidae should remain intact as a monophyletic group or, alternatively, should
be pared to exclude the members of the subfamily Cryptocopinae. Using conventional cladistic
4
methods, the strength of diagnostic characters u tat currently define relationships among members
of the two subfamilies Pseudotanainae and Cryptocopinae are assessed. Questions addressed
include: (1) are the currently accepted genera and subgenera of Pseudotanaidae monophyletic?
and (2) can evidence be provided through the analysis of morphological characters to justify
either further subdivision into additional monophyletic subgroups or eliminating some members
altogether? Secondary objectives to be obtained through this study are the production of uniform
diagnoses of all existing species and the description of new species discovered in material from
the Gulf of Mexico and the Atlantic east coast.
Significance of Study
Since the establishment of the family Pseudotanaidae in 1973, the taxonomic structure of
the family has increased in complexity along with a more than doubling of the number of species
assigned to it. In addition to this increased complexity, the question regarding the status of the
subfamily Cryptocopinae necessitates a revision of the family systematics. Cladistic methods
used in this research reveal evidence of a new working hypothesis of Pseudotanaidae phylogeny
which includes the removal of members of the Cryptocopinae and suggests new hierarchies
among both, the remaining genera and species of Psedotanaidae and the detached species of
Ciyptocopinae. The uniform morphological diagnoses of all known species of Pseudotanaidae
provides a concise basis of comparison for the addition of new species in the future. Finally, the
six new species from the Gulf of Mexico presented in this study represents merely a starting point
for further biogeographical investigations in the deep waters of this region; it is likely that at least
eight to ten additional species of Pseudotanaidae exist there.
5
CHAPTER II
REVIEW OF RELATED LITERATURE
Foundational literature
The family Pseudotanaidae was established by Jurgen Sieg in his 1973 doctoral
dissertation and formally recognized in his tanaid systematics publication in 1976. The
foundation of the family was based in part on a description of Pseudotanais by the Norwegian
zoologist G.O. Sars in an 1882 paper dealing with a revision of the crustacean order Isopoda back
when tanaids were considered a type of isopod, and on some species placed by Karl Lang in the
families Tanaidae (Lang 1949) and Paratanaidae (Lang 1967). At the time, all related species
belonged to the suborder Dikonophora (Lang 1956), which was based on the number of genital
cones in males (two vs. one).
Sieg's 1977 monograph was essentially a publication of the Pseudotanaidae portion of his
1973 dissertation in which he established two new subfamilies, the Cryptocopinae with four
genera (two new) and the Pseudotanainae containing the single genus Pseudotanais and nine new
species. One new genus from his dissertation, Mortensenia, later became Iungentitanais in the
1977 work after Sieg learned that the former was preoccupied by a genus of Echinoderm. In his
monograph, Sieg erected the subgenera Akanthinotanais and Pseudotanais sensu stricto
(indicated hereafter by 'A' and 'P' in parenthesis), to separate species of the Pseudotanainae
based on the presence or absence of carpal blade-like setae on pereopods 2 to 4. In all, Sieg
reviewed and described 21 species, either from his personal collections or from museum material
sent to him from collections made world wide. Complete synonymies, illustrations and
geodistribution data were provided for the following species: Iungentitanais primitivus Sieg,
1977; Paraiungentitanais longidigitatus (Kudinova-Pasternak, 1975); Cryptocope abbreviata
(G.O. Sars, 1866); Cryptocopoides artica (Hansen, 1887); Pseuodotanais {A) longipes Hansen,
1913; Pseudotanais (A) gerlachi Sieg, 1977; Pseudotanais {A) malayensis Sieg, 1977;
Pseudotanais (A) mortenseni Sieg, 1977; Pseudotanais (A) similis Sieg, 1977; Pseudotanais (A)
6
gaussi Vanhoffen, 1914; Pseudotanais (P) affinis Hansen, 1887; Pseudotanais (P) nordenskioldi
Sieg, 1977; Pseudotanais (P) longisetosus Sieg, 1977; Pseudotanais (P) oculatus Hansen, 1913;
Pseudotanais (P) liljeborgi G.O. Sars, 1882; Pseudotanais (P) macrocheles G.O. Sars, 1882;
Pseudotanais (P) unicus Sieg, 1977; Pseudotanais (P) abyssi Hansen, 1913; Pseudotanais (P)
jonesi Sieg, 1977 and Pseudotanais (P)forcipatus (Liljeborg, 1864). Sieg's monograph also
proposed a brief phylogeny of the genera and provided keys to the species and maps of their
world distribution. Two species, Pseudotanais borceai (Bacescu, 1960) and P. vitjazi (KudinovaPasternak, 1966) were listed by Sieg as incertae sedis primarily because he had no specimens of
them to study; the type material for P. borceai had been lost and the description of P. vitjazi was
confusing and incomplete. Surprisingly, P. vitjazi was apparently later recognized by Sieg since
it appears in his 1988 key to the known species of Pseudotanaidae (Sieg and Heard 1988). Their
questionable status notwithstanding, these two species apparently belong to the genus
Pseudotanais by virtue of their well-illustrated carpal blade-like setae and were thus included in
the present study.
The second most important contribution to the knowledge of Pseudotanaidae was by Bird
and Holdich (1989) in their work on the tanaidacean fauna of the northeast Atlantic Ocean in
which they summarized recent work on the subfamily Pseudotanainae in the region and provided
records of 14 species including two new genera and nine new species descriptions. At the time
this represented a substantial portion (40%) of the known species and their nine new species
increased the fauna of the family by 20%. In their discussion of family systematics they drew
attention to problems relating to the accepted taxonomy of the group such as the fact that of the
diagnostic characters used by Sieg (1977), none are in themselves restricted to the Pseudotanaidae
but can be found in other tanaidomorph families. For example, the principle defining character the single pair of oostegites on the fourth pereopod - is also seen in members of the family
Tanaidae, although in that family the oostegites are formed by an enclosed sac-like structure
("ovisac"), rather than the lamelliform plates seen in the Pseudotanaidae. Bird and Holdich
/
(1989) also found evidence that the single oostegite pair character may be limited only to the
subfamily Pseudotanainae since specimens of Cryptocopoides cf. arcticus were examined during
their study with four pairs of oostegites, thus putting into question the presence of the
Cryptocopinae within the family Pseudotanaidae. These authors also provided a dichotomous
key and distributional notes on 16 species encountered including detailed descriptions and
illustrations of the following new species: Mystriocentrus serratus, Parapseudotanais abyssalis,
Pseudotanais (P) scalpellum, Pseudotanais (P) spatula, Pseudotanais (P) longispinus,
Pseudotanais (P) denticulatus, Pseudotanais (P) spicatus, Pseudotanais (P) vulsella, and
Pseudotanais (P)falciculata. In addition, a re-description of Pseudotanais (P) affinis (Hansen,
1887) was provided and a new species, Pseudotanais (P) corollatus was established that resulted
from Sieg's (1977) mistakes in his re-description of P. affinis. Bird and Holdich (1989) proposed
two groups of related taxa - the "forcipatus" group composed of those species with forcipate
chelae, and the "affinis" group, a confusing assemblage of eight species supposedly united by
similarities in molar structure (note: P. corrolatus and P. denticulatus both have broad, truncate
molar structures, whereas in the others the molar structure is narrow and pointed).
Geo-distribution.
The family Pseudotanaidae has been reported from all the world's oceans, but nearly half
of the described species (18) are known from the northern Atlantic and Arctic regions probably
because more studies have been conducted in these regions (Bird and Holdich 1989). Sieg (1977)
incorporated in his monograph (see above) species collected by himself and others including
Hansen (1887, 1913), Liljeborg (1864) and G.O. Sars (1866, 1882). Kudinova-Pasternak (1985)
studied tanaids from collections made on the Great Meteor Seamount in the North Atlantic. From
those taken near the summit (325 - 470 m), she described the new species, Pseudotanais (A)
siegi, an oculate species noted for its long spindly appendages and apparently (based on the
illustration) with slight serrations on the chela incisive margins. Like other works from this
author, the description was incomplete and illustrations were inadequate.
8
From the Black Sea and Mediterranean Sea, there have been described one and two
species respectively. Pseudotanais (P) borcaei, an oculaie species collected near the mouth of
the Black Sea in 60 - 70 m, was described by Bacescu (1960) and incorrectly assigned to a new
genus, Pontonais. Sieg (1977) recognized the species as belonging to Pseudotanais but treated it
as incertae sedis because of "inconsistencies" in the original description and the loss of the type
material (Sieg 1977, Sieg and Heard 1988). Sieg included two Mediterranean species in his 1977
monograph, both oculate species from relatively shallow waters off the coast of Naples, Italy.
One, Pseudotanais (P) unicus, was a new species featuring atypically short third pereopods, and
the other, Pseudotanais (P) mediterraneus, was described earlier by G.O. Sars in 1882.
Four nominal species of Pseudotanaidae are known from the West Indies and Gulf of
Mexico regions of the North Atlantic. In Sieg and Heard (1988), a key to 23 species considered
to be valid at that time was provided along with discussion of why other species were
disregarded. Also included were new illustrations of Iungentitanais primitivus (Sieg 1977) and
Pseudotanais (A) morlenseni (Sieg 1977), both collected from shallow water off the coast of
Florida, and a description of the new species Pseudotanais (P) mexikolpos from shallow water
(72 m) off the Texas coast. Not surprisingly, all three species are oculate. In his monograph of
deep water tanaids of the Gulf of Mexico, Larsen (2005) reviewed the known Gulf species of
Pseudotanaidae discussed by Sieg and Heard (1988) and mentioned that several unidentified
species were discovered that will be covered in the present study. The remaining pseudotanaid
known from the West Indies region, Pseudotanais (P) baresnauti, was taken from deep water
(935 m) near the Barbados Trench in the Caribbean Sea (Bird 1999).
Based on collections from Atlantic subantarctic waters off South America and off
Antarctica, Kudinova-Pasternak (1975) recorded four pseudotanaids: (1) Cryptocope antarctica,
later synonymized with Cryptocopoides arcticus by Sieg (1977); (2) Pseudotanais gaussi, which
was earlier described from a nearby location by Vanhoffen (1914); (3) Pseudotanais
nordenskioldi, which Sieg (1973) had earlier described and named in his doctoral dissertation;
9
and (4) the new species Mortensenia longidigitata, also taken from Sieg's dissertation and which
in his 1977 monograph, he placed in the new genus Paraiungentititanais.
In Antarctic waters of the Indian Ocean, Vanhoffen (1914) reported Pseudotanais abyssi,
a species previously found off Greenland, and described a new species, Pseudotanais ganssi from
samples collected by the German South Pole Expedition of 1901-1903. Sieg (1977) examined
Vanhoffen's specimens of P. abyssi and compared them to Hansen's (1913) specimens from off
Greenland (type locality) and apparently concluded that they were conspecific and considered
them to have a bi-polar distribution. Sieg (1977) described a new species, Pseudotanais (P)
longisetosus based on material from north of Cumberland Bay, South Georgia collected during
the Swedish South Pole expedition to Antarctica . The enigmatic species Pseudotanais {A) guillei
Shiino, 1978, was collected from the vicinity of Kerguelen Island off Antarctica. Shiino based
his description on a single specimen and did not dissect the mouthparts. Sieg (1986), in a study
of tanaids collected aboard the R/V Hero in 1970 in the sub Antarctic region, worked on
additional material of P. guillei and included a description and illustrations of the mandibles and
pars molaris. This oculate species, while superficially similar to P. mortenseni, is shown by the
phylogenetic analysis of the present study to have intrinsic apomorphic differences that set it
apart from all other pseudotanaids. In the same work, Sieg also reported additional occurrences
of P. abyssi, and P. nordenskioldi from Antarctic waters.
Pseudotanais (A) gerlachi, a shallow-water oculate species, was described by Sieg (1977)
from collections made near the Malediven, Addu Attoll in the northern Indian Ocean and is
known only from the type locality. In a study of tanaids collected from deep water (3923 - 5060
m) in the Madagascar Basin, Kudinova-Pasternak (1987) described two species belonging to new
genera, Latitanais beklemishevi and Curtichelia expressa, but apparently did not assign them to
any higher category. They later were placed in Pseudotanaidae, subfamily Cryptocopinae, as a
result of a phylogenetic analysis of the superfamily, Paratanaoidea conducted by Larsen and
Wilson (2002).
10
In the South Pacific Ocean, there are only two records of the family Pseudotanaidae.
Sieg (1977) included in his monograph a description of Pseudotcmais (A) malayensis, an oculate
species similar to P. gerlachi, from shallow water near the Gilbert Islands north of Australia.
More recently, Bamber (2005) included in his account of tanaids from Esperance, Western
Australia, a description of a new shallow water oculate species, Pseudotanais (A) scrappi, which
is similar in some ways to P. guillei but is unique in that the maxillule endites terminate with
seven spiniform setae rather than the usual nine known for other members of the genus.
In the northern Pacific Ocean, pseudotanaids have been reported along the coasts of
California and Alaska and in the northwestern abyssal regions. Dojiri and Sieg (1997) described
two species, Pseudotanais (P) californiensis and Pseudotanais (A) makrothrix from the Santa
Maria Basin off California. Both species were inadequately described and the illustrations are
incomplete. One (P. makrothrix) is an oculate while the other is not, even though they were both
collected from similar depths (90 to 390 m). Pseudotanais (P) falciculata, widespread in the
North Sea and Arctic Ocean, was recorded in the Barents Sea off the coast of Alaska by Stappers
(1911), as cited in Sieg (1977). Also from the Northeast Pacific, Pseudotanais (P) oculatus,
another species common in the North Sea, occurred in studies conducted off the coasts of
Washington by Hatch (1947) and Departure Bay, Northwest Canada by Fee (1926) - both cited in
Sieg (1977).
In the Northwest Pacific, pseudotanaids have been identified in samples collected from
the abyssal plain region and also in the Kurile-Kamchatka and Japan Trenches north of Japan.
Kudinova-Pasternak (1966) examined specimens collected from the abyssal plain south of the
Bering Sea in depths greater than 6,000 m and described the species Pseudotanais (P) vitjazi,
named for the Soviet research vessel, Vitjaz. Sieg (1977) considered P. vitjazi to be incertae
sedis mainly because material was unavailable for examination and the original description was
inadequate. Nevertheless, he included it in his key to the worldwide species (Sieg and Heard,
1988) as distinguished from P. affinis by the subequal length of the uropodal rami and distinct
11
denticulation of the pars molaris. In 1970, Kudinova-Pasternak reported on additional material of
P. vitjazi collected from 5400 m depth in the Kurile-Kamchatka Trench. She reported the
occurrence of Crytocope arctica Hansen, 1887 (= Cryptocopoides arcticus) along with an
unidentified Pseudotanais species and an unidentified male Cryptocope, which she partially
illustrated. More recently, McLelland (2007) examined pseudotanaid specimens collected from
the Kurile-Kamchatka and Japan Trenches by the Japanese research vessel, Hakuho Mara and
reported the occurrence of two new species, Pseudotanais (P) nipponicus and Cryptocopoides
pacificus. He considered it probable that Kudinova-Pasternak's record of Cryptocopoides
arcticus from the Kurile-Kamchatka Trench actually refers to C. pacificus.
Oculate species and vertical distribution
Most pseudotanaid species are blind and occur at depths ranging from 1,000 to 6,050 m;
however, some occupy shallow water habitats from near surface to near the edge of the photic
zone (0 - 300 m). All of the 11 known oculate species (those with either fully functional eyes or
eyelobes with missing visual elements) are from depths shallower than 500 m. Pseudotanais (P)
siegi is the deepest recorded oculate species at 325-470 m in the North Atlantic; the rest are
within the photic zone with most inhabiting waters of less than 100 m. In contrast, the absence of
eyes does not invariantly predict the depth of occurrence as six species without eyes are known
from photic habitats in less than 100 m. For example the non-oculate P. similis and P.jonesi
were collected from the English Channel in 20 and 50 m respectively (Sieg 1977). It is worth
noting that seven of the ten species of the more apomorphic subgenus Akanthinotanais (70%)
have pigmented eyes, which goes along with their shallow water habitat. This gives credence to
Bird and Holdich's (1989) assertion that "this genus originated in shallow water and is currently
invading the deep-sea".
12
CHAPTER III
METHODOLOGY
Material Studied
Data used in the analysis for this research was taken largely from published records on all
known species from the family Pseudotanaidae including two new species described by this
author from the North Pacific (McLelland 2007). Additionally, data was used from six new
species discovered in pre-sorted material from the northern Gulf of Mexico (Appendix A) and
also from a new species discovered in material collected from the South Atlantic Bight off the
U.S. east coast. Descriptions of these new species appear in chapter V. Unless otherwise
indicated, morphological terminology used in this research follows that of Bird and Holdich
(1989) and Larsen (2003).
Laboratory Techniques
Pseudotanaidae specimens were examined using a Wild M5 stereomicroscope and a Leitz
Labrolux 11 compound microscope. Dissections were performed using chemically sharpened
tungsten wire needles on specimens softened with glycerol. Measurements and sketches of whole
animals and dissected appendages were made using an Olympus 1.25X drawing tube attached to
the compound microscope. After illustration, dissected appendages were transferred to glycerin
jelly for permanent storage. Upon formal description, specimens selected as type material will be
deposited in the Smithsonian National Museum of Natural History with additional material placed
in the Gulf Coast Research Laboratory Museum.
Data Management and Character Selection
Meristic data was derived from original descriptions of all known species from the family
Pseudotanaidae, including those discovered during this research, and species used as out-groups
in the phlyogenetic analysis. Measurements to the nearest 0.1 mm were made on enlarged
illustrations of the holotype and dissected paratypes to generate values including body length
percentages of the antennule, cephalothorax, pereon, pleon, pleotelson, and length vs. breadth
13
dimensions of articles of the antennae, pereopods, pieopods, and uropods. Text descriptions from
the literature were used to supplement, and in some cases to ascertain, certain character states
such as simple vs. spiniform setae when original illustrations were poorly rendered. Meristic data
and character descriptions were entered into a DELTA database in order to generate a concise
uniform text description for each species and a nexus data matrix for use in the phylogenetic
analyses. The DELTA (Descriptive Language for Taxonomy) format, designed by Dallwitz et al.
(1997), has been adopted as a standard for data exchange by the International Taxonomic
Databases Working Group. In generating the nexus file, characters were selected that not only
provided diagnostic value in defining a taxon but were considered to be reliable in seeking
monophyletic groups defined by shared, derived character states based on comparison with outgroup polarity, thus defining phylogenetic relationships among the various species being
analyzed. In all, 69 characters were selected that were considered to be unambiguous and that
had diagnostic value at the suborder, family, subfamily, generic, subgeneric and specific levels.
Some characters useful for morphological description such as specific numbers or type of setae or
highly subjective descriptions of body features (e.g., shape of the cephalothorax being "subtrapezoidal") were often too variable or ambiguous and thus were not scored for use in the
phylogenetic analysis.
Data Analysis
Out-groups.
Carpoapseudesprospectnes, a species from a distant apseudomorph family, and three
representatives from more closely allied tanaidomorph families, were selected as out-group taxa
to establish polarity of the selected character states and to increase the chances of identifying
plesiomorphies (ancestral characters). Three intermediately distant taxa (Typholtanais parvus,
Hargeria rapax and Sinelobus stanfordi) were included to strengthen the definition of the ingroup
node, since more closely related taxa increase the number of characters that can be compared
(Nixon and Carpenter 1993), thus assisting the construction of a more robust phylogeny among
14
the established Pseudotanaidae taxa. Two of the out-group taxa, C. prospectnes and S. stanfordi
provided historic perspective in that they were used as out-groups in Larsen and Wilson's (2002)
phyiogenetic analysis of the superfamily Paratanaoidea representing respectively the suborder
Apseudomorpha and the superfamily Tanaioidea.
In-group.
The in-group consisted of 53 species comprising all known species worldwide of the
family Pseudotanaidae plus those newly discovered from this research. Currently the family
consists of the subfamilies Pseudotanainae with 45 and Cryptocopinae with eight species. The
former is comprised of three genera, Mystriocentrus, Parapseudotanais and Pseudotanais.
Pseudotanais, the largest of the three genera, is comprised of two subgenera, Akanthinotanais and
Pseudotanais sensu stricto, which include 10 and 31 species respectively. The Cryptocopinae is
more loosely defined with its eight species distributed among six genera, five of which are
monospecific.
Phyiogenetic analysis.
A heuristic search was conducted in PAUP* 4.0b (Swofford 2002) on unordered and
unweighted morphological characters using a generalized parsimony optimal ity criterion to find
the most parsimonious trees. Stepwise random addition on 100 replicates starting from random
trees with a TBR (tree bisection and reconnection) branch swapping algorithm was employed in
two steps as described by Olmstead et al. (1993) - first without saving multiple trees and again
with multiple trees saved but starting with trees stored in memory from the first step. In addition,
the character state optimization ACCTRAN (Accelerated transformation) was in effect which
puts the character change as close as possible to the point of divergence between the out-groups
and in-group, and MAXTREES was set at 100,000 trees with multiple trees saved. Each
replicate search was continued until 8 x 106 rearrangements were tried. A strict consensus tree
was computed from the most parsimonious trees resulting from the heuristic search described
above and trees resulting from both the heuristic search and computed consensus were saved.
15
Cladograms were generated using the display Print Trees option in PAUP and the resulting PICT
files were modified for display in this research using Photoshop 7 (Adobe Systems, Inc.). Since
more than one out-group was used, out-groups were set to display as paraphyletic to the in-group.
Various tree statistics were computed to indicate how well the character data fit the tree
and the extent to which homoplasy (character state similarity not resulting from shared ancestry)
permeates the tree topology. These included tree length (TL) which is the number of steps
required to achieve the shortest, most parsimonious tree, consistency index (CI), retention index
(RI), rescaled consistency index (RC), and homoplasy index (HI); the latter four being indicators
of the amount of homoplasy present in the tree and how well the tree describes the data set. The
CI, a ratio of the minimal amount to actual amount of character change in the tree, is sensitive to
uninformative characters (autapomorphies and symplesiomorphies) while the RI indicates
character similarities, regardless of the autapomorphies and symplesiomorphies present, by
measuring the amount of synapomorphy (shared derived characters) in the data set based on
homoplasy percentages (actual/maximum possible). The RC is a function of the CI multiplied by
the RI and approaches one as homoplasy decreases. The HJ is merely the reciprocal of CI and
approaches zero with decreasing homoplasy.
In the initial character selection process, CI values were used to assess characters for
possible inclusion in or exclusion from the analysis. Character diagnostics resulting from a
search conducted on an initial 80-character data set revealed some characters with extremely low
(less than 0.08) CI values. These were excluded to arrive at the 69-character array used for
analysis. A diagnostic description of the most parsimonious tree including statistics for each of
the 69 characters used in this analysis is presented in Appendix B.
Optimal trees were evaluated for support using (1) jackknife resampling, which
systematically removes taxa out of resampled data to detect the percent occurrence of a particular
clade, thereby determining how much the tree topology is reliant on a single taxon, (2) bootstrap
analysis which repeatedly re-samples the data set until a comparably sized pseudo-replicate data
16
set can be obtained from which a frequency measurement of the of re-created individual branches
is obtained, and (3) the calculation of decay indices to estimate branch support on the strict
consensus of all trees resulting from parsimony analysis. Also known as "Bremer support"
(Bremer 1994), decay indices indicate how strongly the data supports a particular hypothesis on
the tree based on the number of steps required for the branch or clade to collapse. Jackknife and
Bootstrap analyses were performed as PAUP functions and decay indices were obtained using
TreeRot v.2 (Sorenson 1999). As compared to their use in analyzing molecular data, Jackknife
and bootstrap analyses have been criticized as deficient for analyzing morphological data sets
because they have relatively small numbers of dependently distributed characters (Felsenstein
1985). Data sets with such dependent characters often display erroneously high confidence
estimates with underestimated high and overestimated low values (Hillis and Bull 1993; Kluge
and Wolf 1993; Zharkikh and Li 1995). Notwithstanding, these analyses were employed in this
research to primarily examine the node separating the Cryptocopinae from the remainder of the
family, thus providing a modicum of supplementary support to the more definitive decay indices.
MacClade 4.06 (Maddison and Maddison 2005) was used to trace individual character
evolution among clades observed in a strict consensus of the most parsimonious trees generated
by PAUP. Diagnostic characters delineating subfamilies, genera and subgenera were examined,
and in some cases detected, and marginal, low-consistent characters were evaluated for possible
exclusion from further analysis.
Re-weighted character analysis
Parsimony analyses using differentially weighted characters were also conducted using
PAUP, in which characters were re-weighted based on the highest RC, CI or RI for all equally
parsimonious trees from the original heuristic search. The value of character weighting in
cladistic analysis has been discussed by many authors including Farris (1969, 1982), Carpenter
(1988), and summarized by Goloboff (1993). The objective of these re-weighted analyses was to
test the effect, if any, of minimizing systematic error by maximizing informative data, assuming
17
all characters are neither equally informative nor equally reliable. Those trees which imply the
characters to be more reliable explain the data better (Goloboff 1993). For these analyses all
settings were as in the original, unweighted analysis. Best trees from the re-weighted analyses
were added to a file with the best unweighted tree and the Kishino-Hasegawa (K-H) maximum
likelihood test was used to determine if any of these were significantly different from the original
unweighted set. The K-H test is a parametric test that compares tree lenths between the optimal
tree and alternate trees under consideration by generating paired t-values.
Marsitpium development
Bird and Holdich (1989) reported some specimens of Cryptocopoides arcticus as having
four pairs of oostegites rather than the normal single pair as prescribed for the family, thus casting
doubt on the validity of that species and thus the systematics of the Cryptocopinae as belonging to
Pseudotanaidae. To evaluate the strength of marsupium development as a diagnostic character
for the phylogeny of Pseudotanaidae, an additional heuristic search was conducted using the same
setup as before except for the exclusion of the character for marsupium development. Consensus
trees obtained from this second data set were tested for support using jackknife, bootstrap and
decay indices as before. In addition, best trees from each dataset were placed in a "neutral" file
with a number of characters different from each dataset and a Kishino-Hasegawa test was
conducted to test for significant difference.
18
CHAPTER IV
RESULTS OF DATA ANALYSIS
Parsimony analyses
General Observations.
The heuristic search conducted on 53 species of pseudotanaids and four out-group species
using 69 characters resulted in six equally parsimonious trees with a 'best' tree of length 627
(Table 1). Although a rather high degree of homoplasy was evident (HI = 0.745), a consensus of
the trees (Fig. 1) clearly indicated a separation of the taxa associated with the subfamily
Cryptocopinae from the remaining members of the family. This separation was supported by a
decay index of 4 on the node between Cryptocope abreviata and Pseudotanais (Akanthinotanais)
guillei. The nodes on either side of this separation had decay indices of 2 or less. Further
evidence for this separation was manifested in the jackknife and bootstrap analyses which
produced nearly identical consensus trees (Figs. 2 and 3) with values at this same node of 50 and
47 % respectively, values which were about 4.0 times higher than those of the nodes to either
side.
The ten species of the subgenus Akanthinotanais were positioned toward the ancestral
end of the tree topology and, although showing poor Bremmer support, appeared as paraphyletic
in the consensus of six trees (Fig. 1) with one, Pseudotanais (A) guillei, removed not only from
the clade containing the other nine but outside the clade containing the remainder of the family.
This subgenus grouping broke down completely in the jackknife and bootstrap analysis (Figs. 2
and 3).
The 35 species of Pseudotanainae were loosely grouped in two clades of 19 and 16
species, designated as A and B in Figure 1 and in the figures of Appendix D. Within these
structures, clade A consisted solely of 19 Pseudotanais species whereas clade B contained 12
Pseudotanais along with the strongly supported genera Mystriocentrus and Parapseudotanais,
each clade with a decay index of 4. Clades A and B, though poorly supported statistically, did
19
Table 1
Scores of most parsimonious trees from heuristic searches on datasets with (69-character) and
without (68-character) the character for marsupium development.
adjusted scores
69-char
68-char
69-char
68-char
TL
627
625
815
819
CI
0.255
0.253
.218
.217
Rl
0.521
0.520
.479
.466
RC
0.133
0.131
.103
.101
HI
0.745
0.747
.782
.783
Tree
TL
diff
69-char
815
best
68-char
819
4
0.5419
0.5894
K-H test:
s.d. (diff)
7.138113
* Probability of getting a more extreme T-value under the null hypothesis of no difference between the two trees (two-tailed test). Asterisked values in
table (if any) indicate significant difference at P < 0.05. Note: The Kishino-Hasegawa test was conducted on adjusted scores from best trees after each
was placed in a neutral 80-character file.
reveal some trends of commonality among their respective species based on shared characters.
These trends are examined in detail in Appendix D, which presents a synopsis of each character
used in the analysis along with illustrated character state tracings for informative characters.
Re-weighted character analysis
Heuristic searches were conducted on the data using three re-weighting methods and the
results were compared to the unweighted analysis. The topology of all trees resulting from both
unweighted and re-weighted anayses were highly variable among the more apomorphic
Pseudotanainae species but showed stability at the ancestral end of the tree containing the
20
outgroups and the Cryptocopinae. Unweighted tree scores were sHghtly better (lower TL and
higher CI) than all of the weighted trees (Table 2) indicating that, with the current character set,
the phylogeny is best described using unweighted data. A Kishino-Hasegawa test comparing the
best unweighted tree with trees re-weighted on RC, CI, and RI revealed no significant differences
at the P < 0.05 level of significance. Based on the above considerations, it was decided to center
the hypothesis on the original unweighted data set.
Table 2
Score comparison of unweighted and re-weighted trees
Tree:
unweighted
RC-weighted
Cl-weighted
Rl-weighted
TL:
695
717
705
697
CI:
.239
.232
.235
.238
RI:
.501
.481
492
.500
RC:
.120
.111
.116
.119
HI:
.761
.768
.765
.762
Kishino-Haseg;iwa Tesl
s.d (diff)
p*
t
Tree
Length
Length diff
unweighted
695
(best)
RC-weighted
717
21
15.95
1.3165
0.1922
Cl-weighted
705
9
11.12
0.8096
0.4208
Rl-weighted
697
1
7.73
0.1293
0.8975
Result: there is no significant difference among the trees at the P < 0.05 level
Marsupium development.
An additional heuristic search was conducted to test the effect on the data set of
excluding the character for marsupium development (one pair of oostegites vs. four pairs), a
major diagnostic feature of the Pseudotanaidae. The analysis on the remaining 68 characters
produced six equally parsimonious trees with a slightly shorter tree length (625) but otherwise
21
similar tree scores (Table 1). Bremmer support using TreeRot on the consensus of these trees
was nearly identical to the 69 character analysis with a decay value of 4 at the node between the
Pseudotanainae and the more ancestral species, including the Cryptocopinae (Fig. 4). Jackknife
and Bootstrap analyses likewise demonstrated support with values of 46 and 47 % respectively
although the Pseudotanainae clades collapsed similar to that seen in the 69 character analyses.
Best trees from each dataset were placed in an 80-character file to neutralize the effect of scores
being "adjusted" when they are placed in a file altered by different numbers of characters. A
Kishino-Hasegawa test revealed no significant difference between the 69 character and 68
character trees (Table 1).
22
Pseudoianais
(P)spatula
Pseudoianais (P)sp. O
Pseudoianais (P)sp. P
Pseudoianais (P)affinis
Pseudoianais
(P)scalpsHum
Pseudoianais (P) longispinus
Pseudoianais
(P)denticulatus
Pseudoianais
(P}sphatus
Pseudoianais (P) vitjazi
Pseudoianais (P) nipponhus
Pseudoianais (P) maciocneles
Pseudoianais (P)sp. C
Pseudoianais (P) kmgiseiosus
Pseudoianais (P)coroBatus
Pseudoianais (P)ocuiatus
Pseudoianais (P) mexihotpos
Pseudoianais
(P}boiceai
Pseudoianais (P) tinicus
Pseudoianais (P) tnedibenaneus
Pseudoianais (P) inflatus
Pseudoianais (P)sp. A
Pseudoianais (P) vulsella
Pseudoianais (P) fatcicuiata
Mysiriocentrus senatus
Mystriocentms sp. A
Pseudoianais
(P)abyssi
Pseudoianais (P) bamsnauti
Pseudoianais (P)calik>miensis
Pseudoianais (P) fonzipaius
Pseudoianais (P)jonesi
Pseudoianais
(P}cohnus
Pseudoianais (P) fflljebofgi
Pseudoianais (P} noidenshhtdi
I
Parapseudotanais
abyssatis
'
Paiapseudoianais sp. A
Pseudoianas (A) fcwgipes^Pseudoianais
(A)siegi
Pseudoianais (A) mahmthnx
Pseudoianais (A) mortenseni
Pseudoianais (A)geriachi
Pseudoianais
(A)gaussi
Pseudoianais (A)similis
Pseudoianais (A) malayensis
Pseudoianais
(A)sciappi
— Pseudoianais (A)guiHei ^ M
- Crytncope abre>/mia
Gr/pkxMpoxies »it:fau«!i
— Gtyptttcopokitts
pacifism
— liiytocopoKles sp. A
— lungcntitanais primibvus
— Par7wmQP.ntikwiaK kmgidigitrvitK
— leitikwiFNS beMemishcvi
— Cnriit:halitif.xpifissn
—ftinelobus stanfordi
Hatseha lapax
*
parvus
- Caipoapseudes
prospecines
Figure 1. Strict consensus of six equally parsimonious trees of the family Pseudotanaidae. Decay
indices calculated from TreeRot v.2. appear at nodes with values of 1 or less omitted. Arrow
indicates node with a decay index of 4 that separates Cryptocopinae (shaded) from remainder of
the family. Bracket indicates species of the subgenus Akanthinotanais.
32 |
5
45 |
1
27
1
|
20
0
0
9
23
,
16
0
7
1
21
4
4
19,
24 |
7
24 |
8
12
A
4
I
11
in
9
6
|
|
26
0
i
73
22
6
25
76
MyrJnocWbuv «#> A
Psewteten**. (P) (ittefctilsia
Ps8«*si6n*s (P) w»efta
Pstuastwuas iPJ abys«
P»u**»n»i<f (P) b*r«ti»iub
Pi»^M4**ls^J^fl^ (P) £«N**3**^*ttH<*
pM»s*»l»8iitf» (PJ (arctpato
Psvutfotanaiii (P) COIMM*
f>5©i*<iotii«ssi<; (P) j w i w
PKOlHMIafMM* (PJ illj<*«9l
PMnudetoniN't (PJ l*f«fi«*tn«i>
fjrfHHtotolMlh (P) NPP6MCU4
PM»udatenM<> (P) veolpoluFn
nrnxMimmu (Pj tftoMs
P»ti*fan»is (PJ sp O
PswiSwhm*'- <P) sp P
P&AutMatmii (P) nwcroctistes
PseuaotMiMi. ( p j s.p C
PS8«S«1WIS)*. (PS ftfttfW
f*s»udoisnMii (pj intf«t»
Ps«u4«tarMM» (PJ hp A
35
15
25
25
|
TO
3
50
64 |
13
1)
VaB>
|
P-ietrfstanai"? (A) jjerlaeN
PssuiJetarare if 1 ) nwJttenariews
p<s»t>stet»rt*'. (A) rtfsalayeirei!,
PwudotariMt. (A) i t t ^ ^ t i
Ps#uttolwi«i (P) bwmut
Ps«u*>tafwi<s (P) uncus
P-,mt$tstmmi< (A) puttof
S*umiMmm*» j p j msuawtpes*
P«»<ttiflal»sa<» jp) km^&Ma'Mi,
fHWMkHWW* (P) IKK«««Kt»eW»
f^wKMaruMti (P) bmtttptr',
Pseu<W**w« (PJ uwp
Ps»u**»ni*'. (A) nwirtwwww
PMHitfototMifc (A| fira&wlhtw
HMHidobmaiti (A) <p«KM
PbMHtolansi*! (A) SMKtn
PseudeteHBiB (PJ derAeolMui
P>>fMi*at«»».*i«, (P) spread
PseiKWanw. (P) vtli«t
Pse«**art«* (PJ coreltaiK»5>
P<!«uaa*»twii» j p j aetjisftj';
PurapwwWsirwMS. ativv»s»h;,
Psrapwudotaiwrt vp A
49 |
89 |
52 |
Sunc|«f?l»te*»i!-. iMinsttmii*
P(»»ijfnjei*)(tlim*i NHitfdHjilahA
^••Vpsncoi'wsfc...sp. A
Smelobus starsfofds
Hsrgena rapan
Tyiahtatwats p»rvw
Figure 2. Jackknife consensus of six equally parsimonious trees of the family
Pseudotanaidae. Arrow indicates node separating the species of Cryptocopinae
(shaded) from the Pseudotanainae. Values indicate percent occurrence of a particular
clade upon completion of the jackknife analysis.
24
s (P) longispiius
s (P) mpponious
is (P) scalpelum
is (P) spatula
(P)sp.O
s(P)sp.P
s (P) macrocheles
s<P)sp.C
is (P) aflnis
is (P) denticulatus
is (P) spicalus
is (P) tijazi
is (P) inlatus
s(P)sp.A
is (P) longisetoais
is (P) nordenshioMii
is (P) calHwniensis
s (P)ton:j>atus
s(P)janea
I
'
Mjsbiocennus seiratus
Mjstriocentius sp. A
s (P) lafciculata
is <P) wdsella
s (P) abyss
is (P) bareaiauti
s (P) colonus
s (P) lijeboitf
is (A) gerlachi
s (P) medHeiraneus
is (A) malayenss
is (A) sciappi
s (P) borceai
is (P) uncus
(A guild
Pseudotanai;s (P) meakolpos
Pseudotanai!a (P) coroUatus
Pseudotanai;is (P) oculahis
Pseudstanai!is (A) longipes
Pseudotanai;s (A) aegi
Pseudotanai;is (A) mortenseni
Pseudotanai!Is (A) rnakrothrbc
Pseudotanai!s (A) gausa
Pseudotanai!is (A) smiis
PaiapseudotanaE abyssals
Paiapseudotanais sp. A
Curbchelia expressa
beklemishev
kngentbnais primliws
PaiaiungenttanaE longidlgitalus
Crjpnwopoides pacihcus
Cryptoeopoides sp. A
Cryptocopoides anticus
Crjptocope abretiata
Sinelobus stanttnii
Hargeria rapax
TjphlotanaE panus
Caipoapseudes prospecbies
Figure 3. Bootstrap consensus of six equally parsimonious trees of the
family Pseudotanaidae. Arrow indicates node separating the species of
Cryptocopinae (shaded) from the Pseudotanainae. Values indicate
frequency of re-created individual branches at the completion of the
bootstrap analysis.
25
Pseydtonais (P) Inflate
PMNKtotwiato (P) sp, A
Ps««<J(5tanais (P) denticulatus
Pseudotanais (P) spieatus
Pseudotanais (P) vitjazi
Pseudotanais (P) affinis
Pseudotanais (P) seafpelum
Pseudotanais (P) spatula
Pseudotanais (P) longispinus
Pseudotanais (P) nippanteus
Psewlotiwste (P) sp. O
Pseudotanais (P) sp, P
Pseudotanais (P) macrocheles
Pseaetotanais (P) sp. C
PsBodotanais (P) corallatus
Pseudotanais (P) ocutatus
Pseudotanais (P) lortgisetosus
Pseudotanais (P) caJiforniensts
Pseudotanais (P) forcipatus
Pseudotanais (P) jonesi
Pseudotanais (P) cotonus
Psetidotanais (P) IMjeborgi
Pseudotanais (P) baresnauti
Mystriocentnis setratus
Mystriocentrus sp, A
Pseudotanais (P) falciculaia
Parapseudotanais abyssaiis
Parapseudotanais sp, A
Pseudotanais (P) abyssi
Pseudotanais (P) nofdenskioidii
Pseudotanais (P) vulsella
Pseudotanais (P) borqeai
Pseudotanais (P) unicus
Pseudotanais (P) mediteiraneus
Psetrtotanais (P) rnexiiselpos
Pseudotanais (A) gaussi *
Pseudotanais (A) similis
Pseudotanais (A) lortgJpes
Pseudotanais (A) sisjji
Pseudotanais (A) malayensls
Pseudotanais (A) scrappi
Pseudotanais (A) gedaehi
Pseudotanais (A) makrothrix
Pseudotanais (A) mortwisersi
Pseudotanais (A) guillal mm
CryntoRooe abreviata
Crypiooaaoides arcfisjs
Cryatoccooides pacif cus
Ciyplocasoides so. A
Ij-gentitanais prmil uus
Paraiungerlitanais longidigilatjs
Laatanais be«lemishev
Curtichelia expressa
Sineiootis Stanford!
Hargeria rapax
TypMotanais parvus
Carpoapseudes prospoctnes
Figure 4. Strict consensus of six equally parsimonious trees of the family Pseudotanaidae with
the character for marsupium development omitted from the analysis. Decay indices calculated
from TreeRot v.2. appear at nodes with values of 1 or less omitted. Arrow indicates node with a
decay index of 4 that separates Cryptocopinae (shaded) from remainder of the family. Species
bracketed are those of the subgenus Akanthinotanais.
26
Summary of Results
1. The results of a cladistic analysis using 69 morphological characters on 53 species of
Pseudotanaidae and four out-groups credibly implies that the group of species currently belonging
to the subfamily Cryptocopinae are separate from those of the subfamily Pseudotanainae. The
main evidence for this conclusion is the strong support indices of decay (Fig. 1), jackknife (Fig.
2) and bootstrap (Fig. 3) at the node where the Pseudotanainae diverges.
2. A separate analysis omitting the character for marsupium development resulted in a
tree not significantly different in character diagnostics (Table 1) and with similar support values
at the separating node for the two subfamilies (Fig. 4). This implies that, although some species
of Cryptocopinae may possess more than one pair of oostegites as suggested by Bird and Holdich
(1989), the absence of this character does not alter the hypothesis that the two subfamilies are
separate.
3. The cladogram produced by this research suggests the existence of three assemblages
within the Pseudotanainae which, although lacking in support, are typified by particular suites of
characters, discussed in the previous section, none of which are unique to either clade. These are
indicated in Fig. 1 as clades A and B and with a bracket for the species of the subgenus
Akanthinotanais, and also in illustrated character tracings presented in Appendix D (Figs. 33 92).
4. Clade A, consisting solely of 19 species of the subgenus Pseudotanais sensu stricto,
was characterized by twelve species with short, stout spiniform setae on antenna articles 2 and 3
(character 13 - Fig. 43) and maxilliped endites fused up to 2/3 their length (character 24 - Fig.
50), 14 species having a propodus length exceeding 6.1 times its width (character 42 - Fig. 67),
13 species with long carpal blade-like setae, exceeding half the length of the propodus (character
52 - Fig. 77), and seven species with additional carpal specialized setae (character 60 - Fig. 84).
5. Clade B containing 16 species comprised a mixture of three genera including
Pseudotanais sensu stricto, Mystriocentrus and Parapseudotanais. The clade was typified by
27
eleven species with more derived maxilliped endites which were nearly or completely fused
(character 24 - Fig. 50), seven species with strongly forcipate chelae (character 26 - Fig. 52), and
14 with short carpal blade-like setae of less than half the propodus length (character 52 - Fig. 77).
6. The subgenus Akanthinotanais, consisting of 10 species (bracketed in Fig. 1), is
paraphyletic with one species, Pseudotanais (A) gui/lei, lying outside a clade containing the other
nine. This assemblage is characterized by seven species with eyes (character 2 - Fig. 34), nine
with a simple pointed or whip-like pars molaris on the mandibles (character 18 - Fig. 47), seven
with the maxilliped endites nearly or completely fused (character 24 - Fig. 50), eight with an
apomorphic shortened cheliped propodus of less than or equal to 1.5 times the carpus length
(character 30 - Fig. 56), seven with a dactylus distinctly shorter than the propodus on pereopod 1
(character 45 - Fig. 71), and all with the apomorphic lack of blade-like carpal setae on pereopods
2 to 6 (characters 51 and 59 - Figs. 76 and 83).
7. The eight species of the subfamily Cryptocopinae formed a loosely organized,
paraphyletic assemblage that contained one monophyletic group composed of the monospecific
genera, Curtichelia expressa, Iungentitanais primitivus, Paraiungentitanais longidigitatus, and
Latitanais beklemishevi with fairly good support values (clade C in Figs. 39, 44, and 87). As a
whole, the eight-species assemblage was distinguished by all lacking abbreviated pereonites
(character 3 - Fig. 35), seven with a short first antennular article (character 6 and 7 - Figs. 37 and
38), seven with the apomorphic broad and truncate mandible pars molaris terminating with a ring
of setae or short denticles (characters 17 and 18 - Figs. 46 and 47), all having completely separate
maxilliped endites (character 24 - Fig.50), all with a short and five with a broad cheliped
propodus (characters 29 and 30 — Figs. 55 and 56), seven with 2 inferior setae on the cheliped
fixed finger (character 31 - Fig. 57), seven with two or three propodus setae on pereopod 1
(character 43 - Fig. 68), five with a single merus seta on pereopods 2 and 3 (character 49 - Fig.
74), all lacking the dactylus-unguis fusion on pereopods 4 to 6 (character 62 - fig. 86), and as
with the Akanthinotanais species, all with the apomorphic lack of blade-like carpal setae on
28
pereopods 2 to 6 (characters 51 and 59 - Figs. 76 and 83). The aforementioned four-species
clade within the Cryptocopinae was distinguished from others of the subfamily by having a
longer antennular third aricle (character 8 - Fig. 39) and well-developed pleopods with terminal
and lateral setae (character 63 - Fig. 87).
29
CHAPTER V
SYSTEMATICS AND TAXONOMY
Proposed Systematic Revision
Based on an interpretation of the cladistic analysis presented in chapter IV, a systematic
revision of the current family Pseudotanaidae is proposed in this chapter in which a new family
and four new subfamilies are created. An illustration of this new phylogeny indicating the
position of the subfamilies is presented in Figure 5. Complete diagnoses of all new taxa resulting
from this proposal are presented in the following section, including full descriptions for new
species discovered during this research. All descriptions and diagnoses are based on females
since males for this group remain largely unknown. Keys to the major groups appear within each
sub-level and a key to all species is given at the end of this chapter.
As seen in the consensus of six most parsimonius trees (Fig. 1), and in supplementary
jackknife and bootstrap analyses (Figs. 2 and 3), there is strong evidence for the removal of eight
species currently assigned to the subfamily Crytocopinae from the family Pseudotanaidae. It is
hereby proposed to erect a new family, Cryptocopidae to contain these eight species. Within this
family, substantial (though weaker) support is also evident to erect three subfamilies:
Cryptocopinae (one species), Cryptocopoidinae (three species), and Iungentitanainae (four
species). Cryptocopinae, originally designated by Sieg (1977), contains the genus Cryptocope
and diverges from the node separating it from the others of the family and from the
Pseudotanaidae with a decay value of 2 and bootstrap and jackknife indices of 13 and 11
respectively. Cryptocopoidinae, containing the three species of Cryptocopoides, is supported
with jackknife and bootstrap values of 24 each (Figs. 2 and 3) while Iungentitanainae, containing
the four monospecific genera, are supported by bootstrap and jackknife indices of 21 each and
with a decay value of 2.
The revised Pseudotanaidae with its large number of species presented a weakly
supported structure in the cladistic analysis probably owing to the large amount of homoplasy
30
Pp«;u*i(.tatt<<is
tF)spalutSmmmm'
Pseddotaoais (P)sp. P
Pseudetanate (P)B8atpelftim
Pseudoianais (P) tongfspinits
P?ewdeteisa& (P) de«M6Mlafe5
Pg«(i«J9taftaj$ (pjsplciUsi
Psfeudfttonaig (P)vlliiat
PsfjudotaMts (P) nippttftifcus
Ps guttata it*J$ (P) ffi.i(5r(K!h*!es
PMtHtataMif (P)«|i<C
Psftwdotenais (P) tanjjfeetasus
PseudotoitaSs (P) c«roJ(atns
PseadefaRais (P) ecalaius
Pse«(totenats (P) mexikoljjfts
Psfeudotanafc (P)borceai
Pseuttatanats (P) unicus
Ps guttata nate (P) meditenosetss
Psfeudotanais (P) inflates
PSftwdotoiais (P)sp.A
Pst>u(J6liJ*ais (P) vulsella
Psfc«<tatajMis ^P)f*t)S!6«ldta
PaeudotonaisCM) semi t us
Ps«udoianais(M)^, A
PifgyitatanaiiS (p)flfepg)
PsSMdttiaitajs CP) fcaftssnaiift
Pseaftataaais (P) eaWattiiensis
Pstsudotanats (P) f*reipatus
PssaitatatiaSs (P)j*ne£j
Psfcudotauais (P) e»l<w«s
Pseudotaitais (P) IHIjebarji
Pseudotabes (P) ftodeoskieldji.
Paiapseudotanais abyssal is
Paiapseudotafiais sp, A
Akanihiitetanais(A) longipfl a> "
Afcanthtaotanais (A) s iegi
Akjnthinotanais(A) m«kl>ttilfte
Afcanthittata rials (A) mfcrtens«ni
Akanthittotanais (A) g ftrisehi
Akanthiftotanais (A) § awss i
AJkanthiftfttanais (A) S im His
Akanthinotanais (A) m »lay ins is
f~"
1
,
1
tf
•
•
•
.
.
Afcahthtaotanais (A) s aiappi
Akanthinoianais (CO gyiilei
CffM* <*«(*« abrwiabC^
Ciftaeoptiides arcticus
Ciytacapaides paeifisus
Ciyt>t»ep«id«s sp. A
iungentitana*} piimihnis
Paraiungentitanais I»n0idijStatus
tatttanais bakiermshevi
.
•
•
Sinelolms slmtatJT
i\&t$*t\a sap ax
TypMcttsftajjf pmms
EU>p<MpsiH»des: pro«peet«fiS
LU
<
Q
t
Q
3
CO
Q.
Pa raps.
CRYPTOCOPIDAE
Figure 5. A modification of the consensus of six most parsimonious trees with the proposed
phylogeny of Pseudotanaidae and Cryptocopidae. Included are the locations of subfamilies
Pseudotanainae, Parapseudotanainae (Paraps.), Akanthinotanainae (Akanth.), Cryptocopinae
(CI), Cryptocpoidinae (C2) and Iungentitanainae (lung.). Values shown are decay indices.
among the characters used. Therefore the following proposed taxonomy is largely conjectural
and based more on observed groupings of character traits rather than support values. It is
provisional until a more detailed character analysis, possibly involving molecular techniques, can
be performed that will hopefully elucidate relationships among this highly variable and
widespread family. Three subfamilies are proposed: the Akanthinotanainae (ten species), the
Parapseudotanainae (two species) and the Pseudotanainae (33 species).
The Akanthinotanainae is clearly the more ancestral subfamily and is separated from the
other two subfamilies chiefly by its less derived carpal setae on pereopods 2 to 6 (Figs. 76 and
31
83). The former Pseudotanais subgenus Akanthinotanais is hereby elevated to generic rank and
is the sole representative of the subfamily. Two sub-genera within the Akanthinotanainae are
recognized: Akanthinotanais, s.s. with nine species and Guilleitanais represented by a single
species, A. guillei which features ancestral characters that place it not only outside the
Akanthinotanais clade but outside the clade containing the rest of the family as well. These
characters include plesiomorphic (more complex) setation of the maxilliped endites (Fig. 51) and
less derived setation of some pereopod articles (Figs. 68 and 72).
The Parapseudotanainae is shown as part of clade B (Fig. 1) but has features that
sufficiently distinguish it from the remaining members of the family such that its node could also
emerge prior to the Pseudotanainae; in fact, this is demonstrated in the jackknife and bootstrap
analyses (Figs. 2 and 3). Ancestral features retained by this subfamily include a short antennule
article 1 which is less than half the length of the antennule (Fig. 37), a cheliped fixed finger with
two inferior setae (Fig. 57), and the first pereopod carpus being more setose than those of all but
three of the other family members (Fig. 66). Additionally, this subfamily is unique in that it has
an antenna fifth article which is greater than 6.0 times longer than broad (Fig. 45).
The Pseudotanainae is composed of 33 species found in clades A and B, exclusive of the
genus Parapseudotanais (Fig. 1). Because of its size, it necessarily shows a wide variety of
mostly apomorphic characters but with little or no support among them for further sub-family
level division. A single genus, Pseudotanais, and two sub-genera, Pseudotanais, s.s. and
Mystriocentrus, are recognized. The latter has some unique features including spatulate setae
found on the merus of pereopods 2 and 3 (Fig. 75) and the carpus of pereopods 4 to 6 (Fig. 84),
and serrated incisive margins of the chelae (a character not used in this analysis). Other
distinguishing characters of this subgenus including a short cheliped dactylus (Fig. 58), an
elongate pereopod 4 basis (Fig. 79), and short uropod exopods (Fig. 89) are features shared with
other members of the genus. For this reason the former genus Mystriocentrus is relegated to
subgenus status.
32
Taxonomy of Pseudotanaidae
Family Pseudotanaidae Sieg, 1976
Diagnosis
Eyelobes pigmented, rudimentary or absent. Pereon with six free pereonites, first reduced
in length. Pleon with five free pleonites. Antennule with three articles. Antenna with six
articles, articles 2 and 3 with or without stout spiniform setae. Mandible pars molaris broad or
narrow, with or without terminal setation. Maxillule palp terminating with two setae and endite
terminating with usually nine spiniform setae (two exceptions). Maxilla rudimentary. Maxilliped
bases completely fused and endites completely or partially fused and bearing simple setae, cusps,
or naked. Cheliped attached to body via sclerite. Chelae forcipate or not. Cheliped carpus with
usually two inferior setae (three exceptions).
Cheliped fixed finger usually with one inferior
setae (four exceptions). Cheliped proximal dactylus seta present or absent. Marsupium formed
by one pair of oostegites. Pereopods 2 to 6 carpus with or without modified blade-like setae.
Pereopods 4 to 6 ischium with one or two setae, merus with one or two setae and dactylus fused
with unguis forming claw. Pleopods usually elongate with terminal setae only (three exceptions).
Uropod exopods and endopods with one or two articles or one pseudo-articulate article.
Remarks
The diagnosis was modified after Sieg (1977), Larsen and Wilson (2002) and Larsen
(2005) from their diagnoses of the family Pseudotanaidae which included the subfamilies
Cryptocopinae and Pseudotanainae. The revised family is monophyletic (Fig. 5) and includes the
subfamilies Akanthinotanainae, Parapseudotanainae and Pseudotanainae.
Key to Subfamilies
1.
Modified blade-like carpal setae present on pereopods
Modified blade-like carpal setae absent from pereopods
2.
2
Akanthinotanainae
Blade-like carpal setae on pereopods 4 to 6; cheliped with two setae on fixed finger
33
Parapseudotanainae
Blade like carpal setae on pereopods 2 to 6; cheliped with one or no setae on fixed finger
Pseudotanainae
Subfamily Akanthinotanainae, n. subfam.
Diagnosis
As for family except: Eyelobes pigmented or absent. Antenna articles 2 and 3 lacking
stout, spiniform setae. Mandible pars molaris narrow, simple pointed (with one exception).
Maxillule endite terminating with nine spiniform setae (with one exception). Cheliped carpus
with two inferior setae. Cheliped fixed finger with one inferior seta. Pereopods 2 to 6 carpus
without modified blade-like setae. Pleopods elongate with terminal setae only.
Type Genus
Akanthinotanais, n. gen.
Subgenera Included
Akanthinotanais, s.s., n. subgen.
Guilleitanais, n. subgen.
Remarks
The original name Akanthinotanais was created as a subgenus of Pseudotanais by Sieg
(1977:30) to account for six species with "normal" carpal setae on the pereopods. In this
revision, a new subfamily within Pseudotanaidae is created to contain the ten species formerly of
this subgenus which are in turn elevated to the new genus Akanthinotanais. Within the new
genus, two new subgenera, Akanthinotanais sensu stricto, and Guilleitanais, are created to
account for major differences between the species A. guillei and the other nine species of the
genus.
34
Akanthinotanais, n. gen.
Subgenus Guilleitanais, n. subgen
Diagnosis
As for subfamily except: Eyelobes pigmented. Antenna second article shorter than third
article. Mandible pars molaris narrow, simple pointed. Maxillule endite terminating with nine
spiniform setae. Maxilliped endites completely fused and with more complex setation (with
cusps and setae). Cheliped chelae weakly forcipate, dactylus subequal in width to fixed finger
and possessing proximal seta. Pereopod 1 carpus and propodus each with two setae and dactylus
with unguis subequal in length to propodus. Pereopods 2 and 3 basis with more complex setation
(broom plus simple setae). Uropod exopods 75 percent length of endopods.
Type species
Akanthinotanais (Guilleitanais) guillei (Shiino, 1978)
Remarks
The cladistic analysis presented in chapter IV resulted in the species Pseudotanais (A)
guillei being separate from all other species of the family with a decay value of 2. A comparison
of characters reveals that this species has a combination of plesiomorphic features that are
sufficient to influence its position at the ancestral end of the cladogram and justify the creation of
a new subgenus within Akanthinotanais. These characters include more complex setation of the
maxilliped endites (Fig. 51), a greater number (two) of propodal setae on pereopod 1 (Fig. 68),
and more complex setation on the basis of pereopods 2 to 6 (Figs. 72 and 80).
Akanthinotanais (Guilleitanais) guillei (Shiino, 1978)
Synonymy
Pseudotanais guillei Shiino, 1978
Diagnosis
As for subgenus.
35
Type Locality
Indian Ocean: near Kergulen islands off Antarctica
Depth Range
1 0 - 3 2 m.
Distribution
Off Antarctica in the southern Indian Ocean.
Remarks
Shiino's original description lacked dissection of mouthparts because he had only two
specimens to work with. Sieg (1986) re-illustrated the species based on additional material,
including the mouthparts. This oculate species is similar to A. gerlachi except that the acuminate
pars molaris of the mandible is shorter, the two spiniform setae on the merus of pereopods 4 to 6
are unequal in length and the uropodal exopod is about 75 percent the length of the endopod. It
differs from A. mortenseni primarily by the shorter cheliped carpus. This species is currently the
sole representative of the subgenus.
References
Shiino(1978): 106-111, Figs. 58-60; Sieg (1986): 152, Figs. 100-104.
Subgenus Akanthinotanais sensu stricto, n. subgen.
Diagnosis
As for the subfamily.
Type Species
Akanthinotanais {Akanthinotanais) gerlachi (Sieg, 1977)
Species Included
Akanthinotanais {Akanthinotanais) gerlachi (Sieg, 1977)
Akanthinotanais {A) longipes (Hansen, 1913)
36
Akanthinotanais (A) malayensis (Sieg, 1977)
Akanthinotanais (A) mortenseni (Sieg, 1977)
Akanthinotanais {A) similis (Sieg, 1977)
Akanthinotanais (A) gaussi (Vanhoffen, 1914)
Akanthinotanais {A) scrappi (Bamber, 2005)
Akanthinotanais {A) makrothrix (Dojiri and Sieg, 1977)
Akanthinotanais {A) siegi (Kudinova-Pasternak, 1985)
Akanthinotanais (Akanthinotanais) gerlachi (Sieg, 1977)
Synonymy
Pseudotanais (Akanthinotanais) gerlachi Sieg, 1977
Diagnosis
As for subfamily except: Eyelobes pigmented. Antennule first article about 5.0 times
longer than broad. Antenna second article subequal to third article, fourth article about 6.0 times
longer than broad and fifth article less than 4.0 times longer than broad. Mandible pars molaris
narrow, simple pointed. Maxillule endite terminating with nine spiniform setae. Maxilliped
endites fused except for distal notch and without setation. Cheliped chelae weakly forcipate;
dactylus greater than half the length of the propodus, narrower than fixed finger and possessing
proximal seta. Pereopod 1 basis about 6.0 times longer than broad, with one simple seta; ischium
with two setae, merus without setae, carpus and propodus each with one seta, propodus about 6.0
times longer than broad, and dactylus with unguis distinctly shorter than propodus. Pereopods 2
and 3 basis without setae, ischium with two setae, merus with two setae, dactylus with unguis 2/3
length of propodus. Pereopod 4 basis about 3.0 times longer than broad. Pereopods 4 to 6 basis
without setae, ischium with two equal setae, and merus with two setae. Uropod exopods subequal
in length to endopods.
37
Type Locality
Indian Ocean: Malediven and Addu atolls (off coast of India)
Depth Range
5 - 3 2 m.
Distribution
Indian Ocean from the type locality.
Remarks
This species is designated the type for the subgenus based on its useage by Sieg (1977: 30) as
the type for the subgenus Akanthinolanais under Pseudotanais. An occulate species, A. gerlachi is
distinguished by its long, thin pars molaris on the mandibles, the merus of pereopods 4 to 6 with
two short spiniform setae and with subequal uropodal rami.
Reference
Sieg (1977): 35-38, Figs. 21-23, 38.
Akanthinotanais (Akanthinotanais) longipes (Hansen, 1913)
Synonymy
Pseudotanais longipes Hansen, 1913: 33.
Pseudotanais (Akanthinotanais) longipes Hansen, 1913 - Sieg (1977:30-34)
Diagnosis
As for subfamily except: Eyelobes lacking. Antennule first article greater than 7.0 times
longer than broad. Antenna second article longer than third article, fourth article greater than
12.0 times longer than broad and fifth article 4.0 times longer than broad. Mandible pars molaris
narrow with several blunt terminal and subterminal denticles. Maxillule endite terminating with
nine spiniform setae. Maxilliped endites fused between 1/3 and 2/3 length, each with one short
seta. Cheliped chelae not forcipate, propodus about 4.0 times longer than broad; dactylus greater
38
than half length of propodus, subequal in width to fixed finger and possessing proximal seta.
Pereopod 1 basis about 10.0 times longer than broad, without setae; ischium, rnerus, carpus,
propodus and dactylus each with one seta, propodus about 10.0 times longer than broad and
dactylus with unguis distinctly shorter than propodus. Pereopods 2 and 3 basis without setae,
ischium with one seta, merus with two setae, dactylus with unguis 2/3 length of propodus.
Pereopod 4 basis greater than 7.0 times longer than broad. Pereopods 4 to 6 basis without setae,
ischium with two unequal setae, and merus with two setae. Uropod exopods subequal in length to
endopods.
Type Locality
North Atlantic: southwest of Iceland
Depth Range
1280 m.
Distribution
North Atlantic from type locality.
Remarks
This species is distinguished by its elongate cheliped carpus and propodus (3.3 and 4.1
times longer than broad respectively) and the mandible pars molaris with several distal and
subdistal blunt processes (all other members of the subfamily have a simple pointed pars
molaris). It is further unique among its congenitors for having an extremely long antennule first
article and antenna third article (longer than article 2).
References
Sieg (1977): 30-34. Figs.18-20, 38; Hansen (1913): 33-34, PL. Ill, Figs. 4a-i.
Akanthinotanais {Akanthinotanais) malayensis (Sieg, 1977)
Synonymy
39
Pseudolanais (Akanthinotanais) malayensis Sieg, 1977:38
Diagnosis
As for subfamily except: Eyelobes present, with pigment. Antennule first article about
5.0 times longer than broad. Antenna second article subequal to third article, fourth article about
5.0 times longer than broad and fifth article 2.0 times longer than broad. Mandible pars molaris
narrow with simple point. Maxillule endite terminating with nine spiniform setae. Maxilliped
endites fused except for distal notch, each with one short seta. Cheliped chelae not forcipate,
propodus about 3.0 times longer than broad; dactylus less than half length of propodus, subequal
in width to fixed finger and possessing proximal seta. Pereopod 1 basis about 8.0 times longer
than broad, without setae; ischium with one seta, merus without setae, carpus with one seta,
propodus and dactylus without setae, propodus about 7.0 times longer than broad and dactylus
with unguis distinctly shorter than propodus. Pereopods 2 and 3 basis without setae, ischium with
two equal setae, merus with two setae, dactylus with unguis about half length of propodus.
Pereopod 4 basis less than 4.0 times longer than broad. Pereopods 4 to 6 basis without setae,
ischium with two equal setae, and merus with two setae. Uropod exopods half length of
endopods.
Type Locality
South Pacific Ocean: Gilbert Islands.
Depth Range
Shallow water (no depth provided).
Distribution
Western Pacific Ocean above Australia.
Remarks
40
This oculate species is similar to A. geriachi except for the comparative difference in
uropodal rami length - the exopod is about half as long as the endopod in A. malayensis whereas
the rami are subequal in A. geriachi.
Reference
Sieg (1977): 38-42, Figs. 24-26, 38.
Akanthinotanais {Akanthinotanais) mortenseni (Sieg, 1977)
Synonymy
Pseudotanais (Akathinotanais) mortenseni Sieg, 1977:42
Diagnosis
As for subfamily except: Eyelobes present, with pigment. Antennule first article about
5.0 times longer than broad, distal article 6.0 times longer than broad. Antenna second article
subequal to third article, fourth article about 6.0 times longer than broad and fifth article about 5.0
times longer than broad. Mandible pars molaris narrow with simple point. Maxillule endite
terminating with nine spiniform setae. Maxilliped endites completely fused, without setae.
Cheliped chelae not forcipate, propodus about 4.0 times longer than broad; dactylus greater than
half length of propodus, narrower than fixed finger and possessing proximal seta. Pereopod 1
basis about 8.0 times longer than broad, without setae; ischium and merus with one seta, caipus
with two setae, propodus with one seta and dactylus without setae, propodus about 5.0 times
longer than broad and dactylus with unguis distinctly shorter than propodus. Pereopods 2 and 3
basis without setae, ischium with one seta, merus with two setae, dactylus with unguis 2/3 length
of propodus. Pereopod 4 basis greater than 6.0 times longer than broad. Pereopods 4 to 6 basis
with broom setae, ischium with two unequal setae, and merus with two setae. Uropod exopods
75 percent length of endopods.
Type Locality
41
North Atlantic: St. Thomas (West Indies).
Depth Range
18-25 m.
Distribution
St. Thomas (West Indies), Florida Keys (USA).
Remarks
This oculate species is noted for its long, unusually shaped cheliped, its merus setation on
pereopods 4 to 6 (one long and one short spiniform setae) and its shallow distribution in tropical
Atlantic waters. It is most similar in appearance to A. siegi but differs among other features by
having shorter, more convex pereonites, well-developed, more setose pleopods, and shorter
uropodal rami.
References
Sieg (1977): 42-46, Figs. 27-29, 38; Sieg and Heard (1988): 48, Figs. 5-7.
Akanthinotanais {Akanthinotanais) similis (Sieg, 1977)
Synonymy
Pseudotanais {Akanthinotanais) similis Sieg, 1977:46
Diagnosis
As for subfamily except: Eyelobes lacking. Antennule first article about 5.0 times longer
than broad. Antenna second article subequal to third article, fourth article about 5.0 times longer
than broad and fifth article less than 4.0 times longer than broad. Mandible pars molaris narrow
with simple point. Maxillule endite terminating with nine spiniform setae. Maxilliped endites
fused except for distal notch, each with short seta. Cheliped chelae not forcipate, propodus 2.5
times longer than broad; dactylus greater than half length of propodus, narrower than fixed finger
and possessing proximal seta. Pereopod 1 basis about 5.0 times longer than broad, without setae; .
42
ischium, merus and carpus with one seta, propodus and dactylus without setae, propodus about
6.0 times longer than broad and dactylus with unguis distinctly shorter than propodus. Pereopods
2 and 3 basis with broom setae, ischium and merus with one seta, dactylus with unguis about half
length of propodus. Pereopod 4 basis less than 4.0 times longer than broad. Pereopods 4 to 6 basis
with broom setae, ischium with one seta and merus with two setae. Uropod exopods slightly
shorter than endopods.
Type Locality
North Atlantic: Roscoff/Bloscon - English Channel off the northern coast of France.
Depth Range
20 m.
Distribution
North Atlantic off the northern coast of France (type locality).
Remarks
A non-oculate shallow water species, A. similis is most similar to A. gaussi but differs by
having a single distal spinifonn seta on pereopods 4 to 6 which is shorter than the claw (A. gaussi
has two such setae that extend beyond the claw) and the propodus on the same appendages are
about twice the length of the claw, whereas those on A. gaussi are 3.0 times the length.
Reference
Sieg (1977): 46-49, Figs. 30-33, 38.
Akanthinotanais (Akanthinotanais) gaussi (Vanhoffen, 1914)
Synonymy
Pseudotanais gaussi Vanhoffen, 1914:438
Pseudotanais {Akanthinotanais) gaussi Vanhoffen, 1914 - Sieg (1977): 50
Diagnosis
43
As for subfamily except: Eyelobes lacking. Antennule first article about 6.0 times longer
than broad. Antenna second article subequal to third article, fourth article about 10.0 times longer
than broad and fifth article about 3.0 times longer than broad. Mandible pars molaris narrow with
simple point. Maxillule endite terminating with nine spiniform setae. Maxilliped endites fused
between 1/3 and 2/3 length, each distally with long simple, seta. Cheliped chelae not forcipate,
propodus 2.5 times longer than broad; dactylus greater than half length of propodus, narrower
than fixed finger and possessing proximal seta. Pereopod 1 basis about 8.0 times longer than
broad, with broom seta; ischium and merits with one seta, carpus and propodus without setae and
dactylus with short seta, propodus about 11.0 times longer than broad and dactylus with unguis
distinctly shorter than propodus. Pereopods 2 and 3 basis with broom setae, ischium and merus
with one seta, dactylus with unguis subequal to length of propodus. Pereopod 4 basis about 6.0
times longer than broad. Pereopods 4 to 6 basis with simple and broom setae, ischium and merus
with one seta. Uropod exopods about 2/3 length of endopods.
Type Locality
South Atlantic: off Antarctica.
Depth Range
385-1425m.
Distribution
South Atlantic Ocean off Antarctica.
Remarks
This species is distinctive with its two long infero-distal spiniform setae on the propodus
which are longer than the claw which is, in turn, about 1/3 the length of the propodus.
References
Vanhoffen (1914): 438, Figs. 20a-c; Sieg (1977): 50-55, Figs. 34-38.
44
Akanthinotanais {Akanthinotanais) scrappi (Bamber, 2005)
Synonymy
Pseudotanais (Akanthinotanais) scrappi Bamber, 2005: 720
Diagnosis
As for subfamily except: Eyelobes present, with pigment. Antennule first article about
3.0 times longer than broad. Antenna second article shorter than third article, fourth article about
5.0 times longer than broad and fifth article about 3.0 times longer than broad. Mandible pars
molaris narrow with simple point. Maxillule endite terminating with seven spiniform setae.
Maxilliped endites completely fused, without setae. Cheliped chelae not forcipate, propodus
about 3.0 times longer than broad; dactylus less than half length of propodus, subequal in width to
fixed finger and lacking proximal seta. Pereopod 1 basis about 7.0 times longer than broad,
without setae; ischium, merus, carpus, propodus and dactylus without setae, propodus about 7.0
times longer than broad and dactylus with unguis subequal in length to propodus. Pereopods 2
and 3 basis with simple seta (on pereopod 3), ischium with two equal setae, merus with two setae,
dactylus with unguis about half length of propodus. Pereopod 4 basis less than 4.0 times longer
than broad. Pereopods 4 to 6 basis without setae, ischium with two equal setae, and merus with
two setae. Uropod exopods subequal in length to endopods.
Type Locality
South Pacific: Esperance, South Australia
Depth Range
38 m.
Distribution
Pacific Ocean off the coast of Esperance, South Australia (type locality)
Remarks
45
In appearance and pereopod setation, this oculate species is most similar to A. guillei
from the Subantarctic but differs by having a stouter body size of 3.8 times longer than broad
(opposed to 5.0 times for A. guillei), bi-articulate uropodal rami, and more elongate antennal
articles. The unusual armament of the maxillule endite (seven terminal spines rather than the
normal nine) is unique for the subfamily.
Reference
Bamber (2005): 720-723, Figs. 55, 56.
Akanthinotanais (Akanthinotanais) makrothrix (Dojiri and Sieg, 1977)
Synonymy
Pseudotanais makrothrix Dojiri and Sieg, 1977: 258
Diagnosis
As for subfamily except: Eyelobes present, with pigment. Antennule first article about
6.0 times longer than broad. Antenna not described. Mandible pars molaris narrow with simple
point. Maxillule not described. Maxilliped endites completely fused, each with short seta.
Cheliped chelae not forcipate, propodus about 3.0 times longer than broad; dactylus greater than
half length of propodus, narrower than fixed finger and possessing proximal seta. Pereopod 1 not
described. Pereopod 2 basis with broom seta, ischium with one setae, merus with two setae,
dactylus with unguis about 2/3 length of propodus. Pereopods 4 and 5 not described. Uropod
exopods subequal in length to endopods.
Type Locality
North Pacific: Santa Maria Basin off California, USA.
Depth Range
98-393 m.
Distribution
46
Pacific Ocean off California, USA (type locality).
Remarks
The original description by Dojiri and Sieg is inadequate because the body length and
descriptions and illustrations of most appendages were omitted (of the pereopods, only pereopod
2 and part of pereopod 6 were illustrated). As a result, the precision of the analysis was likely
affected due to missing data. The authors, in comparing this oculate species to the similar A.
guillei, stated that it is different by "its slender and longer propodal spines, most noticeable on
periopod 6" which in A. makrothrix are "more than half as long as the propodus". They
erroneously termed these structures as propodal spines when, in fact, they were referring in the
illustrations to the carpal spiniform setae which do not appear to be much different than those
illustrated for A. guillei by Sieg (1986, Fig. 102); the latter species appears to have caipal
pereopod spiniform setae that are, in pereopods 2 and 3, subequal to the propodus.
Reference
Dojiri and Sieg (1977): 258, Fig. 3.29.
Akanthinotanais {Akanthinotanais) siegi (Kudinova-Pasternak, 1985)
Synonymy
Pseudotanais siegi Kudinova-Pasternak, 1985:60
Diagnosis
As for subfamily except: Eyelobes present, with pigment. Antennule first article 6.5
times longer than broad, distal article about 9.5 times longer than broad. Antenna second article
subequal to third article, fourth article about 11.0 times longer than broad and fifth article more
than 6.0 times longer than broad. Mandible pars molaris narrow with simple point. Maxillule
endite terminating with nine spiniform setae. Maxilliped endites completely fused, each with
short seta. Cheliped chelae not forcipate, propodus about 5.0 times longer than broad; dactylus
47
greater than half length of propodus, narrower than fixed finger and lacking proximal seta.
Pereopod 1 basis about 9.0 times longer than broad, without setae; ischium, merus, carpus and
propodus with one seta and dactylus without setae, propodus about 9.0 times longer than broad
and dactylus with unguis distinctly shorter than propodus. Pereopods 2 and 3 basis without setae,
ischium and merus with one seta, dactylus with unguis 2/3 length of propodus. Pereopod 4 basis
about 9.0 times longer than broad. Pereopods 4 to 6 basis without setae, ischium with one seta,
and merus with two setae. Uropod exopods 2/3 length of endopods.
Type Locality
North Atlantic: Great Meteor Seamount.
Depth Range
325-470m.
Distribution
North Atlantic Ocean near the Great Meteor seamount (type locality).
Remarks
This species is similar to A. mortenseni in some features (see remarks for A. mortenseni).
Unique characteristics for A siegi include a long seta on the merus of the cheliped supported by a
small protuberance, serrate incisive margins on the chela, notably long, spindly pereopods and
antennae, and an extremely long distal carpal seta on pereopods 4 to 6 that extends beyond the
dactylus. Setation on the pereopods was inadequately described and illustrated and was therefore
assumed to be simple setae. Original description is in Russian.
Reference
Kudinova-Pasternak (1985): 60-63, Figs. 5,6.
Subfamily Parapseudotanainae, n. subfam.
Diagnosis
48
As for family except: Eyelobes absent. Cephalothorax longer than pereonites 1 to 3.
Pereonites 4 and 5 distinctively elongate. Antennule first article less than half length of entire
antennule. Antenna second and third articles with simple setae, fifth article greater than 6.0 times
longer than broad. Mandible pars molaris narrow, bifid-tipped or with marginal setules.
Maxillule endite terminating with nine spiniform setae. Maxilliped endites fused between 1/3 to
2/3 length, bearing cusps or setae. Chelae not forcipate. Cheliped propodus greater than 1.5
times longer than carpus, carpus and fixed finger with two inferior setae, dactylus subequal in
width to fixed finger and bearing proximal seta. Pereopod 1 merus with one seta, carpus with
three setae, dactylus with exceptionally long seta and dactylus with unguis subequal to length of
propodus. Pereopods 2 and 3 carpus with modified blade-like setae, half or less length of
propodus; dactylus with unguis about half length of propodus. Pereopods 4 to 6 basis without
setae, ischium with one seta, merus with two setae, carpus lacking modified blade-like setae and
propodus lacking broom setae. Pleopods lacking or elongate with terminal setae. Uropod
exopods with two articles, subequal in length to endopods; endopods with one or two articles.
Type Genus
Parapseudotanais Bird and Holdich, 1989
Remarks
This subfamily currently comprises a single genus and has features that sufficiently
distinguish it from the other two subfamilies of Pseudotanaidae. Although poorly supported in
the cladistic analysis, as was also the case for Pseudotanainae, it possesses ancestral characters
that place it somewhere between the Akanthinotanainae and the more derived Pseudotanainae.
These include a short antennule article 1 which is less than half the length of the antennule (Fig.
37), a cheliped fixed finger with two inferior setae (Fig. 57), and a first pereopod carpus that is
more setose than most of the other family members (Fig. 66). Additionally, this subfamily is
unique in that it has an apomorphic fifth antenna article which is greater than 6.0 times longer
than broad (Fig. 45).
49
Species Included
Parapseudotanais abyssalis Bird and Holdich, 1989
Parapseudotanais sp. A., n. sp.
Parapseudotanais abyssalis Bird and Holdich, 1989
Diagnosis
As for subfamily except: Antennule distal article about 11.0 times longer than broad.
Antenna 75 percent length of antennule, second article longer than third, fourth article 7.0 times
longer than broad, and fifth article nearly 8.0 times longer than broad. Mandible pars molaris
narrow, bifid-tipped. Maxilliped endites bearing marginal cusps. Cheliped carpus twice as long
as broad, propodus about 3.0 times longer than broad and dactylus about 60% of propodus length.
Pereopod 1 basis 9.0 times longer than broad, without setae; ischium with one seta and propodus
5.0 times longer than broad, with two setae. Pereopods 2 and 3 basis without setae and ischium
and merus each with one seta. Pereopod 4 basis 3.0 times longer than broad. Pleopods lacking.
Type Locality
Northeast Atlantic: northern Bay of Biscay.
Depth Range
4226-4327 m.
Distribution
Northeast Atlantic Ocean in the Bay of Biscay (type locality).
Remarks
This species is distinguished by the presence of two inferior setae on the fixed finger of
the chela and small blade-like spiniform setae on the carpus of pereopods 2 and 3 which are
replaced by stout serrate spiniform setae on pereopods 4-6.
Reference
50
Bird and Holdich (1989): 282-285, Figs. 23, 24.
Parapseudotanais sp. A, n. sp. (Figures 6 - 9 )
Diagnosis
As for the subfamily and P. abyssalis except:
Antennule distal article 8.0 times longer than broad. Antenna 82 percent length of antennule,
second article shorter than third and fifth article 7.0 times longer than broad.
Mandible pars molaris narrow, bifid-tipped and laterally setulate. Maxilliped endites each
bearing short seta. Cheliped carpus 2.5 times longer than broad. Pereopod 1 basis 7.0 times
longer than broad, with simple setae; ischium without setae and propodus 6.0 times longer than
broad, with one seta. Pereopods 2 and 3 basis with simple setae, ischium without setae and merus
with two setae. Pereopod 4 basis 4.0 times longer than broad. Pleopods elongate, with terminal
setae.
Type Locality
North Atlantic: Gulf of Mexico, Texas A&M Deepwater Gulf of Mexico Station S35-1
in the western Gulf of Mexico.
Depth Range
533-1460m.
Distribution
Gulf of Mexico
Material Examined
Holotype, preparatory female, 1.9 mm., DGoM Station S35-1, 29°20.0500'N
87°03.3758'W, western Gulf of Mexico, 11 June 2000, 658 m. Paratypes: one preparatory female,
1.8 mm and two non-ovigerous females, 1.35 mm - one dissected and slide mounted, from type
locality. Other paratypes: DGoM Station W4—2, one female, 2.2 mm; DGoM Station S35-5, two
females, 0.9 mm; DGoM Station BH-5, two females, 1.5 mm, - one dissected and slide mounted.
51
Description
Based on holotype (Fig. 6A,B) and non-ovigerous female paratypes. Body 1.4 to 2.2
mm, 4.8 times longer than broad.
Cephalothorax (Fig. 6A) longer than pereonites 1-3, subtriangular, becoming narrow
anteriorly, with pair of antero-lateral setae, 17.2 percent total length; eyelobes absent.
Pereon 48.6 percent total length, pereonites 1-3 abreviated; pereonites 1 and 3-6 with
pair of dorso-lateral setae (those of pereonite 1 more prominent).
Pleon shorter than pereonites 5+6, 15.3 percent total length, pleonites 4 and 5 with pair of
dorso-lateral setae.
Pleotelson (Fig. 7B) subequal to pleonites 4-5, 7.3 percent total length, subtriangular,
with pair of dorso-lateral setae, pair of ventro-lateral setae near emergence of uropod basal article,
and four apical setae.
Antennule (Fig. 7D) 26 percent of body length, with three articles; first article 46.4
percent of total length, 4.3 times longer than broad, with two short medial simple setae, five short
hair-like medial setae, four short hair-like distal setae and one long distal simple seta; second
article 2.8 times longer than broad, with one long distal simple seta and two hair-like distal setae;
distal article 7.9 times longer than broad, terminating with four simple setae (three long, one
short) and three aesthetascs.
Antenna (Fig. 7E) with six articles, 82.3 percent length of Al; second article 0.6 times
longer than broad, about 2/3 length of third article, both articles distally with short, slender
spiniform seta; fourth article 6.8 times longer than broad, with four short distal simple setae and
one distal broom seta; fifth article 6.7 times longer than broad, distally with one long simple seta;
sixth article small, terminating with one aesthetasc and three simple setae (one long, distally bent,
apparently fused to aesthetasc).
Mouthparts. Labrum (Fig. 8C) broad, cap-shaped, non-setose. Mandibles (Fig. 8A,B)
with distal margins coarsely denticulate (left) and finely denticulate (right), right lacinia mobilis
52
represented by short, pointed process, left lacinia mobilis well developed and coarsely
denticulate, pars molaris acuminate, minutely bifid (left only) and with 7-10 setules along
anterior margin. Labium (Fig. 8D) broad, medially cleft, lobes non-setulose. Maxillule endite
(Fig. 8E) terminating with nine spiniform setae and pair of long sub-terminal accessory setae.
Maxilla not observed. Maxilliped (Fig. 8F) bases completely fused, with small seta near each
palp articulation; endites fused between 1/3 and 2/3 length, each with one short seta, with lateral
margins smooth; palp with four articles; article 1 without setae; article 2 with two unequal inneredge simple setae and one outer-edge simple seta; article 3 with three inner-edge spiniform setae
and one inner-edge simple seta; article 4 with one small outer-edge simple seta and five inneredge simple setae. Epignath not observed
Cheliped (Fig. 7A) large, somewhat elongate, chela not forcipate; sclerite well
developed; basis about same size as sclerite and 2.0 times longer than broad, with distal superior
seta; merus triangular, with inferior seta; carpus 2.5 times longer than broad, with two medial
inferior setae, one distal superior seta and one medial superior seta; propodus 3.0 times longer
than broad, about 1.8 times longer than carpus, palm with one long and seven short comb setae;
fixed finger with two inferior setae, three superior setae, one seta near articulation with dactylus
and two low profile teeth on incisive margin; dactylus 57.1 percent of propodus length, width
subequal to fixed finger, with proximal simple seta and subdistal low profile tooth on incisive
margin.
Pereopod 1 (Fig. 9A) more slender and longer than pereopods 2 and 3; coxa with seta;
basis 7.0 times longer than broad, with medial superior seta and proximal superior seta; ischium
without setae; merus 1.3 times longer than broad, distally with short inferior seta; carpus 2.0
times longer than broad, 1.4 times length of merus, distally with three short setae; propodus 5.8
times longer than broad, with superior marginal spinules, distally with short inferior seta; dactylus
with long seta, length with unguis subequal to length of propodus.
53
Pereopod 2 (Fig. 9B) coxa with seta; basis 5.5 times longer than broad, with short
proximal inferior seta; ischium with setae lacking; merus 2.1 times longer than broad, distally
with two unequal inferior setae; carpus 2.2 times longer than broad, about 0.9 times length of
merus, distally with one short superior spiniform seta, one short, broad posterior spiniform seta,
inferior bayonet-like seta of about 50 percent length of propodus and four transverse rows of
inferior marginal setules; propodus 3.4 times longer than broad, with one short distal inferior
spiniform seta (about third length of dactylus-unguis); dactylus stout, completely fused with
unguis, and without setae, length about half that of propodus.
Pereopod 3 (Fig. 9C) carpus 1.6 times longer than broad; otherwise similar to pereopod
2.
Pereopod 4 (Fig. 9D) basis 4.1 times longer than broad, tumid, with setae lacking;
ischium with one short simple seta; merus 2.0 times longer than broad, distally with two short
unequal serrate spiniform setae; carpus 5.4 times longer than broad, 2.7 times longer than merus,
distally with one short superior simple seta, four short unequal serrate spiniform setae and four
semicircular rows of inferior marginal setules; propodus 5.1 times longer than broad, slightly
shorter than carpus, distally with one long superior terminal spiniform seta, two unequal inferior
spiniform setae, superior marginal spinules and setulose terminal margin at emergence of
dactylus; dactylus fused with unguis to form claw, length with unguis less than half length of
propodus.
Pereopod 5 (Fig. 9E) similar to pereopod 4.
Pereopod 6 (Fig. 9F) carpus 3.9 times longer than broad; distally with one less serrate
spiniform seta, propodus with additional terminal spiniform seta, otherwise similar to pereopods 4
and 5 but proportionally shorter.
Pleopods (Fig. 7C) rami elongate, with terminal setae only; endopod about 4/5 length of
exopod, with five terminal setae; exopod with seven terminal setae.
54
Uropods (Fig. 7B) rami slender, elongate; exopod with two articles, about 80 percent
length of endopod, proximal article about half length of ramus, with one distal simple seta, distal
article with two unequal simple setae; endopod with one article, with two medial broom setae,
one long sub-terminal simple seta, three long and two short terminal simple setae, and one
terminal broom seta.
Remarks
This new species is similar to P. abyssalis from the northeast Atlantic, sharing a
characteristic body shape with abbreviated first three pereonites, similar proportions and setation
for antennae and pereopods, and nearly identical chelipeds. Basic differences that distinguish the
new species from its congenitor include (1) having a uni-articulate uropodal endopod, (2) more
modified mandible pars molaris with marginal setae, and (3) maxillipeds with basal setae, endites
with small setae instead of cusps, and one fewer inner edge setae on the first article of the palp. In
addition, the new species appears to have modified bayonet-like spiniform setae on the carpus of
pereopods 2 and 3 similar in appearance to the "blade-like" setae illustrated by Bird and Holdich
(1989:284). This might just be a matter of interpretation. It should also be noted those authors
based their description on just five poorly preserved, non-ovigerous specimens collected from a
small area in the Bay of Biscay in much deeper water (below 4000 m) than that observed for the
Gulf of Mexico species (533-1460 m).
55
Figure 6. Parapseudotanais sp. A. Holotype, female. A, dorsal view. B,
lateral view. Scale = 0.5 mm
56
Figure 7. Parapseudotanais sp. A. Parataype, female. A, Cheliped inner face. B, Pleotelson
and uropods, ventral view. C, Pleopods. D, Antennule. E, Antenna. Scale = 0.2mm
57
Figure 8. Parapseudotanais sp. A. Paratype, female. A, Left mandible, inset shows detail of
pars molaris. B, Right mandible. C, Labrum. D, Labium. E, Maxillule endite. F, Maxillipeds.
Scale = 0.2mm.
58
Figure 9. Parapseudotanais sp. A. Paratype, female. A-F, Pereopods 1 to 6 respectively.
Scale = 0.2mm
59
Subfamily Pseudotanainae Sieg, 1977
Diagnosis
As for family except: Antenna fifth article less than 6.0 times longer than broad.
Maxillule endite terminating with usually nine spiniform setae (one exception). Cheliped
propodus greater than 1.5 times longer than carpus and fixed finger usually with one inferior setae
(two exceptions).
Pleopods usually elongate with terminal setae only (two exceptions).
Type Genus
Pseudotanais G.O. Sars, 1882
Subgenera Included
Mystriocentrus, n. subgen.
Pseudotanais, s.s. Sieg, 1977:55
Remarks
The diagnosis presented here differs from those presented by Sieg (1977), Bird and
Holdich (1989) and Larsen (2005) largely because of the exclusion of the previously discussed
subfamilies. It is based entirely on characters used in the cladistic analysis and is necessarily
broad given the large number of species (33) with numerous divergent characters.
Key to Subgenera
1. Chela strongly forcipate with serrate incisive margins; spatulate-tipped setae present on merus
of pereopods 2 and 3 and carpus of pereopods 4 to 6
Mystriocentrus
2. Chela may be forcipate but incisive margins not serrate; without spatulate-tipped setae as
above
Subgenus Mystriocentrus, n. subgen.
Diagnosis
Pseudotanais
60
As for family and subfamily except: Eyelobes absent. Antenna article 2 subequa! in
length to article 3, both without stout spiniform setae. Mandible pars tnolaris narrow, with or
without bifid tip. Maxillule endite terminating with nine spiniform setae. Maxilliped bases and
endites completely fused, endites bearing cusps or naked. Chelae strongly forcipate, with serrate
incisor margins. Cheliped carpus with two inferior setae, fixed finger with one inferior seta;
dactylus greater than 63 percent of propodus and subequal in width to fixed finger. Pereopod 1
basis without setae, merus with and carpus with one seta, propodus with two setae, dactylus with
unguis distinctly shorter than propodus. Pereopods 2 and 3 ischium with one seta, merus with
one modified spatulate-tipped seta, carpus with one modified blade-like seta less than half length
of propodus. Pereopod 4 basis greater than 6.0 times longer than broad. Pereopods 4 to 6 basis
with broom setae, ischium and merus with one seta, carpus with one blade-like seta and one
spatulate-tipped seta and propodus with broom seta. Pleopods elongate with terminal setae only.
Type Species
Pseudolanais (Mystriocentrus) serratus (Bird and Holdich, 1989)
Species Included
Pseudotanais {Mystriocentrus) serratus (Bird and Holdich, 1989)
Pseudolanais (Mystriocentrus) sp. A, n. sp.
Remarks
Members of this subgenus possess some unique features such as the derived spatulatetipped setae and serrate chelae; however, nearly all other diagnostic characters are shared by at
least some other members of the family. Therefore the former genus Mystriocentrus was
relegated to subgenus status.
Pseudotanais (Mystriocentrus) serratus (Bird and Holdich, 1989)
61
Synonymy
Mystriocentrus serratus Bird and Holdich, 1989: 27'/
Diagnosis
As for subgenus except: Cephalothorax subequal to pereonites 1 to 3. Pereonite 3
abreviated (shorter than pereonite 2). Antennule 93 percent length of antenna, first article about
6.0 times longer than broad. Antenna fourth article about 9.0 times longer than broad. Mandible
pars molaris simple pointed. Maxilliped endites completely fused, naked. Cheliped carpus about
2.5 times longer than broad; propodus greater than 1.5 times longer than carpus, about 3.0 times
longer than broad; dactylus 72 percent of propodus length, without proximal seta. Pereopod 1
ischium and dactylus without setae. Pereopods 2 and 3 basis without setae, merus with two setae
(in addition to spatulate-tipped setae) and dactylus with unguis distinctly less than half length of
propodus. Pereopod 4 basis about 8.0 times longer than broad. Uropod exopods with one
pseudo-articulated article, about 75 percent length of endopods; endopods likewise with one
pseudo-articulated article.
Type Locality
Northeast Atlantic: Rockall Trough, west of Hebrides Terrace Seamount.
Depth Range
1378-4632m.
Distribution
North Atlantic Ocean west of Ireland.
Remarks
This species is characterized by forcipate chelae with serrate incisive margins and
distinctive spatulate setae on pereopods 2 to 3 (merus) and 4 to 6 (carpus). In addition to the
diagnosis, this species also possesses thickened setae on the sixth antennal article and also on the
62
fourth article of the maxilliped palp. Bird and Holdich (1989) found this species to be
widespread both geographically and bathymetrically in Northeast Atlantic stations off Ireland.
Reference
Bird and Holdich (1989): 277-281, Figs. 1C, U, 2C, 21-22.
Pseudotanais (Mystriocentrus) sp. A, n. sp. (Figures 10 - 13)
Diagnosis
As for subgenus and P. serratus except: Cephalothorax longer than pereonites 1 to 3.
Pereonite 3 not abbreviated. Antennule 82 percent length of antenna, first article about 4.0 times
longer than broad. Antenna fourth article about 6.0 times longer than broad. Mandible pars
molaris minutely bifid-tipped. Maxilliped endites completely fused, bearing 2 short cusps.
Cheliped propodus about 2.5 times longer than broad; dactylus 65 percent of propodus length,
with proximal seta. Pereopod 1 ischium and dactylus each with one seta. Pereopods 2 and 3
basis with broom and simple setae, merus with three setae (in addition to spatulate-tipped setae)
and dactylus with unguis about half length of propodus. Pereopod 4 basis about 11.0 times longer
than broad. Uropod exopods with two articles, half length of endopods; endopods with two
articles.
Type Locality
North Atlantic: Gulf of Mexico, Texas A&M Deepwater Gulf of Mexico Station W2-2
in the western Gulf of Mexico.
Depth Range
212-3007 m.
Distribution
Gulf of Mexico
Material Examined
63
Holotype, preparatory female, 0.85 mm, DGoM station W2-2, 27°24.7019'N 93°20.3849'
W, western Gulf of Mexico, 14 May 2000, 625 m. Paratypes: DGoM Station HP-3, one female,
0.95 mm, dissected and slide mounted; DGoM Station MT3-1, one manca; DGoM Station C7-2,
one female, one manca; DGoM Station MT2-1, eight females (two preparatory), two specimens,
0.9 mm, mouthparts dissected and slide mounted; DGoM Station MT3-1, two females (one
preparatory); DGoM Station MT3-2, one female; DGoM Station RW1-2, two females; DGoM
Station RW2-1, one female; DGoM Station RW2-2, two females; DGoM Station RW3-1, one
ovigerous female; DGoM Station RW4-1, one female; DGoM Station S35-1, two females, 0.6
mm; DGoM Station S39-1, one preparatory female, 1.0 mm, dissected and slide mounted; DGoM
Station W2-1, three females, 0.85 mm, one possible male morph; DGoM Station Cl-2, four
females (one with spent marsupium); DGoM Station C7-2, two females with marsupium
remnants; DGoM Station S35-2, one female; DGoM Station AC 1-3, one female; DGoM Station
C4-2, one female; DGoM Station MT2-2, five females, 1.0 mm; DGoM Station MT5-3, one
female; DGoM Station RW3-3, one female; DGoM Station W2-3, two females, 0.9 mm,
dissected and slide mounted; DGoM Station W3-2, two females, 0.8 mm; DGoM Station S44-3,
one female 0.6 mm; DGoM Station RW1-3, two mancas (one with nematode parasite); DGoM
Station W2-3, one female.
Description based on holotype (Fig. 10A, B) and non-ovigerous female paratype. Body
0.85-1 mm, 3.2 times longer than broad.
Cephalothorax longer than pereonites 1-3, subtriangular, becoming narrow anteriorly,
with slight rostrum and pair of antero-lateral setae, 20.1 percent total length; eyelobes absent.
Pereon 54.7 percent total length, pereonite 1 abreviated; each pereonite with pair of small
dorso-lateral setae.
Pleon shorter than pereonites 5+6, 16 percent total length, each pleonite with pair of
small dorso-lateral setae.
64
Pleotelson subequal to pleonites 3-5, 9.4 percent total length, apex slightly produced
ventrally, with two apical setae.
Antennule (Fig. 1 IB) 24.2 percent of body length, with three articles; first article 57.1
percent of total length, 3.8 times longer than broad, with four short medial simple setae, three
short distal simple setae and one long distal simple seta; second article 1.8 times longer than
broad, with one long distal simple seta and one short distal simple seta; distal article 2.5 times
longer than broad, terminating with two simple setae (one long, one short), one aesthetasc, one
bifid-tipped seta, two spatulate setae and one hook-tipped seta.
Antenna (Fig. 11C) with six articles, 122 percent length of Al; second article 1.4 times
longer than broad, equal in length to third article, both articles distally with short, slender simple
seta; fourth article 6.2 times longer than broad, with four short distal simple setae and one long
distal simple seta; fifth article 3.8 times longer than broad, distally with one long simple seta;
sixth article small, terminating with one spatulate seta and three simple setae (one long, distally
bent).
Mouthparts. Labrum (Fig. 12C) cap-shaped, with marginal ridge. Mandibles (Fig. 12A,
B) with distal margins denticulate, right lacinia mobilis represented by short, pointed process, left
lacinia mobilis well developed and denticulate, pars molaris acuminate, minutely bifid. Labium
(Fig. 12D) medially cleft, lobes non-setose. Maxillule (Fig. 12F) palp with two equal setae,
terminating with nine spiniform setae, one much thinner than other eight. Maxilla not observed.
Maxilliped (Fig. 12E) bases completely fused, without seta near palp articulation; endites
completely fused (no notch), each with two short cusps, with lateral margins smooth; palp with
four articles; article 1 without setae; article 2 with one medio-distal extremely long simple seta
and one inner-edge simple seta; article 3 with five unequal inner-edge simple setae; article 4 with
one small outer-edge simple seta, four inner-edge simple setae, and one inner-edge thick
spiniform seta. Epignath not observed.
65
Cheliped (Fig. 11A) chela strongly forcipate; sclerite small; basis 1.5 times longer than
broad, without setae; merus triangular, with inferior seta; carpus 2.4 times longer than broad, with
two unequal medial inferior setae, one distal superior seta and one proximal superior seta;
propodus 2.4 times longer than broad, about 1.8 times longer than carpus, palm with seven comb
setae (one longer than others); fixed finger with one inferior seta, three superior setae, one seta
near articulation with dactylus, two low profile teeth on incisive margin and serrate incisive
margin; dactylus 64.7 percent of propodus length, width subequal to fixed finger, with proximal
spiniform seta, two low profile sub-terminal teeth on incisive margin and serrate incisive margin.
Pereopod 1 (Fig. 13A) basis 8.0 times longer than broad, with no setae; ischium with
small seta; merus 2.2 times longer than broad, distally with long inferior seta; carpus 3.0 times
longer than broad, 1.2 times length of merus, distally with one short superior seta; propodus 5.3
times longer than broad, distally with short superior seta and short inferior seta; dactylus with
seta, length with unguis 80 percent length of propodus.
Pereopod 2 (Fig. 13B) basis 5.0 times longer than broad, with short proximal superior
seta; ischium with short inferior seta; merus 1.8 times longer than broad, distally with one long
inferior simple seta, one short inferior spiniform seta, long spatulate seta, and two transverse rows
of inferior marginal spinules; carpus 1.5 times longer than broad, about 0.8 times length of merus,
distally with one short superior simple seta, one short anterior simple seta, one inferior blade-like
seta of about 35 percent length of propodus and six transverse rows of inferior marginal setules;
propodus 5.3 times longer than broad, with one short distal inferior spiniform seta (about half
length of dactylus-unguis), one short distal inferior simple seta, five transverse rows of inferior
marginal spinules and setulose terminal margin at emergence of dactylus; dactylus without setae,
length with unguis about half length of propodus.
Pereopod 3 (Fig. 13C) basis 5.8 times longer than broad and with superior medial broom
seta; ischium with short superior simple seta; merus with three spatulate distal setae, short distal
spiniform seta, and no transverse marginal spinules, merus 2.5 times longer than broad; carpus
66
2.3 times longer than broad, carpus with inferior blade-like seta of about 45 percent length of
propodus and no transerse marginal spinules; with short distal superior seta; dactylus length with
unguis about 75 percent length of propodus; otherwise similar to pereopod 2.
Pereopod 4 (Fig. 13D) basis 10.6 times longer than broad, with one medial broom seta;
ischium with one short simple seta; merus 2.0 times longer than broad, distally with one long
inferior simple seta; carpus 4.4 times longer than broad, 2.2 times longer than merus, distally with
one short anterior simple seta, one short inferior spiniform seta and one superior spatulate seta;
propodus 4.7 times longer than broad, slightly shorter than carpus, distally with one long superior
terminal spiniform seta, two long inferior spiniform setae and one superior broom seta; dactylus
fused with unguis to form claw, with small hook at tip of unguis, length with unguis less than half
length of propodus.
Pereopod 5 (Fig. 13.E) basis 7.1 times longer than broad, carpus distally with additional
short spiniform seta and inferior blade-like seta of about 25 percent of propodus length, propodus
5.9 times longer than broad and with three transverse rows of superior marginal spinules,
otherwise similar to P4.
Pereopod 6 (Fig. 13F) basis 6.6 times longer than broad and lacking broom seta, carpus
distally with additional inferior spiniform seta, propodus with additional terminal spiniform seta
and lacking broom seta, otherwise similar to peropod 4.
Pleopods (Fig. 11D) rami elongate, with terminal setae only; endopod slightly shorter
than exopod, with four terminal setae; exopod with six to eight terminal setae.
Uropods (Fig. 11E) exopod with two articles, half length of endopod, proximal article
about 2/3 length of ramus, with one distal simple seta, distal article with one long simple seta;
endopod with two articles, proximal article slightly greater than half length of ramus, with two
distal unequal simple setae, distal article with four long and two short simple setae.
Remarks
67
This new species is very similar in many respects to Pseudotanais (M) serratus especially
in the presence of serrate, forcipate chelae and the spatulate setae on pereopods 2 to 6. It differs
however by lacking the abbreviated pereonite 3, having a longer cephalothorax in comparison to
pereonites 1 to 3 (subequal in P. serratus), having a shorter pleon and longer pleotelson, and
having minutely hooked unguis on pereopods 4 to 6. The mandibles show subtle differences in
that the pars molaris appears to be minutely bifid (single pointed in P. serratus). The maxillipeds
are similar in that both species have completely fused endites; however the new species lacks
setae on the bases near the articulation point of the palps and possesses a pair of minute cusps
submarginally on each endite which are lacking on P. serratus. The maxilliped palps are similar
in both species including the presence of a thickened spiniform seta on the fourth article. The
forcipate chelipeds are nearly identical in the two species except the dactyl in the new species
possesses a short proximal spiniform seta (lacking in P. serratus) and lacks the inferior peg-like
structure described for P. serratus. Additional pereopod characters of the new species which
differ from P. serratus include (1) perepod 1 dactylus with a small seta; (2) pereopod 2 with
maginal spinules on the merus similar to those on the carpus but fewer in number, carpus with
fewer rows of marginal spinules and a blade setae that is longer (35 percent of the propodus
length rather than about 20 percent), and a slightly greater dactylus-unguis length; and (3)
pereopod 3 with a broom seta on the basis, short seta on the ischium, and a large blade seta (45
percent of Propodus length) on the carpus.
68
Figure 10. Pseudotanais (Mystriocentrns) sp. A. Holotype, female. A,B,
dorsal and lateral views. Scale = 0.5 ram
69
Figure 11. Pseudotanais {Mystriocentrus) sp. A. Paratype, female. A, Cheliped inner face. B,
Antennule. C, Antenna. D, Pleopods. E, Uropods. Scale = 0.2 mm.
70
Figure 12. Pseudotanais (Mystriocentrus) sp. A. Paratype, female. A, Left mandible. B, Right
mandible (both showing detail of pars molaris). C, Labrum. D, Labium. E, Maxillipeds. F,
Maxillule. Scale = 0.2 mm.
71
Figure 13. Pseudotanais (Mystriocentrus) sp. A. Paratype, female. A-F, Pereopods 1 to 6
respectively. D showing detail of hooked unguis. Scale = 0.2 mm.
72
Subgenus Pseudotanais sensu stricto Sieg, 1977
Diagnosis
As for family and subfamily except: Antennule first article greater than half the length of
the entire antennule. Antenna article 2 shorter than or subequal to article 3, with or without stout
spiniform setae. Pereopods 2 to 6 carpus each with modified blade-like seta.
Type Species
Pseudotanais (Pseudotanais) forcipaius (Lilljeborg, 1864)
Species Included
Pseudotanais
Pseudotanais
abyssi Hansen, 1913
Pseudotanais
Pseudotanais
affmis Hansen, 1887
Pseudotanais
Pseudotanais
baresnauti Bird, 1999
Pseudotanais
Pseudotanais
borceai (Bacescu, 1960)
Pseudotanais
Pseudotanais
californiensis Dojiri and Sieg, 1997
Pseudotanais
Pseudotanais
colonus Bird and Holdich, 1989a
Pseudotanais
Pseudotanais
corollalus Bird and Holdich, 1989
Pseudotanais
Pseudotanais
denticulatus Bird and Holdich, 1989
Pseudotanais
Pseudotanais
falciculata Bird and Holdich, 1989
Pseudotanais
Pseudotanais
forcipatus (Lilljeborg, 1864)
Pseudotanais
Pseudotanais
inflatus Kudinova-Pasternak, 1973
Pseudotanais
Pseudotanais
jonesi Sieg, 1977
Pseudotanais
Pseudotanais
lilljeborgi, G.O. Sars, 1882
Pseudotanais
Pseudotanais
longisetosus Sieg, 1977
Pseudotanais
Pseudotanais
longispinus Bird and Holdich, 1989
Pseudotanais
Pseudotanais
macrochelis G.O. Sars, 1882
Pseudotanais
Pseudotanais
mediterraneus G.O. Sars, 1882
73
Pseudotanais {Pseudotanais) mexikolpos Sieg and Heard, 1988
Pseudotanais {Pseudotanais) nipponicus McLelland, 2007
Pseudotanais {Pseudotanais) nordenskioldi Sieg, 1977
Pseudotanais {Pseudotanais) oculatus Hansen, 1913
Pseudotanais {Pseudotanais) scalpellum Bird and Holdich, 1989
Pseudotanais {Pseudotanais) spatula Bird and Holdich, 1989
Pseudotanais {Pseudotanais) spicatus Bird and Holdich, 1989
Pseudotanais {Pseudotanais) unicus Sieg, 1977
Pseudotanais {Pseudotanais) vitjazi Kudinova-Pasternak, 1966
Pseudotanais {Pseudotanais) vulsella Bird and Holdich, 1989
Pseudotanais {Pseudotanais) sp. A, n. sp.
Pseudotanais {Pseudotanais) sp. C, n. sp.
Pseudotanais {Pseudotanais) sp. O, n. sp.
Pseudotanais {Pseudotanais) sp. P, n. sp.
Remarks
With its 27 nominal and four new species, this subgenus is the largest, most diverse taxon
in the family. Found in all oceans at all studied depths, it exhibits a wide variety of
morphological specializations which is evident in the large degree of homoplasy seen in the
phylogenetic analysis.
Pseudotanais {Pseudotanais) abyssi Hansen, 1913
Synonymy
Pseudotanais abyssi Hansen, 1913:25
Diagnosis
74
As for the family, subfamily and subgenus except: Cephalothorax longer than pereonites
1 to 3. Eyelobes absent. Antennuie first article 60 percent of total antennule length, about 6.0
times longer than broad; distal article about 4.0 times longer than broad. Antenna 92 percent
length of antennule; second article subequal in length to third article, both with long slender
spiniform setae; fourth article 7.0 times longer than broad and fifth article about 4.0 times longer
than broad. Mandible pars molaris narrow and simple pointed. Maxillule endite terminating with
nine spiniform setae. Maxilliped bases without seta near palp articulation; endites fused except
for marginal notch, each bearing pair of simple setae. Cheliped with strongly forcipate chelae;
carpus 2.0 times longer than broad, with three inferior setae; propodus nearly 3.0 times longer
than broad, fixed finger lacking inferior setae; dactylus about 60 percent of propodus length,
subequal in width to fixed finger and possessing a proximal seta. Pereopod 1 basis 9.0 times
longer than broad, with simple and broom setae; ischium, merus and carpus each with one seta;
propodus 6.0 times longer than broad, with one seta; dactylus with short seta, length with unguis
subequal to propodus length. Pereopods 2 and 3 basis with simple and broom setae; ischium with
one seta, merus with two setae, carpus with blade seta less than half length of propodus and
dactylus with unguis 2/3 length of propodus. Pereopod 4 basis 6.0 times longer than broad.
Pereopods 4 to 6 basis with broom setae, ischium with one seta, merus with one seta and
propodus lacking broom seta. Pleopods elongate with terminal setae only. Uropod exopods and
endopods each with two articles; exopods 75 percent length of endopods.
Type Locality
North Atlantic: Davis Strait, Arctic Ocean
Depth Range
40-2702 m.
Distribution
75
Known from Davis Strait at the entrance to the Arctic Ocean (type locality) and from
coastal Antarctica below the Indian Ocean.
Remarks
This species is one of only two pseudotanaids with known bi-polar distribution, the other
being Cryptocopoides arcticus. It is distinctive by having strongly forcipate chelae which are
similar to those of Pseudotanais. forcipatus, but differs from that species by possessing pleopods
(missing in P. forcipatus) and having longer, more slender appendages. In addition, P. abyssi
bears a resemblance to P. jonesi but differs first with uropodal exopods extending to about 75
percent the length of the endopods whereas in P. jonesi the exopods are only about half the length
of the endopods, and second by having maxilliped endites which possess a pair of small setae and
are fused except for a marginal notch whereas, in P. jonesi the maxilliped endites are non-setose
and completely fused. Vanhoffen (1914) stated that the Antarctic specimens differed from
specimens from the Arctic population only by the more elongate dactylus-unguis on pereopod 1
but attributed the feature as growth related.
References
Hansen (1913): 25-26, PL II, Figs. 4a-i; Vanhoffen (1914): 483; Sieg (1977): 89-93,
Figs. 65-68.
Pseudotanais {Pseudotanais) affinis Hansen, 1887
Synonymy
Pseudotanais affinis Hansen, 1887: 207-208
Pseudotanais affinis: Sieg (1977): 28, 55-59, (non Hansen, 1887)
Pseudotanais (Pseudotanais) affinis Bird and Holdich (1989): 241-246.
Diagnosis
76
As for the family, subfamily and subgenus except: Cephalothorax subequal to pereonites
1 to 3. Eyelobes absent. Antennule first article 55 percent of total antennule length, about 5.0
times longer than broad; distal article about 5.0 times longer than broad. Antenna same length as
antennule; second article subequal in length to third article, both with short, stout spiniform setae;
fourth article about 8.0 times longer than broad and fifth article 4.5 times longer than broad.
Mandible pars molaris narrow, with four unequal accessory denticles. Maxillule endite
terminating with nine spiniform setae. Maxilliped bases with seta near palp articulation; endites
fused between 1/3 and 2/3 length, each bearing simple seta. Cheliped with non-forcipate chelae;
carpus 2.0 times longer than broad, with two inferior setae; propodus about 2.5 times longer than
broad, fixed finger with one inferior seta; dactylus about 60 percent of propodus length, subequal
in width to fixed finger and possessing proximal seta. Pereopod 1 basis about 12.0 times longer
than broad, with simple seta; ischium and merus each with one seta; carpus with two setae;
propodus about 8.0 times longer than broad, without setae; dactylus with short seta, length with
unguis about 2/3 propodus length. Pereopods 2 and 3 basis with broom setae; ischium with one
seta, merus with two setae, carpus with blade seta about half length of propodus and dactylus with
unguis half length of propodus. Pereopod 4 basis 4.5 times longer than broad. Pereopods 4 to 6
basis with broom setae, ischium with two equal setae, merus with one seta and propodus lacking
broom seta. Pleopods elongate with terminal setae only. Uropod exopods and endopods each
with two articles; exopods 75 percent length of endopods.
Type Locality
Arctic Ocean: Kara Sea north of Russia (lectotype).
Depth Range
40-2970 m.
Distribution
North Atlantic and Arctic Oceans: North Greenland, Kara Sea, Davis Straits,
Norwegian Sea, Faeroe-Shetland Channel.
77
Remarks
Bird and Holdich redescribed Pseudotanais affinis from type material from the Kara Sea
and other material from the Norwegian Sea after finding that Sieg's description was based on a
"variety" that Hansen had included with his specimens. This variety had sufficient distinguishing
characters, in particular the mandible pars molaris structure and antenna setation to justify
establishing a new species under the name P. corollatus. Pseudotanais affinis is distinguished by
its acuminate pars molaris that bears terminal denticles, a feature it shares with six other species
in the "affinis group" (Bird and Holdich, 1989: 241), and the short, stout spiniform setae on the
second and third antenna articles.
References
Hansen (1887): 207-208, PL 21, Fig. 2; Hansen (1913): 30-33; PL III, Figs. 2a-j, 1-m, o;
male?; Hansen (1913) :32-33, Figs. 3a-b 'variety' and 31, 32, Figs . 2k {non Hansen, 1887); Sieg
(1977): 28, 55-59, Figs 39-41, non Hansen, 1887; Bird and Holdich (1989): 241-246, Figs IB,
D, 3-5 (redescription)
Pseudotanais {Pseudotanais) baresnauti Bird, 1999
Diagnosis
As for the family, subfamily and subgenus except: Cephalothorax longer than pereonites
1 to 3. Eyelobes absent. Antennule 93 percent length of antenna, first article 61 percent of total
antennule length, about 4.0 times longer than broad; distal article about 3.0 times longer than
broad. Antenna second article subequal in length to third article, both with short, stout spiniform
setae; fourth and fifth articles each about 4.0 times longer than broad. Mandible pars molaris
narrow with bifid tip. Maxillule endite terminating with nine spiniform setae. Maxilliped bases
with seta near palp articulation; endites fused except for marginal notch, each bearing short
simple seta. Cheliped with non-forcipate chelae; carpus about 1.5 times longer than broad, with
78
two inferior setae; propodus nearly 3.0 times longer than broad, fixed finger with one inferior
seta; dactylus 62 percent of propodus length, subequal in width to fixed finger and possessing
proximal seta. Pereopod 1 basis 8.0 times longer than broad, with simple setae; ischium and
merus each with one seta; carpus with two setae; propodus about 4.5 times longer than broad,
with one seta; dactylus with short seta, length with unguis subequal to propodus length.
Pereopods 2 and 3 basis without setae; ischium with one seta, merus with two setae, carpus with
blade seta less than half length of propodus and dactylus with unguis 2/3 length of propodus.
Pereopod 4 basis about 4.5 times longer than broad. Pereopods 4 to 6 basis with broom setae,
ischium and merus each with one seta and propodus lacking broom seta. Pleopods elongate with
terminal setae only. Uropod exopods and endopods each with two articles; exopods slightly
shorter than endopods.
Type Locality
North Atlantic: Seaward of Barbados Trench, Caribbean Sea.
Depth Range
5000 m.
Distribution
Known only from the type locality in the Caribbean Sea near the Barbados Trench.
Remarks
Bird (1999) compared this species to similarities with Pseudotanais macrocheles and P.
lilljeborgi, both from northern Atlantic waters. Pseudotanais baresnauti differs from the former
by the structure of the uropods (uni-articulate in P. macrocheles) and by having smaller
pereopodal carpal blade-like setae. It differs from P. lilljeborgi by possessing pleopods, having a
longer uropodal exopod and having short, stout spiniform setae distally on the second and third
antenna articles (P. lilljeborgi has short simple setae).
Reference
79
Bird (1999): 449-451, Figs. 1 and 2.
Pseudotanais {Pseudotanais) borceai (Bacescu, 1960)
Synonymy
Pontonais borceai Bacescu, 1960: 113
Diagnosis
As for the family, subfamily and subgenus except: Cephalothorax longer than pereonites
1 to 3. Eyelobes present, with pigment. Antennule 94 percent length of antenna, first article 55
percent of total antennule length, about 4.0 times longer than broad; distal article about 3.0 times
longer than broad. Antenna second article subequal in length to third article, both with short,
slender simple setae; fourth article 5.0 times longer than broad and fifth article about 3.5 times
longer than broad. Mandible pars molaris narrow, simple pointed. Maxillule endite setation not
described. Maxilliped basis setation not described; endites fused except for marginal notch,
naked. Cheliped with non-forcipate chelae; carpus about 1.3 times longer than broad, with two
inferior setae; propodus about 2.5 times longer than broad, fixed finger with one inferior seta:
dactylus 57 percent of propodus length, narrower than fixed finger and lacking proximal seta.
Pereopod 1 basis length not described, without setae; ischium, merus and carpus without setae;
propodus about 4.5 times longer than broad, without setae; dactylus without setae, length with
unguis distinctly shorter than propodus. Pereopods 2 and 3 basis, ischium and merus setation not
described; carpus with blade seta greater than half length of propodus and dactylus with unguis
about half length of propodus. Pereopods 4 to 6 basis, ischium and merus not described.
Pleopods elongate with terminal setae only. Uropod exopods and endopods each with one article;
exopods 2/3 length of endopods.
Type Locality
Black Sea.
Depth Range
80
60-70 m.
Distribution
Known only from the type locality (Black Sea).
Remarks
Sieg (1977:105) treated this species as incertae sedis because Bacescu's illustrations were
"contradictory" and he had no access to the material. Later, he stated that the type material had
been lost (personal communication from Bacescu) and that "inconsistencies in the original and
only description" led him to exclude P. borceai from his key to the worldwide genera and species
of Pseudotanaidae (Sieg and Heard 1988:44). Bacescu published the original description in
Romanian and the illustrations are either poorly rendered or some appendages were not drawn;
however from the author's illustration of the modified blade-like carpal seta on pereopod 3, it is
clear that this species belongs to the subgenus Pseudotanais s.s.
References
Bacescu (1960): 113-117, Figs. 4 and 5.
Pseudotanais (Pseudotanais) californiensis Dojiri and Sieg, 1997
Diagnosis
As for the family, subfamily and subgenus except: Cephalothorax longer than pereonites
1 to 3. Eyelobes absent. Antennule first article 60 percent of total antennule length, about 6.0
times longer than broad; distal article about 3.0 times longer than broad. Antenna not described.
Mandible pars molaris narrow, simple pointed. Maxillule not described. Maxilliped bases
without seta near palp articulation; endites completely fused, each bearing short simple seta.
Cheliped with strongly forcipate chelae; carpus slightly longer than broad, with two inferior setae;
propodus about 2.0 times longer than broad, fixed finger with one inferior seta; dactylus 58
percent of propodus length, subequal in width to fixed finger and possessing proximal seta.
81
Pereopod 1 basis 5.0 times longer than broad, without setae; ischium and merus each with one
seta; carpus without setae; propodus about 5.0 times longer than broad, with one seta; dactylus
without seta, length with unguis distinctly shorter than propodus. Pereopods 2 and 3 basis not
described; ischium and merus with one seta, carpus with blade seta about half length of propodus
and dactylus with unguis about half length of propodus. Pereopods 4 to 6 not described.
Pleopods elongate with terminal setae only. Uropod exopods and endopods each with one article
bearing pseudo-articulation; exopods about half length of endopods.
Type Locality
North Pacific: Purisima Pt., Santa Maria Basin off California, USA.
Depth Range
90-291 m.
Distribution
Known only from type locality in the Santa Maria Basin off the coast of California.
Remarks
According to the authors, this species is most similar to Pseudotanais jonesi and P. abyssi
because of its forcipate chelae but differs from these by the "strong distal tergal spine" on the
propodus of pereopods 2 and 3. As in the case for other species described by these authors,
illustrations and descriptions of some appendages are lacking or incomplete, thus diminishing to a
degree the accuracy of the phylogenetic analysis conducted on the family as a whole.
Reference
Dojiri and Sieg (1997): 260-264, Figs. 3.30, 3.31.
Pseudotanais {Pseudotanais) colonus Bird and Holdich, 1989a
Diagnosis
82
As for the family, subfamily and subgenus except: Cephalothorax longer than pereonites
I to 3. Eyelobes absent. Antennule first article 60 percent of total antennule length, about 3.5
times longer than broad; distal article about 3.0 times longer than broad. Antenna 88 percent
length of antennule, second article subequal in length to third article, both with short, stout
spiniform setae; fourth article about 5.0 times longer than broad and fifth article about 3.0 times
longer than broad. Mandible pars molaris narrow, simple pointed. Maxillule endite terminating
with nine spiniform setae. Maxilliped bases with seta near palp articulation; endites fused except
for marginal notch, without setae. Cheliped with non-forcipate chelae; carpus about 1.3 times
longer than broad, with two inferior setae; propodus about 2.5 times longer than broad, fixed
finger with one inferior seta; dactylus 60 percent of propodus length, subequal in width to fixed
finger and possessing a proximal seta. Pereopod 1 basis about 6.5 times longer than broad, with
simple setae; ischium with one seta; merus without setae; carpus with two setae; propodus about
5.5 times longer than broad, with two setae; dactylus without seta, length with unguis slightly
longer than propodus. Pereopods 2 and 3 basis with broom setae; ischium with one seta, merus
with two setae, carpus with blade seta about half length of propodus and dactylus with unguis 75
percent length of propodus. Pereopod 4 basis about 5.0 times longer than broad. Pereopods 4 to
6 basis with broom setae, ischium with one seta, merus with two setae, and propodus lacking
broom seta. Pleopods elongate with terminal setae only. Uropod exopods with one article, 1/3
length of endopod; endopods with two articles.
Type Locality
North Atlantic: Meriadzeck Terrace, northern Bay of Biscay.
Depth Range
2175 m.
Distribution
Known only from the type locality in the northern Bay of Biscay.
83
Remarks
Specimens of Pseudotanais colonus were collected as part of a deep sea re-colonization
experiment from enriched artificial sediment suspended at depth. Several unidentified swimming
males were also present. This species was deemed most similar to P. mediterraneus and P. unicus
by virtue of its simple acute molar process and non-forcipate chelae but differs by having shorter
antennae and a short, uni-articulate uropodal exopod.
Reference
Bird and Holdich (1989a): 310-313, Figs. 2 and 3.
Pseudotanais {Pseudotanais) corollatus Bird and Holdich, 1989
Synonymy
Pseudotanais ajfinis Hansen, 1887: Sieg (1977 - in part)
Diagnosis
As for the family, subfamily and subgenus except: Cephalothorax subequal to pereonites
1 to 3. Eyelobes absent. Antennule first article 49 percent of total antennule length, about 3.0
times longer than broad; distal article about 5.5 times longer than broad. Antenna 91 percent
length of antennule, second article shorter than third article, both with short, slender spiniform
setae; fourth article 6.0 times longer than broad and fifth article about 4.0 times longer than
broad. Mandible pars molaris broad with numerous accessory denticles and two short marginal
setae. Maxillule endite terminating with nine spiniform setae. Maxilliped bases without seta near
palp articulation; endites fused except for marginal notch, naked. Cheliped with non-forcipate
chelae; carpus about 2.0 times longer than broad, with three inferior setae; propodus about 3.0
times longer than broad, fixed finger with one inferior seta; dactylus 54 percent of propodus
length, subequal in width to fixed finger and possessing a proximal seta. Pereopod 1 basis 5.5
times longer than broad, without setae; ischium, merus and carpus each with one seta; propodus
84
about 6.5 times longer than broad, with one seta; dactylus without seta, length with unguis
distinctly shorter than propodus. Pereopods 2 and 3 basis without setae; ischium with one seta,
merus with two setae, carpus with blade seta greater than half length of propodus and dactylus
with unguis 2/3 length of propodus. Pereopod 4 basis 5.0 times longer than broad. Pereopods 4
to 6 basis without setae, ischium with two equal setae, merus with two setae and propodus lacking
broom seta. Pleopods elongate with terminal setae only. Uropod exopods and endopods each
with two articles; exopods 75 percent length of endopods.
Type Locality
Northwest Atlantic: south of Davis Strait, 'Ingolf Station 25.
Depth Range
1064 m.
Distribution
Recorded only from the type locality in the Northwest Atlantic.
Remarks
Sieg (1977) originally described this species as Pseudotanais (P.) qffinis but it was shown
by Bird and Holdich (1989) that the 'variety' Sieg described was in fact a distinct species (see
remarks under P. affmis). In addition to the differences in pars molaris and antenna setation
mentioned earlier, another immediate difference between P. affmis and P. corrolatus is that the
inferior distal seta of pereopods 2 and 3 are much shorter than dactyus with unguis on the latter
whereas in P. qffinis this terminal seta is subequal to the tip of the unguis. According to Bird and
Holdich (1989), P. corrolatus is most similar to P. denticulatus with differentiation shown in
pereonite 2 length, pereopod 1 propodus length, and setation on antenna artices 2 and 3 and the
merus of pereopods 2 to 6.
References
85
Bird and Holdich (1989): 258 (nam. nov.); Hansen (1913): 31-31, Figs. 2k,n - as P.
affinis; Sieg (1977): 28, 55-59, figs. 17, 39-41, 58 (non Hansen 1887) - as P. affmis.
Pseudotanais (Pseudotanais) denticulalus Bird and Holdich, 1989
Synonymy
Pseudotanais affinis Holdich and Bird (1985): 445, non Hansen 1887.
Diagnosis
As for the family, subfamily and subgenus except: Cephalothorax longer than pereonites
1 to 3. Eyelobes absent. Antennule first article 60 percent of total antennule length, about 6.0
times longer than broad; distal article about 5.0 times longer than broad. Antenna about same
length as antennule, second article subequal to third article, both with short, stout spiniform setae;
fourth article 9.0 times longer than broad and fifth article about 4.0 times longer than broad.
Mandible pars molaris broad with numerous accessory denticles and three short marginal setae.
Maxillule endite terminating with nine spiniform setae. Maxilliped bases with two medially
located setae; endites fused between 1/3 and 2/3 length, each with one seta. Cheliped with nonforcipate chelae; carpus 1.5 times longer than broad, with two inferior setae; propodus about 2.5
times longer than broad, fixed finger with one inferior seta; dactylus 61 percent of propodus
length, narrower than fixed finger and possessing proximal seta. Pereopod 1 basis about 6.0
times longer than broad, with simple setae; ischium without setae; merus with one seta and carpus
with four setae; propodus about 6.0 times longer than broad, with one seta; dactylus without seta,
length with unguis subequal in length to propodus. Pereopods 2 and 3 basis with simple setae;
ischium with one seta, merus with two setae, carpus with blade seta about half length of propodus
and dactylus with unguis 2/3 length of propodus. Pereopod 4 basis about 3.5 times longer than
broad. Pereopods 4 to 6 basis with broom setae, ischium with two equal setae, merus with two
86
setae and propodus with broom seta. Pieopods elongate with terminal setae only. Uropod
exopods and endopods each with two articles; exopods 75 percent length of endopods.
Type Locality
Northeast Atlantic: Porcupine Seabight, I.O.S. station 9754/3.
Depth Range
1484 m.
Distribution
Northeast Atlantic Ocean.
Remarks
Bird and Holdich (1989) considered this species to be the most closely allied to
Pseudotanais corollatus and found it to be the most common and abundant tanaid in their
Northeast Atlantic study. In addition to differences between P. denticulatus and P. corollatus
mentioned earlier, P. denticulatus can also be distinguished by overal size, the lengths of the
cephalothorax and pereonites 1 to 3, and slightly different terminal denticulation on the
mandibular pars molaris.
Reference
Bird and Holdich (1989): 258, Figs. IF, 13-14.
Pseudotanais (Pseudotanais) falciculata Bird and Holdich, 1989
Synonymy
Pseudotanais abyssi (Holdich and Bird, 1985): 445, non Hansen, 1913.
Diagnosis
As for the family, subfamily and subgenus except: Cephalothorax shorter than pereonites
1 to 3. Eyelobes absent. Antennule 90 percent length of antenna, first article 62 percent of total
87
antennule length, about 6.0 times longer than broad; distal article about 4.0 times longer than
broad. Antenna second article subequal to third article, both with long, slender spiniform setae;
fourth article about 6.0 times longer than broad and fifth article about 4.0 times longer than broad.
Mandible pars molaris narrow with small denticles along one margin. Maxillule endite
terminating with nine spiniform setae. Maxilliped bases with seta near each palp articulation;
endites fused except for notch on distal margin, each with two short cusps. Cheliped with
strongly forcipate chelae; carpus 1.3 times longer than broad, with two inferior setae; propodus
about 2.5 times longer than broad, fixed finger with one inferior seta; dactylus 51 percent of
propodus length, subequal in width to fixed finger and possessing proximal seta. Pereopod 1
basis about 10.0 times longer than broad, with broom setae; ischium, merus and carpus each with
one seta; propodus about 5.0 times longer than broad, with two setae; dactylus with short seta,
length with unguis distinctly shorter than propodus. Pereopods 2 and 3 basis without setae;
ischium with one seta, merus with two setae, carpus with blade-like seta less than half length of
propodus and dactylus with unguis about half length of propodus. Pereopod 4 basis about 5.5
times longer than broad. Pereopods 4 to 6 basis with broom setae, ischium with two unequal
setae, merus with two setae and propodus with broom seta. Pleopods elongate with terminal setae
only. Uropod exopods and endopods each with two articles; exopods subequal to length of
endopods.
Type Locality
Northeast Atlantic: Rockall Trough, Scottish Marine Biological Association, Permanent
Station no. 540, west of Ireland.
Depth Range
2719-4829m.
Distribution
Northeast Atlantic Ocean, at type locality and Bay of Biscay Abyssal Plains.
88
Remarks
This species is distinguished, according to Bird and Holdich (1989: 277), by its short
dactylus and fixed finger of the cheliped. It is most similar to Pseudtotanais vulsella, another
forcipate-chela species, differring, not only by the shorter cheliped fingers, but by having smaller
cusps on the maxilliped endites and shorter carpal blade-like setae on pereopods 2 to 6.
Reference
Bird and Holdich (1989): 273-277, Figs. 19 and 20.
Pseudotanais (Pseudotanais) forcipatus (Lilljeborg, 1864)
Synonymy
Tanais forcipatus Lilljeborg, 1864: 16
Paratanais forcipatus Meinert, 1877: 87
Pseudotanais forcipatus Sars, 1882: 46
Pseudotanais lilljeborgi (non Sars) Hansen, 1913: 28
Diagnosis
As for the family, subfamily and subgenus except: Cephalothorax longer than pereonites
1 to 3. Eyelobes absent. Antennule first article 64 percent of total antennule length, about 5.5
times longer than broad; distal article about 3.5 times longer than broad. Antenna about same
length as antennule, second article subequal to third article, both with short, slender spiniform
setae; fourth article about 4.5 times longer than broad and fifth article about 2.5 times longer than
broad. Mandible pars molaris narrow, simple pointed. Maxillule endite terminating with nine
spiniform setae. Maxilliped bases without seta near each palp articulation; endites completely
fused, naked. Cheliped with strongly forcipate chelae; carpus 1.4 times longer than broad, with
two inferior setae; propodus about 2.5 times longer than broad, fixed finger with one inferior seta;
dactylus 57 percent of propodus length, subequal in width to fixed finger and possessing proximal
89
seta. Pereopod 1 basis about 7.5 times longer than broad, with simple setae; ischium and inerus
each with one seta and carpus without setae; propodus about 5.5 times longer than broad, with
one seta; dactylus without setae, length with unguis distinctly shorter than propodus. Pereopods 2
and 3 basis with broom and simple setae; ischium with one seta, merus with two setae, carpus
with blade-like seta less than half length of propodus and dactylus with unguis about 75 percent
length of propodus. Pereopod 4 basis about 4.5 times longer than broad. Pereopods 4 to 6 basis
with broom setae, ischium with one unequal seta, merus with two setae and propodus without
broom seta. Pleopods absent. Uropod exopods and endopods each with one pseudo-articulated
article; exopods 75 percent length of endopods.
Type Locality
Northeast Atlantic: Coast of Norway, off Molde
Depth Range
15-375 m.
Distribution
Widespread in the North Sea and Arctic Ocean off Britain, Norway, Greenland and in the
North Pacific south of Alaska.
Remarks
The lack of pleopods and the uni-articulate uropods sets this species apart from all others
with forcipate chelae.
Reference
Sieg(1977): 97-105, Figs. 72-79.
Pseudotanais {Pseudotanais) inflatus Kudinova-Pasternak, 1973
Diagnosis
90
As for the family, subfamily and subgenus except: Cephaiothorax longer than pereonites
1 to 3. Eyelobes absent. Antennule 70 percent length of antenna, first article 60 percent of total
antennule length, about 8.0 times longer than broad; distal article about 10.0 times longer than
broad. Antenna second article shorter than third article, both with short, slender simple setae;
fourth article about 11.0 times longer than broad and fifth article about 5.0 times longer than
broad. Mandible pars molaris broad, truncate, with numerous accessory denticles. Maxillule
endite terminating with seven spiniform setae. Maxilliped bases without seta near each palp
articulation; endites completely separate, each with two setae and two short cusps. Cheliped with
weakly forcipate chelae; carpus 1.8 times longer than broad, with two inferior setae; propodus
about 4.0 times longer than broad, fixed finger without inferior seta; dactylus 60 percent of
propodus length, subequal in width to fixed finger and possessing proximal seta. Pereopod 1
basis 10.0 times longer than broad, without setae; ischium with one seta; merus without setae;
carpus with one seta; propodus about 6.0 times longer than broad, without setae; dactylus without
setae, length with unguis subequal to length of propodus. Pereopods 2 and 3 basis, ischium and
merus not described; carpus with blade-like seta less than half length of propodus and dactylus
with unguis distinctly less than half length of propodus. Pereopod 4 basis about 4.0 times longer
than broad. Pereopods 4 to 6 setation not described. Pleopods elongate, with terminal setae only.
Uropod exopods and endopods each with two articles; exopods subequal in length to endopods.
Type Locality
North Pacific: Aleutian Trench off Alaska.
Depth Range
3610 m.
Distribution
Known only from type locality, Aleutian Trench off Alaska in the North Pacific.
Remarks
91
This species was poorly described and illustrated by Kudinova-Pasternak. Most pereopod
setal details were omitted from the above diagnosis, and from the phyogenetic analysis, because
their number and structure is unclear. Further inconsistencies include maxilliped endites that are
illustrated as completely separate and a maxillule endite with seven terminal spiniform setae
rather than the usual nine. Sieg (1977:59) synonomized Pseudotanais inflatus with P. affinis
primarily because of its similar pars molaris, broad with numerous accessory denticles, as
illustrated by Kudinova-Pasternak. This synonymy was shown to be invalid by Bird and Holdich
(1989) in their redescription of P. afflnis and description of/*, corollatus (see remarks under these
two species). These authors (1989: 263) suggest that P. inflatus, if found to be valid upon reexamination, should be grouped with P. denticulatus and P. corollatus based on the structure of
the pars molaris. The placement of this species in Pseudotanais is legitimate owing to the
apparent presence of blade-like carpal setae, but until further material can be examined its
validity as a species should probably remain questionable.
Reference
Kudinova-Pasternak (1973): 164-166, Fig. 14.
Pseudotanais {Pseudotanais) jonesi Sieg, 1977
Diagnosis
As for the family, subfamily and subgenus except: Cephalothorax longer than pereonites
1 to 3. Eyelobes absent. Antennule first article 55 percent of total antennule length, about 5.0
times longer than broad; distal article about 3.0 times longer than broad. Antenna 95 percent
length of antennule, second article subequal in length to third article, both with short, slender
spiniform setae; fourth article about 5.0 times longer than broad and fifth article about fout times
longer than broad. Mandible pars molaris narrow, simple pointed. Maxillule endite terminating
with nine spiniform setae. Maxilliped bases without seta near palp articulation; endites
completely fused, naked. Cheliped with strongly forcipate chelae; carpus about 1.5 times longer
than broad, with two inferior setae; propodus about 2.5 times longer than broad, fixed finger
without inferior seta; dactylus 68 percent of propodus length, broader than fixed finger and
possessing proximal seta. Pereopod 1 basis about 6.0 times longer than broad, with simple setae;
ischium with one seta; merus with two setae; carpus without setae; propodus about 5.5 times
longer than broad, with one seta; dactylus without setae, length with unguis distinctly shorter than
propodus. Pereopods 2 and 3 basis with broom setae, ischium and merus with one seta, carpus
with blade seta about half length of propodus and dactylus with unguis subequal to length of
propodus. Pereopod 4 basis about 4.5 times longer than broad. Pereopods 4 to 6 basis with
broom setae, ischium and merus with one seta, and propodus lacking broom seta. Pleopods
elongate with terminal setae only. Uropod exopods and endopods each with two articles;
exopods half length of endopods.
Type Locality
North Atlantic: off Plymouth, England (UK).
Depth Range
50 m.
Distribution
Known only from the type locality near the Isle of Man off Plymouth, England, UK.
Remarks
This Northeast Atlantic species is distinguished from other forcipate-chelae species by its
short uropodal exopod which is only half the length of the endopod, its non-setose, completely
fused maxilliped endites, and by its lack of an inferior seta on the cheliped fixed finger. In
addition, it is the only species in the family in which the cheliped dactylus is broader, if only
slightly, than the fixed finger.
References
93
Sieg (1977): 94-97, Figs. 69-71, 79.
Pseudotanais {Pseudotanais) lilljeborgi, G.O. Sars, 1882
Diagnosis
As for the family, subfamily and subgenus except: Cephalothorax longer than pereonites
1 to 3. Eyelobes rudimentary, without pigment. Antennule first article 60 percent of total
antennule length, about 5.5 times longer than broad; distal article about 3.0 times longer than
broad. Antenna about same length as antennule, second article shorter than third article, both
with short, slender spiniform setae; fourth article about 4.5 times longer than broad and fifth
article about 3.0 times longer than broad. Mandible pars molaris narrow, with three blunt
terminal teeth. Maxillule endite terminating with nine spiniform setae. Maxilliped bases with
seta near palp articulation; endites fused except for marginal notch, each with short seta.
Cheliped with non-forcipate chelae; carpus about 1.3 times longer than broad, with two inferior
setae; propodus about 3.0 times longer than broad, fixed finger with one inferior seta; dactylus 51
percent of propodus length, subequal in width to fixed finger and possessing proximal seta.
Pereopod 1 basis about 5.0 times longer than broad, with simple setae; ischium and merus with
one seta; carpus with two setae; propodus about 4.5 times longer than broad, with one seta;
dactylus without setae, length with unguis distinctly longer than propodus. Pereopods 2 and 3
basis with simple setae; ischium with one seta, merus with two setae, carpus with blade seta less
than half length of propodus and dactylus with unguis 75 percent length of propodus. Pereopod 4
basis about 4.0 times longer than broad. Pereopods 4 to 6 basis without setae, ischium with one
seta, merus with two setae, and propodus lacking broom seta. Pleopods absent. Uropod exopods
and exopods each with two articles; exopods half length of endopods.
Type Locality
North Atlantic: Varanger Fjord at Vadso off northern Norway (Barent's Sea).
94
Depth Range
7-536 m.
Distribution
Known from Several locations off Iceland, Greenland, Norway, Denmark and the Barents
Sea.
Remarks
This species is one of two species of Pseudotanais, the other being P.forcipatus that
completely lack pleopods. It differs from P.forcipatus primarly by having non-forcipate chelae
but also by having complete articulation in the uropods and a more complex mandible pars
molaris. Except for the lack of pleopods it is also similar to P. baresnauti (see remarks for that
species).
References
Sieg (1977): 72-76, Figs. 52-54, 68; Sars (1889), 40-41, PL. XVII, Fig. 2.
Pseudotanais {Pseudotanais) longisetosus Sieg, 1977
Diagnosis
As for the family, subfamily and subgenus except: Cephalothorax longer than pereonites
1 to 3. Eyelobes absent. Antennule first article 55 percent of total antennule length, about 4.5
times longer than broad; distal article about 4.0 times longer than broad. Antenna 94 percent
length as antennule, second article shorter than third article, both with short, slender spiniform
setae; fourth article 5.0 times longer than broad and fifth article 3.5 times longer than broad.
Mandible pars molaris narrow with terminal spine-like tooth and six subterminal denticles.
Maxillule endite terminating with nine spiniform setae. Maxilliped bases with seta near each palp
articulation; endites fused exept for notch on distal margin, each with one short seta. Cheliped
with non-forcipate chelae; carpus 1.7 times longer than broad, with two inferior setae; propodus
95
about 3.0 times longer than broad, fixed finger with one inferior seta; dactyl us 49 percent of
propodus length, subequal in width to fixed finger and possessing proximal seta. Pereopod 1
basis about 6.5 times longer than broad, with simple setae; ischium, merus and carpus each with
one seta; propodus about 7.0 times longer than broad, without setae; dactylus without setae,
length with unguis subequal in length to propodus. Pereopods 2 and 3 basis with broom and
simple setae; ischium with one seta, merus with two setae, carpus with blade seta greater than half
length of propodus and dactylus with unguis subequal to length of propodus. Pereopod 4 basis
about 4.5 times longer than broad. Pereopods 4 to 6 basis with simple and broom setae, ischium
with two equal setae, merus with one seta, carpus with additional exceptionally long superior seta
and propodus without broom seta. Pleopods elongate with terminal setae only. Uropod exopods
with one article, 2/3 length of endopods; endopods with two articles.
Type Locality
Indian Ocean: North of Cumberland Bay, South Georgia off Antarctica (Swedish South
Pole Expedition, station 34).
Depth Range
497-6150 m.
Distribution
Known only from type locality off Antarctica.
Remarks
This Antarctic species resembles Pseudotanais nordenskioldi with its similar mandibular
molar process but is distinguished from that species by having an extremely long carpal superior
seta on pereopods 4 to 6 which is equal to or exceeds the length of the propodus and uni-articulate
uropod exopods. It is also apparently closely related to P. longispinus from the Northeast
Atlantic, bearing similarities in pars molaris stucture and the aforementioned carpal superior
setae, but differing, among other features, by the short slender spiniform setae on the second and
96
third antenna articles (short and stout in P. longispinus), and also by having uni-articulate uropod
exopods.
Reference
Sieg (1977): 64-68, Figs. 45-47, 58.
Pseudotanais {Pseudotanais) longispinus Bird and Holdich, 1989
Synonymy
Pseudotanais longisetosus, Holdich and Bird (1985).
Diagnosis
As for the family, subfamily and subgenus except: Cephalothorax subequal in length to
pereonites 1 to 3. Eyelobes absent. Antennule first article 57 percent of total antennule length,
about 5.0 times longer than broad; distal article about 5.0 times longer than broad. Antenna 97
percent length as antennule, second article subequal in length to third article, both with short,
stout spiniform setae; fourth article 11.0 times longer than broad and fifth article 4.0 times longer
than broad. Mandible pars molaris narrow with terminal setulate spine-like tooth and about eight
subterminal denticles. Maxillule endite terminating with nine spiniform setae. Maxilliped bases
with two medially located setae; endites fused between 1/3 and 2/3 length, each with one strong
seta and two short cusps. Cheliped with non-forcipate chelae; carpus 1.7 times longer than broad,
with two inferior setae; propodus 3.5 times longer than broad, fixed finger with one inferior seta;
dactylus 49 percent of propodus length, narrower than fixed finger and possessing proximal seta.
Pereopod 1 basis about 8.0 times longer than broad, with numerous simple setae; ischium and
merus each with one seta and carpus with two setae; propodus about 11.0 times longer than
broad, with two setae; dactylus with short seta, length with unguis subequal in length to propodus.
Pereopods 2 and 3 basis with broom and simple setae; ischium with one seta, merus with two
setae, carpus with blade seta greater than half length of propodus and dactylus with unguis 75
97
percent length of propodus. Pereopod 4 basis about 5.5 times longer than broad. Pereopods 4 to
6 basis with simple and broom setae, ischium with two equal setae, merus with one seta, carpus
with additional exceptionally long superior seta and propodus with broom seta. Pleopods
elongate with terminal setae only. Uropod exopods and endopods each with two articles;
exopods subequal to length of endopods.
Type Locality
Northeast Atlantic: Northern Bay of Biscay.
Depth Range
2644-4829 m.
Distribution
Rockall Trough, Porcupine Seabight, northern Bay of Biscay (type locality), and the
Porcupine and Biscay Abyssal Plains, all in the Northeast Atlantic.
Remarks
Since this species was deemed an important member of the "affinis group", its description
by Bird and Holdich (1989) emphasized only those characters differing from Pseudotanais
affinis. The authors suggested that its similarity in pars molaris structure allies it closely to the
Antarctic species, P. longisetosus and P. nordenskioldi, however its somewhat more complex
(setose) pars molaris, as well as its deeper distribution, indicates that it is possibly the most
plesiomorphic (ancient) member of that group. The long superior carpal seta of periopods 5 and
6 is a feature shared with P. longisetosus as well as with P. spatula.
Reference
Bird and Holdich (1989): 254-255, Figs. 10 and 11.
Pseudotanais (Pseudotanais) macrochelis G.O. Sars, 1882
Diagnosis
98
As for the family, subfamily and subgenus except: Cephalothorax longer than pereonites
1 to 3. Eyelobes absent. Antennuie 95 percent length of antenna, first article 52 percent of total
antennule length, about 4.0 times longer than broad; distal article about 5.0 times longer than
broad. Antenna second article shorter than third article, both with short, stout spiniform setae;
fourth article 6.5 times longer than broad and fifth article 3.5 times longer than broad. Mandible
pars molaris narrow with bifid tip. Maxillule endite terminating with nine spiniform setae.
Maxilliped bases without setae near palp articulation; endites fused between 1/3 and 2/3 length,
each with one short seta. Cheliped with non-forcipate chelae; carpus 2.0 times longer than broad,
with three inferior setae; propodus 3.5 times longer than broad, fixed finger with one inferior seta;
dactylus 57 percent of propodus length, narrower than fixed finger and possessing proximal seta.
Pereopod 1 basis about 6.0 times longer than broad, with simple seta; ischium and merus each
with one seta and carpus with two setae; propodus about 7.5 times longer than broad, without
setae; dactylus without setae, length with unguis distinctly shorter than propodus. Pereopods 2
and 3 basis with simple setae; ischium and merus each with one seta, carpus with blade seta
greater than half length of propodus and dactylus with unguis half length of propodus. Pereopod
4 basis about 5.0 times longer than broad, with simple seta. Pereopods 4 to 6 ischium with two
equal setae, merus with two setae and propodus without broom seta. Pleopods elongate with
terminal setae only. Uropod exopods and endopods each with two articles; exopods half length of
endopods.
Type Locality
North Atlantic: coast of Norway.
Depth Range
110-135 m.
Distribution
Known only from off the coast of Norway: Verangerfjord, Bergen and Raunefjorden.
99
Remarks
This species bears resemblance to Pseudolanais baresnauti by its acuminate pars molaris
with bifid tips but differs primarily by its larger carpal blade-like spiniform setae on pereopods 2
and 3 which extend for more than half the length of the propodus.
References
Sieg (1977): 76-81, Figs. 55-58.
Pseudolanais {Pseudolanais) medilerraneus G.O. Sars, 1882
Diagnosis
As for the family, subfamily and subgenus except: Cephalothorax longer than pereonites
1 to 3. Eyelobes present, with pigment. Antennule first article 55 percent of total antennule
length, about 4.0 times longer than broad; distal article about 5.5 times longer than broad.
Antenna 95 percent length of antennule, second article subequal in length to third article, both
with short, slender spiniform setae; fourth article 6.0 times longer than broad and fifth article 4.5
times longer than broad. Mandible pars molaris narrow, simple pointed. Maxillule endite
terminating with nine spiniform setae. Maxilliped bases without setae near palp articulation;
endites fused except for notch on distal margin, naked. Cheliped with non-forcipate chelae;
carpus 1.5 times longer than broad, with two inferior setae; propodus 2.5 times longer than broad,
fixed finger with one inferior seta; dactylus 53 percent of propodus length, subequal in width to
fixed finger and possessing proximal seta. Pereopod 1 basis about 7.5 times longer than broad,
with simple seta; ischium and merus with two setae; carpus with one seta; propodus about 6.0
times longer than broad, with seta; dactylus without seta, length with unguis subequal to
propodus. Pereopods 2 and 3 basis without setae; ischium with two equal setae, merus with two
setae, carpus with blade seta less than half length of propodus and dactylus with unguis subequal
to length of propodus. Pereopod 4 basis about 3.5 times longer than broad. Pereopods 4 to 6
100
basis with simple and broom setae, ischium with two equal setae, merus with two setae and
propodus without broom seta. Pleopods elongate with terminal setae only. Uropod exopods and
endopods each with two articles; exopods subequal to length of endopods.
Type Locality
Mediterranean Sea: Off the coast of Spezia, Northwest Itally.
Depth Range
Not published.
Distribution
Mediterranean Sea off the coast of Italy (type locality).
Remarks
This oculate species is only one of two pseudotanaids known from the Mediterranean, the
other being another oculate species, Pseudotanais unicus from which it differs by having a wider
cephalothorax and a longer third pereopod. It is also similar to the oculate P. mexikolpos from the
Gulf of Mexico differing mainly by the length of the superior terminal seta on the propodus of
pereopods 4 to 6 which is about as long as the claw on P. mediterraneous and much longer than
the claw on P. mexikolpos. Pseudotanais mediterraneus is unique among members of the
subfamily Pseudotanainae for having two ischium setae on the first pereopod, all others have one
or none.
Reference
Sieg (1977): 81-85, Figs. 59-61, 68.
Pseudotanais {Pseudotanais) mexikolpos Sieg and Heard, 1988
Synonymy
Paratanais sp. A, anonymous (Texas A&M University), 1978: 772.
Diagnosis
101
As for the family, subfamily and subgenus except: Cephalothorax longer than pereonites
I to 3. Eyelobes present, with pigment. Antennule first article 56 percent of total antennule
length, about 4.5 times longer than broad; distal article about 4.0 times longer than broad.
Antenna about same length as antennule, second article shorter than third article, both with short,
slender spiniform setae; fourth article about 4.5 times longer than broad and fifth article 3.0 times
longer than broad. Mandible pars molaris narrow, simple pointed. Maxillule endite terminating
with nine spiniform setae. Maxilliped bases with seta near each palp articulation; endites
completely fused, each with four setae in two rows. Cheliped with weakly-foreipate chelae;
carpus 1.7 times longer than broad, with two inferior setae; propodus about 3.0 times longer than
broad, fixed finger with one inferior seta; dactylus 54 percent of propodus length, subequal in
width to fixed finger and possessing proximal seta. Pereopod 1 basis about 5.0 times longer than
broad, without setae; ischium, merus and carpus each with one seta; propodus about 5.0 times
longer than broad, with one seta; dactylus without setae, length with unguis subequal in length to
propodus. Pereopods 2 and 3 basis with broom setae; ischium with one seta, merus with two
setae, carpus with blade seta greater than half length of propodus and dactylus with unguis half
length of propodus. Pereopod 4 basis about 4.0 times longer than broad. Pereopods 4 to 6 basis
with simple and broom setae, ischium with two equal setae, merus with two setae and propodus
without broom seta. Pleopods elongate with terminal setae only. Uropod exopods and endopods
each with two articles; exopods subequal to length of endopods.
Type Locality
North Atlantic: Gulf of Mexico, East Flower Garden Bank off Texas Coast.
Depth Range
72 m.
Distribution
102
Known only from the type locality and adjacent locations off the Texas coast in the Gulf
of Mexico.
Remarks
This species appears to be very similar to two Mediterannean species, Pseudotanais
unicus and P. mediterraneus. It is distinguished from the former by having a shorter
cephalothorax and longer periopod 3, and from the latter by a much longer superio-terminal
spiniform seta on the propodus of pereopods 4 to 6 which extends far beyond the claw. All three
species are oculate, from relatively shallow depths and have similar mouthparts with acuminate
pars molaris; however P. mexikolpos has narrowly forcipate chelae whereas the other two have
chelae which are non-forcipate.
Reference
Sieg and Heard (1988): 48-57, Figs. 8-12.
Pseudotanais {Pseudotanais) nipponicus McLelland, 2007
Diagnosis
As for the family, subfamily and subgenus except: Cephalothorax shorter than pereonites
1 to 3. Eyelobes absent. Antennule first article 56 percent of total antennule length, about 5.5
times longer than broad; distal article about 5.0 times longer than broad. Antenna 96 percent
length of antennule, second article subequal in length to third article, both with short, stout
spiniform setae; fourth article about 10.0 times longer than broad and fifth article about 4.0 times
longer than broad. Mandible pars molaris narrow, with four to five small subdistal denticles.
Maxillule endite terminating with nine spiniform setae. Maxilliped bases with small seta near
each palp articulation and two longer medially located setae; endites fused between 1/3 and 2/3
length, each with one short seta and two short cusps. Cheliped with non-forcipate chelae; carpus
about 2.0 times longer than broad, with two inferior setae; propodus 3.7 times longer than broad,
103
fixed finger with one inferior seta; dactylus 61 percent of propodus length, narrower in width to
fixed finger and possessing proximal seta. Pereopod 1 basis about 7.5 times longer than broad,
with numerous marginal simple setae; ischium and merus each with one seta; carpus with two
setae; propodus about 7.5 times longer than broad, with one seta; dactylus without setae, length
with unguis subequal in length to propodus. Pereopods 2 and 3 basis with broom and simple
setae; ischium with one seta, merus with two setae, carpus with blade seta greater than half length
of propodus and dactylus with unguis half length of propodus. Pereopod 4 basis about 4.5 times
longer than broad. Pereopods 4 to 6 basis with simple and broom setae, ischium with two
unequal setae, merus with two setae, carpus with additional exceptionally long superior seta and
propodus with broom seta. Pleopods elongate with terminal setae only. Uropod exopods and
endopods each with two articles; exopods subequal to length of endopods.
Type Locality
North Pacific: Kamchatka Trench north of Japan.
Depth Range
3145-3858m.
Distribution
Known only from the type locality in the Kamchatka Trench, North Pacific Ocean.
Remarks
This species appears to be in the group with Pseudotanais longisetosus, P. longispinus,
and P. nordenskioldi as mentioned by Bird and Holdich (1989). In fact, it is nearly indentical with
their description of P. longispinus except for minor differences in the setation of the mandible
pars molaris, the presence of dorsal and lateral setae on the thoracic and abdominal segments, a
larger spiniform seta on antenna second article, and unequal merus and carpal superior setae on
pereopod 1 (equal lengths in P. longispinus). Both species have a pars molaris with subterminal
teeth except that P. nipponicus apparently has less setation with only 4-5 small subterminal teeth
104
whereas P. longispinus has "one long and about eight short terminal denticles" the longest of
which is 4 times the length of the others and has secondary spinules. Additionally, the distal half
of the long terminal setae on the propodus of pereopods 4-6 are heavily setulose on P.
nipponicus, a feature either absent in P. longispinus or not mentioned by Bird and Holdich, and a
few of the spiniform setae of the maxillule endite are equipped with a subterminal setule, a
feature also not observed in P. longispinus. The setation of the pereopods is nearly identical in
both species, differing from P. nordenskioldi by having a long superior seta on the carpus of
pereopods 4 and 5. The two species appear to have identical maxilliped armature which differs
from that of P. longisetosus by having two small cusps on the endites.
Reference
McLelland (2007): 93-98, Figs. 4-6.
Pseudotanais (Pseudotanais) nordenskioldi Sieg, 1977
Diagnosis
As for the family, subfamily and subgenus except: Cephalothorax longer than pereonites
1 to 3. Eyelobes absent. Antennule first article 56 percent of total antennule length, about 5.0
times longer than broad; distal article about 4.5 times longer than broad. Antenna 92 percent
length of antennule, second article subequal in length to third article, both with short, stout
spiniform setae; fourth article about 4.5 times longer than broad and fifth article about 3.0 times
longer than broad. Mandible pars molaris narrow, with terminal spine-like tooth and six
subterminal denticles. Maxillule endite terminating with nine spiniform setae. Maxilliped bases
with seta near each palp articulation; endites fused exept for notch on distal margin, each with
two setae. Cheliped with non-forcipate chelae; carpus about 1.7 times longer than broad, with
two inferior setae; propodus 3.4 times longer than broad, fixed finger with one inferior seta;
dactylus 58 percent of propodus length, narrower in width to fixed finger and possessing
105
proximal seta. Pereopod 1 basis about 5.5 times longer than broad, with simple seta; ischium
with one seta; merus with three setae; carpus with one seta; propodus about 5.5 times longer than
broad, with one seta; dactylus with short seta, length with unguis subequal in length to propodus.
Pereopods 2 and 3 basis with broom and simple setae; ischium with one seta, merus with two
setae, carpus with blade seta greater than half length of propodus and dactylus with unguis 75
percent length of propodus. Pereopod 4 basis about 4.0 times longer than broad. Pereopods 4 to
6 basis with broom setae, ischium with one seta, merus with two setae and propodus without
broom seta. Pleopods elongate with terminal setae only. Uropod exopods and endopods each
with two articles; exopods subequal to length of endopods.
Type Locality
Indian Ocean: North of Cumberland Bay, South Georgia off Antarctica (Station 34 of the
Swedish South Pole Expedition).
Depth Range
497-6150 m.
Distribution
Antarctica (type locality), Southwest Atlantic (nine sites off Antarctica) and South
America.
Remarks
This species is similar to Pseudotanais longispinus and P. longisetosus by virtue of its
nearly identical pars molaris armature. It differs from both species by lacking the long distal
superior carpal seta on pereopods 4 to 6. See remarks for P. longispinus and P. nipponicus
regarding its relationship with the "affinis group". Pseudotanais nordenskioldii is unique among
the family members by having three pereopod 1 merus setae; all others have two or less.
Reference
Sieg(1977): 60-64, Figs. 42-44, 58.
106
Pseudotanah (Pseudotanais) oculatus Hansen, 1913
Synonymy
Paratanais nanaimoensis Fee, 1926: 17.
Diagnosis
As for the family, subfamily and subgenus except: Cephalothorax subequal to pereonites
1 to 3. Eyelobes present, with pigment. Antennule first article 58 percent of total antennule
length, about 6.5 times longer than broad; distal article about 4.5 times longer than broad.
Antenna 87 percent length of antennule, second article shorter than third article, both with long,
slender spiniform setae; fourth article about 5.5 times longer than broad and fifth article about 2.5
times longer than broad. Mandible pars molaris narrow with with four blunt subterminal
denticles. Maxillule endite terminating with nine spiniform setae. Maxilliped bases with seta
near each palp articulation; endites fused except for marginal notch, each with one short seta.
Cheliped with weakly forcipate chelae; carpus about 1.5 times longer than broad, with two
inferior setae; propodus about 3.0 times longer than broad, fixed finger with one inferior seta;
dactylus 50 percent of propodus length, subequal in width to fixed finger and possessing proximal
seta. Pereopod 1 basis 5.5 times longer than broad, with broom seta; ischium, merus and carpus
each with one seta; propodus about 6.5 times longer than broad, with one seta; dactylus without
seta, length with unguis subequal to propodus. Pereopods 2 and 3 basis with broom setae;
ischium and merus each with one seta, carpus with blade seta less than half length of propodus
and dactylus with unguis 75 percent length of propodus. Pereopod 4 basis about 5.5 times longer
than broad. Pereopods 4 to 6 basis with simple and broom setae, ischium with two equal setae,
merus with two setae and propodus lacking broom seta. Pleopods elongate with terminal setae
only. Uropod exopods and endopods each with two articles; exopods subequal in length to
endopods.
107
Type Locality
North Atlantic: Davis Strait off Western Greenland.
Depth Range
101-123 m.
Distribution
North Atlantic Ocean off Greenland and Iceland; North Pacific off Departure Bay,
Canada.
Remarks
This oculate species is distinguished from other oculates in the subgenus by its more
complex mandibular pars molaris with subterminal denticulations as opposed to the simple
pointed pars molaris in Pseudotanais unicus, P. mexikolpos, and P. mediterraneus. In addition,
the carpal blade-like setae on pereopods 2 and 3 of P. oculatus are shorter in relation to the
propodus than on the other three species.
References
Sieg (1977): 68-72, Figs. 48-51, 68; Hansen (1913): 29-30, PL. II, Figs 6a-d, PL. Ill,
Figs. la-d.
Pseudotanais {Pseudotanais) scalpellum Bird and Holdich, 1989
Diagnosis
As for the family, subfamily and subgenus except: Cephalothorax subequal to pereonites
1 to 3. Eyelobes absent. Antennule first article 53 percent of total antennule length, about 6.5
times longer than broad; distal article about 6.5 times longer than broad. Antenna about same
length as antennule; second article subequal in length to third article, both with short, stout
spiniform setae; fourth article about 11.5 times longer than broad and fifth article 4.5 times longer
than broad. Mandible pars molaris narrow, with two short subterminal denticles and two longer,
108
unequal terminal denticles. Maxillule endite terminating with nine spiniform setae. Maxilliped
bases with seta near palp articulation; endites fused between 1/3 and 2/3 length, each with one
strong seta. Cheliped with non-forcipate chelae; carpus 2.0 times longer than broad, with two
inferior setae; propodus about 3.5 times longer than broad, fixed finger with one inferior seta;
dactylus about 55 percent of propodus length, narrower in width to fixed finger and possessing
proximal seta. Pereopod 1 basis about 8.0 times longer than broad, with numerous simple seta;
ischium without setae; merus with one seta; carpus with two setae; propodus about 9.0 times
longer than broad, with seta; dactylus with short seta, length with unguis subequal to propodus
length. Pereopods 2 and 3 basis with broom and simple setae; ischium with one seta, merus with
two setae, carpus with blade greater than half length of propodus and dactylus with unguis half
length of propodus. Pereopod 4 basis about 6.5 times longer than broad. Pereopods 4 to 6 basis
with broom setae, ischium and merus with one seta, and propodus with broom seta. Pleopods
elongate with terminal setae only. Uropod exopods and endopods each with two articles;
exopods 75 percent length of endopods.
Type Locality
Northeast Atlantic: Rockall Trough, west of Hebrides Terrace Seamount.
Depth Range
2081-2634 m.
Distribution
Rockall Trough (type locality) and Porcupine Seabight in the Northeast Atlantic.
Remarks
This species is closely related to Pseudotanais affinis, according to Bird and Holdich
(1989), with small differences seen in the setation of pereopods and size of the cheliped. The
blade-like seta of periopods 2 and 3 in P. scalpellum are longer (about 70% of the propodus
109
length) than in P. affmis (about 50%). The terminal dentieulation of the mandibular pars molaris
is nearly indentical in the two species.
Reference
Bird and Holdich (1989): 246-249, Figs. 1G, 6, 7.
Pseudotanais {Pseudotanais) spatula Bird and Holdich, 1989
Synonymy
Pseudotanais longisetosus, Holdich and Bird (1985): 446^447, non Sieg, 1977.
Diagnosis
As for the family, subfamily and subgenus except: Cephalothorax shorter than pereonites
1 to 3. Eyelobes absent. Antennule first article 57 percent of total antennule length, about 7.0
times longer than broad; distal article about 6.0 times longer than broad. Antenna slightly longer
than antennule; second article subequal in length to third article, both with short, stout spiniform
setae; fourth article about 10.0 times longer than broad and fifth article 5.5 times longer than
broad. Mandible pars molaris narrow, with four short subterminal denticles and two longer,
unequal terminal denticles. Maxillule endite terminating with nine spiniform setae. Maxilliped
bases with seta near palp articulation; endites fused between 1/3 and 2/3 length, each with short
seta. Cheliped with non-forcipate chelae; carpus 1.8 times longer than broad, with two inferior
setae; propodus about 3.0 times longer than broad, fixed finger with one inferior seta; dactylus
about 59 percent of propodus length, narrower in width to fixed finger and possessing proximal
seta. Pereopod 1 basis about 9.5 times longer than broad, with numerous simple seta; ischium
without setae; merus with one seta; carpus with three setae; propodus about 10.5 times longer
than broad, with seta; dactylus with short seta, length with unguis subequal to propodus length.
Pereopods 2 and 3 basis with simple setae; ischium with one seta, merus with two setae, carpus
with blade greater than half length of propodus and dactylus with unguis 2/3 length of propodus.
110
Pereopod 4 basis about 4.5 times longer than broad. Pereopods 4 to 6 basis with broom and
simple setae, ischium and merus each with one seta, carpus with additional exceptionally long
superior seta, and propodus with broom seta. Pleopods elongate with terminal setae only.
Uropod exopods and endopods each with two articles; exopods 75 percent length of endopods.
Type Locality
Northeast Atlantic: Porcupine Seabight, west of Ireland.
Depth Range
1400-2209m.
Distribution
Porcupine Seabight (type locality), Hebridean Slope, Celtic Slope and southern Biscay,
all in the Northeast Atlantic.
Remarks
Pseudotanais spatula is another species in the 'affinis' group from the Northeast Atlantic
with only minor differences from P. affinis including longer carpal setae on pereopods 1, 5 and 6.
The mandible pars molaris has two additional subterminal denticles but is otherwise similar to
that of P. affinis and P. scalpellum. The elongate superior carpal setae on pereopods 5 and 6 is a
character shared by P. longispinus but that species has a distinctive pars molaris featuring a single
large terminal spine-like tooth similar to that possessed by P. longisetosus and P. nordenskioldi.
Reference
Bird and Holdich (1989): 249-252, Figs. 8 and 9.
Pseudotanais {Pseudotanais) spicatus Bird and Hold ich, 1989
Synonymy
Pseudotanais sp. B Holdich and Bird (1985).
Ill
Diagnosis
As for the family, subfamily and subgenus except: Cephalothorax longer than pereonites
1 to 3. Eyelobes absent. Antennule first article 63 percent of total antennule length, about 6.0
times longer than broad; distal article about 4.5 times longer than broad. Antenna slightly longer
than antennule, second article subequal to third article, both with short, stout spiniform setae;
fourth article 7.5 times longer than broad and fifth article about 5.0 times longer than broad.
Mandible pars molaris narrow with several medio-distal rows of setules and numerous terminal
and subterminal dentritic denticles. Maxillule endite terminating with nine spiniform setae.
Maxilliped bases with seta near each palp articulation; endites fused between 1/3 and 2/3 length,
each with one short spiniform seta. Cheliped with non-forcipate chelae; carpus 1.5 times longer
than broad, with two inferior setae; propodus about 3.0 times longer than broad, fixed finger with
one inferior seta; dactylus 56 percent of propodus length, subequal in width to fixed finger and
lacking proximal seta. Pereopod 1 basis 10.0 times longer than broad, with simple seta; ischium
and merus each without setae; carpus with four setae; propodus about 6.5 times longer than
broad, with one seta; dactylus without seta, length with unguis distinctly longer than propodus.
Pereopods 2 and 3 basis without setae; ischium with one seta, merus with two setae, carpus with
blade seta less than half length of propodus and dactylus with unguis half length of propodus.
Pereopod 4 basis about 3.5 times longer than broad. Pereopods 4 to 6 basis with broom setae,
ischium with two equal setae, merus with two setae and propodus with broom seta. Pleopods
elongate with terminal setae only. Uropod exopods with two articles, subequal in length to
endopods; endopods with one article.
Type Locality
Northeast Atlantic: Porcupine Seabight, west of Ireland.
Depth Range
2227-4829 m.
112
Distribution
Porcupine Seabight (type locality), Rockall Trough, Porcupine Abyssal Plain, north and
south Biscay, all in Northeast Atlantic.
Remarks
This species is unusually large for a pseudotanaid (up to 4 mm) but is otherwise typical of
the genus. It is superficially very similar to Pseudolanais denticulatus, also from the Northeast
Atlantic, but can be distinguished upon dissection by the drastically different mandible pars
molaris, a unique feature of P. spicatus. Bird and Holdich further commented that P. spicatus
bears a close similarity to P. vitjazi from the North Pacific, differing only in setation features of
the antenna and the merus of pereopods 4 to 6.
Reference
Bird and Holdich (1989): 264-268Figs. 2A-B, 15, 16.
Pseudolanais (Pseudotanais) unicus Sieg, 1977
Diagnosis
As for the family, subfamily and subgenus except: Cephalothorax longer than pereonites
1 to 3. Eye lobes present, with pigment. Antennule, first article 54 percent of total antennule
length, about 4.0 times longer than broad; distal article about 3.5 times longer than broad.
Antenna about same length as antennule, second article subequal in length to third article, both
with short, slender spiniform setae; fourth article about 4.0 times longer than broad and fifth
article about 3.0 times longer than broad. Mandible pars molaris narrow, simple pointed.
Maxillule endite terminating with nine spiniform setae. Maxilliped with seta near each palp
articulation; endites fused exept for notch on distal margin, each with one short seta. Cheliped
with non-forcipate chelae; carpus about 1.4 times longer than broad, with two inferior setae;
propodus about 3.0 times longer than broad, fixed finger with one inferior seta; dactylus 52
113
percent of propodus length, narrower th;m fixed finger and possessing proximal seta. Pereopod 1
basis about 6.0 times longer than broad, without setae; ischium with one seta; merus and carpus
each without setae; propodus about 3.5 times longer than broad, with short seta; dactylus without
setae, length with unguis subequal to propodus. Pereopods 2 and 3 basis without setae, ischium
and merus each with one seta, carpus with blade seta greater than half length of propodus and
dactylus with unguis subequal to length of propodus. Pereopod 3 disproportionately smaller in
size than pereopod 2. Pereopod 4 basis about 3.5 times longer than broad. Pereopods 4 to 6 basis
with broom setae, ischium with two equal setae, merus with two setae and propodus without
broom seta. Pleopods elongate with terminal setae only. Uropod exopods and endopods each
with two articles; exopods 75 percent length of endopods.
Type Locality
Mediterranean Sea: off Naples, Italy in the vicinity of Villa Reale.
Depth Range
52 m.
Distribution
Known only from the type locality in the Mediterranean Sea off Naples, Italy.
Remarks
Pseudotanais unicus is distinguished from other oculate members of the subgenus by its
undersized pereopod 3 and elongate carapace (longer than broad).
Reference
Sieg (1977): 85-89, Figs. 62-64, 68.
Pseudotanais {Pseudotanais) vitjazi Kudinova-Pasternak, 1966
Diagnosis
114
As for the family, subfamily and subgenus except: Cephalothorax longer than pereonites
1 to 3. Eyelobes absent. Antennule llrst article 55 percent of total antennule length, about 6.0
times longer than broad; distal article about 6.0 times longer than broad. Antenna slightly longer
than antennule, second article subequal to third article, both with short, stout spiniform setae;
fourth article 8.5 times longer than broad and fifth article about 5.0 times longer than broad.
Mandible pars molaris narrow with several medio-distal rows of setules and numerous terminal
and subterminal dentritic denticles. Maxillule endite terminating with nine spiniform setae.
Maxilliped bases without seta near each palp articulation; endites fused between 1/3 and 2/3
length, each with one short seta. Cheliped with weakly forcipate chelae; carpus 1.6 times longer
than broad, with two inferior setae; propodus about 3-5 times longer than broad, fixed finger with
one inferior seta; dactylus 55 percent of propodus length, subequal in width to fixed finger and
lacking proximal seta. Pereopod 1 basis about 9.0 times longer than broad, with simple seta;
ischium and merus each without setae; carpus with two setae; propodus about 7.0 times longer
than broad, without setae; dactylus without seta, length with unguis subequal to propodus.
Pereopods 2 and 3 basis without setae; ischium not described or illustrated, merus with two setae,
carpus with blade seta less than half length of propodus and dactylus with unguis distinctly less
than half length of propodus. Pereopod 4 basis about 5.0 times longer than broad. Pereopods 4 to
6 basis without setae, ischium with one seta, merus setation inconclusively illustrated, and
propodus without broom seta. Pleopods elongate with terminal setae only. Uropod exopods and
endopods each with two articles; exopods subequal in length to endopods.
Type Locality
North Pacific: abyssal region south of the Bering Sea.
Depth Range
6065 m.
Distribution
115
Known only from the type locality in the north central Pacific Ocean.
Remarks
As in other species accredited to Kudinova-Pasternak, Pseudotanais vitjazi is
inadequately described and poorly illustrated. Pereopod 2 was illustrated without showing the
ischium so that proportions assumed for the basis might be inaccurate. Pseudotanais vitjazi can
definitely be assigned to the subgenus pseudotanais, s.s. because of the obvious blade-like setae
seen in the illustration of pereopod 2. Pseudotanais vitjazi has a noticeably similar pars molaris
structure to that of P. spicalus from the Northeast Atlantic as noted by Bird and Holdich (1989:
268). They stated that the only differences between the two species are longer denticles on the
pars molaris and stronger spiniform setae on the merus of pereopods 4 to 6 of P. spicatus, and
stronger spiniform setae on antenna articles 2 and 3 of P. vitjazi (see remarks under P. spicatus);
however, given the inadequacies of Kudinova-Pasternak's illustrations, all these character
differences are questionable. Sieg (1977: 105) considered P. vitjazi to be incertae sedis mainly
because he had no material to examine, but likely also because of the meagre description given by
its author. Nevertheless, he did include it in his key to the worldwide species (Sieg and Heard,
1988: 43) separating it from P. affmis by highlighting the subequal length of the uropodal rami
and distinct denticulation of the pars molaris.
References
Kudinova-Pasternak (1966): 532-534, Fig. 12; Sieg and Heard (1988): 43 (key).
Pseudotanais {Pseudotanais) vulsella Bird and Holdich, 1989
Diagnosis
As for the family, subfamily and subgenus except: Cephalothorax longer than pereonites
1 to 3. Eyelobes absent. Antennule first article 57 percent of total antennule length, about 6.5
times longer than broad; distal article about 5.5 times longer than broad. Antenna about same
116
length as antennule, second article subequal to length of third article, both with short, slender
spiniform setae; fourth article about 7.0 times longer than broad and fifth article about 3.0 times
longer than broad. Mandible pars molaris narrow, with small denticles along one margin.
Maxillule endite terminating with nine spiniform setae. MaxiJliped bases with seta near each palp
articulation; endites fused exept for notch on distal margin, each with two short cusps. Cheliped
with strongly forcipate chelae; carpus 1.6 times longer than broad, with two inferior setae;
propodus about 3.0 times longer than broad, fixed finger with one inferior seta; dactylus 60
percent of propodus length, narrower in width to fixed finger and possessing proximal seta.
Pereopod 1 basis about 12.5 times longer than broad, without setae; ischium, merus and carpus
each with one seta; propodus about 6.0 times longer than broad, with two setae; dactylus with
short seta, length with unguis subequal to length of propodus. Pereopods 2 and 3 basis with
broom setae, ischium with one seta, merus with two setae, carpus with blade-like seta greater than
half length of propodus and dactylus with unguis half length of propodus. Pereopod 4 basis about
4.5 times longer than broad. Pereopods 4 to 6 basis without setae, ischium with two equal setae,
merus with two setae and propodus without broom seta. Pleopods elongate, with terminal setae
only. Uropod exopods and endopods each with two articles; exopods subequal in length to
endopods.
Type Locality
Northeast Atlantic: Porcupine Seabight, west of Ireland.
Depth Range
1028-1640m.
Distribution
North Feni Ridge, Rockall Trough, Hebridean Slope, Celtic Slope and Porcupine
Seabight (type locality), all in the Northeast Atlantic west of Ireland.
Remarks
117
This Northeast Atlantic species can be distinguished from the closely related
Pseudotanais fakiculata, another forcipate-cheiae species, by the longer cheliped dactylus (60%
length of the propodus/fixed finger) and longer carpal blade-like setae on pereopods 2 and 3.
Among other forcipate members of the subgenus, the maxilliped endites are completely fused in
P. forcipatus, P. jonesi, and P. californiensis and fused except for a distal notch in P. abyssi;
however in the latter species each endite is armed with a small marginal seta rather than the
triangular cusps of P. vulsella.
Reference
Bird and Holdich (1989): 269-273, Figs. IE, 17, 18.
Pseudotanais {Pseudotanais) sp. A, n. sp. (Figures 14-17)
Diagnosis
As for the family, subfamily and subgenus except: Cephalothorax longer than pereonites
1 to 3. Eyelobes absent. Antennule first article-60 percent of total antennule length, about 5.5
times longer than broad; distal article about 3.0 times longer than broad. Antenna slightly longer
than antennule, second article shorter than third article, both with short, stout spiniform setae;
fourth article about 7.0 times longer than broad and fifth article about 3.0 times longer than broad.
Mandible pars molaris broad, truncate, with numerous accessory denticles and two short marginal
setae. Maxillule endite terminating with nine spiniform setae. Maxilliped bases with seta near
each palp articulation; endites fused between 1/3 and 2/3 length, each with one short seta and two
short cusps. Cheliped with non-forcipate chelae; carpus 1.4 times longer than broad, with two
inferior setae; propodus about 2.5 times longer than broad, fixed finger with one inferior seta;
dactylus 56 percent of propodus length, narrower in width to fixed finger and possessing
proximal seta. Pereopod 1 basis about 11.0 times longer than broad, with simple seta; ischium
without setae; merus and carpus each with one seta; propodus about 5.0 times longer than broad,
with two setae; dactylus without setae, length with unguis subequal to length of propodus.
118
Pereopods 2 and 3 basis with broom setae, ischium and merus each with one seta, carpus with
blade-like seta less than half length of propodus and dactylus with unguis distinctly less than half
length of propodus. Pereopod 4 basis about 2.5 times longer than broad. Pereopods 4 to 6 basis
with broom setae, ischium with one seta, merus with two setae and propodus with broom seta.
Pleopods elongate, with terminal setae only. Uropod exopods and endopods each with two
articles; exopods subequal in length to endopods.
Type Locality
North Atlantic: Gulf of Mexico, Texas A&M Deepwater Gulf of Mexico Station RW3-5
in the western Gulf of Mexico.
Depth Range
379-297'4 m.
Distribution
Gulf of Mexico.
Material examined.
Holotype, non-ovigerous female, 1.8 mm, DGoM station RW3-5, 27° 00.5014N 95°
5417W, western Gulf of Mexico, 22 May 2000, 1335 m. Paratypes: one manca from type
localilty; DGoM Station Bl-5, one preparatory female, one female with spent marsupium;
DGoM Station B2-1, two females, one manca; DGoM Station B2-2, four females; DGoM
Station B2-3, seven females, one manca; DGoM Station B3-3, one manca; DGoM Station BH-5,
two females, one female dissected and slide mounted; DGoM Station C4-1, one female, one
manca; DGoM Station C4-2, one large female, one female dissected and slide mounted; DGoM
Station C4-3 one female, one manca; DGoM Station C7-1, four females; DGoM Station C7-2,
two females; DGoM Station HP-3, four females, one female with spent marsupium; DGoM
Station MT3-1, two females; DGoM Station MT4-2, three females, ten mancas; DGoM Station
NB3-3, one large female, one manca; DGoM Station RW2-1, four females; DGoM Station
RW2-3, six females, one with marsupium lacking ova; DGoM Station RW3-3, one small female;
119
DGoM Station RW4-1, three females; DGoM Station RW5-1, two females; DGoM Station
RW5-2, five females, two mancas; DGoM Station RW5-3, one large female; DGoM Station
S35-1, one large preparatory female, four smaller females; DGoM Station S35-2, two females,
one manca; DGoM Station S36-1, two females; DGoM Station S41-2, one female; DGoM
Station S42-1, one female; DGoM Station W l - 1 , four small females; DGoM Station W3-2, one
manca; DGoM Station W4-1, one small female; DGoM Station W4-5, four mancas; DGoM
Station WC12-1, one small female; DGoM Station WC12-2, one female, two mancas; DGoM
Station WC5-1, one manca.
Description
Based on holotype (Figs. 14A, B) and non-ovigerous female paratype. Body 0.8-2.2
mm, 4.1 times longer than broad.
Cephalothorax longer than pereonites 1-3, subtriangular, becoming narrow anteriorly,
with pair of antero-lateral setae, 19.4 percent total length; eyelobes absent.
Pereon 51.8 percent total length, pereonite 1 abreviated.
Pleon shorter than pereonites 5+6, 17.5 percent total length.
Pleotelson subequal to pleonites 3-5, 11 percent total length, apex slightly produced,
with one pair of distolateral setae.
Antennule (Fig. 15C) 29.1 percent of body length, with three articles; first article 59.7
percent of total length, 5.4 times longer than broad, with one short medial simple seta, one long
medial simple seta, one medial broom seta, one short distal simple seta, one long distal simple
seta and two distal broom setae; second article 2.6 times longer than broad, with one long distal
simple seta and one short distal simple seta; distal article 2.8 times longer than broad, terminating
with three simple setae, one aesthetasc and three bidid-tipped setae.
Antenna (Fig. 15D) with six articles, 104.2 percent length of Al; second article 1.4 times
longer than broad, about 75 percent length of third article, both articles distally with short, stout
spiniform seta; fourth article 6.8 times longer than broad, with one short distal simple seta, one
120
long distal simple seta, one long distal spiniform seta, and three distal broom setae; fifth artiele
2.9 times longer than broad, distally with one long simple seta; sixth article small, terminating
with six simple setae (two short, four long).
Moulhparls. Labrum (Fig. 16C) cap-shaped, distal margin finely setose. Mandibles
(Figs. 16A, B) with distal margins coarsely denticulate (left) and finely denticulate (right), right
lacinia mobilis represented by short, pointed process, left lacinia mobilis well developed and
denticulate, pars molaris broad, with numerous accessory denticles and two short marginal setae.
Maxillule (Fig. 19E) palp with two equal setae, terminating with nine spiniform setae, one with
subterminal setule and two accessory setae. Maxilliped (Fig. 16D) bases completely fused, with
seta near each palp articulation; endites fused between 1/3 and 2/3 length, each with one short
seta and two short cusps, with lateral margins setulose; palp with four articles; article 1 without
setae; article 2 with three unequal inner edge simple setae and one outer-edge simple seta; article
3 with four unequal inner-edge simple setae; article 4 with one small outer-edge simple seta and
five inner-edge simple setae. Epignath (Fig. 16F) falciform, tip unarmored.
C'heI'iped (Fig. 15A) strongly built, chela not forcipate; sclerite well developed; basis 2.0
times longer than broad, without setae; merus triangular, with inferior seta; carpus 1.4 times
longer than broad, with two unequal medial inferior setae and one distal superior seta; propodus
2.5 times longer than broad, about 2.0 times longer than carpus, palm with seven comb setae (one
longer than others); fixed finger with one inferior seta, three superior setae (two simple, one
limbate) and one seta near articulation with dactylus; dactylus 55.6 percent of propodus length,
width narrower than fixed finger, with proximal simple seta.
Pereopod 1 (Fig. 17A) basis 11.2 times longer than broad, with proximal superior seta;
ischium without setae; merus 1.9 times longer than broad, distally with short inferior seta; carpus
2.9 times longer than broad, 1.5 times length of merus, distally with one short superior spiniform
seta; propodus 4.9 times longer than broad, distally with short superior seta and long inferior seta;
dactylus without seta, length with unguis slightly longer than propodus.
121
Pereopod 2 (Fig. 17B) basis 7 times longer than broad, with proximal inferior broom
seta; ischium with short inferior spiniform seta; merus 2.2 times longer than broad, distaily with
one short inferior spiniform seta; carpus 3.9 times longer than broad, about 1.5 times length of
merus, distaily with one short superior spiniform seta, one short superior simple seta, one inferior
blade-like seta of about 30 percent length of propodus and one short, broad inferior spiniform
seta; propodus 9.5 times longer than broad, with one long distal inferior spiniform seta (near
length of dactylus-unguis); dactylus without setae, length with unguis less than half length of
propodus.
Pereopod 3 (Fig. 17C) basis 5.3 times longer than broad and without broom seta; carpus
3 times longer than broad; propodus 6.2 times longer than broad; dactylus bifid-tipped; otherwise
similar to pereopod 2.
Pereopod 4 (Fig. 17D) basis 2.4 times longer than broad, tumid, with setae lacking;
ischium with one short spiniform seta; merus 2.5 times longer than broad, distaily with two short
inferior spiniform setae; carpus 2.8 times longer than broad, 1.5 times longer than merus, distaily
with one short superior simple seta, two unequal serrate spiniform setae and one inferior bladelike spiniform seta of about 20 percent length of propodus; propodus 5.7 times longer than broad,
1.2 times longer than carpus, distaily with one long superior terminal spiniform seta, two short
inferior spiniform setae and one superior broom seta; dactylus fused with unguis to form claw,
length with unguis less than half length of propodus.
Pereopod 5 (Fig. 17E) basis with proximal broom seta; merus distaily with two short
unequal spiniform setae; otherwise similar to P4.
Pereopod 6 (Fig. 17F) merus distaily with two short unequal spiniform setae; carpus 1.9
times longer than broad; propodus with additional terminal spiniform seta and superior marginal
spinules, otherwise similar to peropod 4.
Pleopods (Fig. 15E) rami elongate, with terminal setae only; endopod about 4/5 length of
exopod, with seven terminal setae; exopod with eleven terminal setae.
122
Uropods (Fig. 15B) exopod with two articles, slightly shorter than endopod, proximal
article slightly less than half length of ramus, with one distal simple seta, distal article with two
unequal simple setae; endopod with two articles, proximal article about half length of ramus, with
three long distal simple setae, distal article with four long and two short simple setae.
Remarks
This new species is clearly in the same group as Pseudotanais denticulatus, P. corollatus
and probably P. inflatus as evidenced by the similar mandibular molar process which is broad and
terminally ringed with denticles and setae. Of the three, species A appears to be most similar to P.
denticulatus which also has short, stout spiniform setae on the antenna second and third articles,
whereas those of P. corollatus are long, slender spiniform and those of P. inflatus are apparently
short, slender simple setae. Except where noted below, the appendages are nearly indentical in
proportions and setation to P. denticulatus, including the bifid-tipped dactyl on pereopod 3. The
new species can be distinguished from P. denticulatus by (1) the notably longer percentage of
lengths of the pleotelson and antennule to total body length, (2) antenna articles 4 and 5 being
more stout, (3) a long distal spiniform seta on antenna article 4 (lacking in P. denticulatus and
others of the group), (4) maxilliped endites each bearing two cusps (lacking in P. denticulatus),
(5) a limbate seta among the three superior setae on the cheliped fixed finger (all normal simple
setae on P. denticulatus), (6) the basis of pereopod 1 longer and more slender, (7) a much longer
inferior seta on propodus of pereopod 1, (8) one seta rather than two on merus of pereopods 2 and
3, (9) the blade-like carpal seta on pereopods 2 and 3 are about 30 percent of the length of the
propodus (50 percent in P. denticulatus), (10) the dactylus-unguis length of pereopods 2 and 3 is
less than half the length of the propodus (greater than half in P. denticulatus), (11) the ischium of
pereopods 4 to 6 has a single spiniform seta (an additional simple seta is present in P.
denticulatus), (12) the merus of pereopod 4 has two short, stout spiniform setae of equal length
(only one is present in P. denticulatus which has an additional simple seta of differing length),
123
and (13) the uropodal exopod is nearly the same length as the endopod (about 75 percent the
length of endopod in P. denticulatus).
124
Figure 14. Pseudotanais (Pseudotanais) sp. A. Holotype, female. A, Dorsal view. B, Lateral
view. Scale = 0.5 mm
125
Figure 15. Pseudotanais {Pseudotanais) sp. A. Paratype, female. A, Cheliped inner face. B,
Uropod. C, Antennule. D, Antenna. E, Pleopod. Scale = 0.2 mm
126
Figure 16. Pseudotanais {Pseudotanais) sp. A. Paratype, female. A, Left mandible, inset shows
detail of pars molaris tip. B, Right mandible. C, Labrum. D, Maxillipeds. E, Maxillule. F,
Epignath. Scale = 0.2 mm
127
Figure 17. Pseudotanais {Pseudotanais) sp. A. Paratype, female. A - F, Pereopods 1 to 6
respectively. Scale = 0.2 mm
128
Pseudotanais (Pseudolanais) sp. C, n. sp. (Figures 18-21)
Diagnosis
As for the family, subfamily and subgenus except: Cephalothorax longer than pereonites
1 to 3. Eyelobes absent. Antennule first article 55 percent of total antennule length, about 8.5
times longer than broad; distal article about 6.0 times longer than broad. Antenna 96 percent
length of antennule, second article shorter than third article, both with short, stout spiniform
setae; fourth article 9.5 times longer than broad and fifth article 3.5 times longer than broad.
Mandible pars molaris narrow with long terminal spiniform process and 3 subtermina! denticles.
Maxillule endite terminating with nine spiniform setae. Maxilliped bases with two medially
located setae; endites fused between 1/3 and 2/3 length, each with one short seta. Cheliped with
non-forcipate chelae; carpus 1.4 times longer than broad, with two inferior setae; propodus 3.8
times longer than broad, fixed finger with one inferior seta; dactylus 62 percent of propodus
length, narrower than fixed finger and possessing proximal seta. Pereopod 1 basis about 8.0
times longer than broad, with simple seta; ischium and merus each with one seta and carpus with
two setae; propodus about 10.5 times longer than broad, with seta; dactylus without seta, length
with unguis distinctly shorter than propodus. Pereopods 2 and 3 basis with broom and simple
setae; ischium and merus each with one seta, carpus with blade seta greater than half length of
propodus and dactylus with unguis half length of propodus. Pereopod 4 basis about 5.0 times
longer than broad, with simple and broom setae. Pereopods 4 to 6 ischium with two unequal
setae, merus with two setae, carpus with additional bifid-tipped setae and propodus with broom
seta. Pleopods elongate with terminal setae only. Uropod exopods and endopods each with two
articles; exopods 2/3 length of endopods.
Type Locality
North Atlantic: Gulf of Mexico, Texas A&M Deepwater Gulf of Mexico Station B3-2 in
the western Gulf of Mexico.
Depth Range
129
212-3145 m.
Distribution
Gulf of Mexico.
Material Examined
Holotype, non-ovigerous female, 1.1 mm, DGoM Station B3-2,26°09.9750'N
91°43.9772'W, western Gulf of Mexico, 10 May 2000, 2650 m. Paratypes: four females and one
manca from type localilty; DGoM Station S42-2, two mancas; DGoM Station C7-2, one manca;
DGoM Station B2-1, three females, one with nematode parasite; DGoM Station B2-2, one
preparatory female; DGoM Station CI—1, four females, one manca; DGoM Station C4-1, three
females; DGoM Station RW5-1, six females; DGoM Station RW6-1, one female; DGoM Station
S36-1, ten females, one manca; DGoM Station S37-1, four females; DGoM Station S41-1, one
female; DGoM Station W2-1, two females; DGoM Station W2-2, four females, one with
nematode parasite; DGoM Station W4-1, two females; DGoM Station W6-2, one female; DGoM
Station B l - 1 , one ovigerous female, three females, two mancas; DGoM Station NB5-2, one
preparatory female dissected and slide mounted, four females; DGoM Station S35-2, one small
female; DGoM Station S37-2, ten females, three mancas; DGoM Station AC1-3, two females;
DGoM Station B3-3, one female, one manca; DGoM Station RW5-2, one female; S36-3, one
female, three mancas; DGoM Station S3 7-3, one preparatory female, four females, six mancas;
DGoM Station S38-2, one preparatory female; DGoM Station W2-3, three females, three
mancas; DGoM Station W5-2, two females, one dissected and slide mounted; DGoM Station
WC12-3, two females, three mancas; DGoM Station MT6-4, one small female; DGoM Station
S44-3, two females; DGoM Station RW6-5, one female; DGoM Station Bl-5, one female;
DGoM Station W2-3, one female, one manca; DGoM Station WC5-5, two females.
Description
Based on holotype (Figs. 18A, B) and non-ovigerous and preparatory female paratypes.
Body 0.8-1.3 mm, 3.2 times longer than broad.
130
Cephalothorax longer than pereonites 1-3, subtriangular, becoming narrow anteriorly,
slightly narrower than pereon, with slight rostrum and pair of antero-lateral setae, 19.1 percent
total length; eyelobes absent.
Pereon 53.3 percent total length, pereonite 1 abreviated; pereonites 1, 4 and 5 with pair of
dorso-lateral setae; pereonites 3-6 with additional pair of ventro-lateral setae. Pleon subequal to
pereonites 5 + 6, 23.6 percent total length, pleonites 2-5 with pair of dorso-lateral setae, pleonites
1-5 with additional pair of ventro-lateral setae.
Pleotelson subequal to pleonites 4-5, 8 percent total length, apex slightly produced, with
two apical setae and one pair of distolateral setae.
.»-
Anlennule (Fig. 19B) 35.8 percent of body length, with three articles; first article 55.4
percent of total length, 8.5 times longer than broad, with one long medial simple seta, six short
medial simple setae, two short distal simple setae, one long distal simple seta and two distal
broom setae; second article 3.4 times longer than broad, with one long distal simple seta, one
short distal simple seta and one distal broom seta; distal article 6.2 times longer than broad,
terminating with three simple setae (two long, one short), one aesthetasc and three bidid-tipped
setae.
Antenna (Fig. 19C) with six articles, 96.1 percent length of Al; second article 1.6 times
longer than broad, about 75 percent length of third article, both articles distally with short, stout
spiniform seta; fourth article 9.3 times longer than broad, with one long medial simple seta, one
short distal simple seta, one long distal spiniform seta, three distal broom setae, and one sub-distal
broom seta; fifth article 3.6 times longer than broad, distally with one long simple seta; sixth
article small, terminating with one aesthetasc and four simple setae.
Mouthparts. Labrum (Fig. 20C) cap-shaped, distal central margin finely setose.
Mandibles (Figs. 20A, B) with distal margins coarsely denticulate (left) and finely denticulate
(right), right lacinia mobilis represented by short, pointed process, left lacinia mobilis well
developed and coarsely denticulate, pars molaris acuminate, with long terminal spiniform process
131
and three subterminal denticles. Labium (Fig. 20D) medially cleft with submarginal ridges.
Maxillule (Fig. 20G) palp with two equal setae, terminating with nine spiniform setae, one with
subterminal setule and two accessory setae. Maxilla not observed. Maxilliped (Fig. 20E) bases
completely fused, with two medially located setae; endites fused between 1/3 and 2/3 length, each
with one short seta, with lateral margins smooth; palp with four articles; article 1 without setae;
article 2 with three unequal inner edge simple setae and one outer-edge spiniform seta; article 3
with four unequal inner-edge simple setae; article 4 with one small outer-edge simple seta and
five inner-edge simple setae. Epignath (Fig. 20F) elongate, sausage-shaped, tip unarmored.
Che I iped (Fig. 19A) strongly built, chela not forcipate; sclerite well developed; basis 1.5
times longer than broad; merus triangular, with inferior seta; carpus 1.4 times longer than broad,
with two unequal medial inferior setae, one distal superior seta and one medial superior seta;
propodus 3.8 times longer than broad, about 2.1 times longer than carpus, palm with five short
setulate comb setae; fixed finger with one inferior seta, three superior setae and one seta near
articulation with dactylus; dactylus 62.1 percent of propodus length, width narrower than fixed
finger, with proximal spiniform seta.
Pereopod I (Fig. 21 A) coxa with seta; basis 8.0 times longer than broad, with medial
inferior seta, proximal superior seta, and distal inferior seta; ischium with small seta; merus 2.4
times longer than broad, distally with short superior seta; carpus 3.9 times longer than broad, 1.5
times length of merus, distally with two short simple setae; propodus 10.4 times longer than
broad, distally with short inferior seta; dactylus without seta, length with unguis half length of
propodus.
Pereopod 2 (Fig. 21B) coxa with seta; basis 7.7 times longer than broad, with medial
superior broom seta, short proximal inferior seta, and short distal inferior seta; ischium with short
inferior seta; merus 2.4 times longer than broad, distally with long inferior spiniform seta; carpus
4.5 times longer than broad, about 1.2 times length of merus, distally with one short superior
spiniform seta, one inferior blade-like seta of about 65 percent length of propodus and one short
132
spiniform seta with broad basal collar; propodus 8.9 times longer than broad, with inferior
marginal spinuies, one long distal inferior spiniform seta (near length of dactylus-unguis) and
setulose terminal margin at emergence of dactylus; dactylus without setae, length with unguis
slightly longer than half length of propodus.
Pereopod 3 (Fig. 21C) basis lacking proximal inferior seta; carpus 2.6 times longer than
broad; propodus 6.4 times longer than broad; dactylus length with unguis about half length of
propodus; otherwise similar to pereopod 2.
Pereopod 4 (Fig. 2ID) coxa with seta; basis 5.2 times longer than broad, with one medial
broom seta, one proximal broom seta, and one medial inferior simple seta; ischium with two
unequal setae, longer seta about half length of merus; merus 2.3 times longer than broad, distally
with one short inferior spiniform seta and one long inferior spiniform seta; carpus 3.2 times
longer than broad, 1.7 times longer than merus, distally with one short inferior simple seta, one
anterior spiniform seta, one inferior blade-like spiniform seta of about 55 percent length of
propodus, one bifid-tipped simple seta of about half length of propodus, inferior marginal
spinuies and four transverse rows of superior marginal setules (very faint); propodus 5.2 times
longer than broad, about as long as carpus, distally with one long supero-terminal spiniform seta
with heavily setose distal half, two unequal inferior spiniform setae, one superior broom seta,
inferior marginal spinuies and setulose terminal margin at emergence of dactylus; dactylus fused
with unguis to form claw, margins slightly setulose, length with unguis about half length of
propodus.
Pereopod 5 (Fig. 21E) similar to pereopod 4.
Pereopod 6 (Fig. 2IF) basis lacking broom seta, carpus distally with inferior simple seta
non-bifid tipped, about 2/3 length of propodus, propodus with additional terminal spiniform seta
and lacking broom seta, otherwise similar to peropod 4.
Pleopods (Fig. 19D) rami elongate, with terminal setae only; endopod about 4/5 length of
exopod, with five terminal setae; exopod with eight terminal setae.
133
Uropods (Fig. 19E) exopod with two articles, 2/3 length of endopod, proximal article
about half length of ramus, with one distal simple seta, distal article with two unequal simple
setae; endopod with two articles, proximal article slightly greater than half length of ramus, with
one long distal simple seta or one broom seta, distal article with four long and two short simple
setae.
Remarks
This new species is similar to those of the "affinis" group of Pseudotanais as presented
by Bird and Holdich (1989) owing to several characters, primarly its acuminate mandibular molar
process with terminal denticulation. By virtue of its inferodistal propodal spiniform setae being
subequal in length to the dactylus-unguis of pereopods 2 and 3, it is allied to four members of the
group: P. affinis, P. scalpellum, P. spatula and P. longispinus. Of these, it appears closest to the
subgroup which includes P. affinis and P. scalpellum because the superior carpal seta of
pereopods 5 and 6 are only about half the length of the propodus (as long or longer in the P.
spatula and P. longispinus). It is similar to P. affinis because it possesses a short spiniform seta as
well as a longer simple seta on the ischium of pereopods 4 to 6 (spiniform seta lacking on P.
scalpellum) and is similar to P. scalpellum by having cephalothorax slightly longer than
pereonites 1-3 (shorter in P. affinis). It differs from both these species by lacking a long simple
seta on the merus of pereopod 1, and having a unique bifid-tip on the aforementioned superior
carpal seta of pereopods 5 and 6. The mouthparts of species C appear nearly identical in most
respects to other members of the "affinis" group. The molar process terminates in a more elongate
process than P. scalpellum and P. affinis, and as such resembles more the apparatus of P.
longispinus and P. spatula; however the maxillipeds exhibit more derived characters (less
setation) than P. longispinus and also P. nordenskioldi.
134
Figure 18. Pseudotanais {Pseudotanais) sp. C. Holotype, female. A, Dorsal view. B, Lateral
view. Scale = 0.5 mm
135
Figure 19. Pseudotanais (Pseudotanais) sp. C. Paratype, female. A, Cheliped inner face, inset
shows detail of comb seta. B, Antennule. C, Antenna. D, Pleopod. E, Uropod. Scale = 0.2 mm
136
Figure 20. Pseudotanais (Pseudotanais) sp. C. paratype, female. A, Left mandible, inset
shows detail of pars molaris (arrow). B, Right mandible. C, Labrum. D, Labium. E,
Maxillipeds. F, Epignath. G, Maxillule, inset shows detail of spiniform seta with setule
(arrow). Scale = 0.2 mm
137
Figure 21. Pseudotanais (Pseudotanais) sp. C. Paratype, female. A - F, Pereopods 1 to 6,
respectively. Insets at D and E show detail of bifid-tipped setae. Scale = 0.2 mm
138
Pseudotanais {Pseudotanais) sp. O, n. sp. (Figures 22 - 25)
Diagnosis
As for the family, subfamily and subgenus except: Cephalothorax subequal in length to
pereonites 1 to 3. Eyelobes absent. Antennule first article 55 percent of total antennule length,
about 3.0 times longer than broad; distal article about 7.5 times longer than broad. Antenna 95
percent length of antennule; second article shorter than third article, both with short, stout
spiniform setae; fourth article about 11.0 times longer than broad and fifth article 3.5 times longer
than broad. Mandible pars molaris narrow, with with three subterminal denticles. Maxillule
endite terminating with nine spiniform setae. Maxilliped bases with two medially located setae;
endites fused between 1/3 and 2/3 length, each with short seta. Cheliped with non-forcipate
chelae; carpus 1.8 times longer than broad, with two inferior setae; propodus about 3.0 times
longer than broad, fixed finger with one inferior seta; dactylus about 62 percent of propodus
length, narrower in width to fixed finger and possessing proximal seta. Pereopod 1 basis about
9.5 times longer than broad, with three simple seta; ischium with one seta; merus with two setae;
carpus with three setae; propodus about 10.0 times longer than broad, with seta; dactylus without
setae, length with unguis subequal to propodus length. Pereopods 2 and 3 basis with broom and
simple setae; ischium with one seta, merus with two setae, carpus with blade greater than half
length of propodus and dactylus with unguis 2/3 length of propodus. Pereopod 4 basis about 4.5
times longer than broad. Pereopods 4 to 6 basis with broom and simple setae, ischium with two
unequal setae, merus with one seta, carpus with additional exceptionally long superior seta and
propodus with broom seta. Pleopods elongate with terminal setae only. Uropod exopods and
endopods each with two articles; exopods 75 percent length of endopods.
Type Locality
North Atlantic: Gulf of Mexico, Texas A&M Deepwater Gulf of Mexico Station MT2-1
in the western Gulf of Mexico.
Depth Range
139
625-1401 m.
Distribution
Gulf of Mexico.
Material Examined
Holotype, non-ovigerous female, 1.4 mm DGoM Station MT2-1,28°27.0646'N
89°40.3563'W, western Gulf of Mexico, 17 June 2000, 676 m. Paratypes: ten females from type
locality; DGoM Station S42-2, one manca; DGoM Station MT3-5, one preparatory female;
DGoM Station C7-1, two mancas; DGoM Station MT3-1, five females, four mancas; DGoM
Station MT3-2, two females - one dissected and slide mounted, one manca; DGoM Station S3 5 1, one female, one manca; DGoM Station W3-1, three females - one dissected and slide mounted,
one with nematode parasite; DGoM Station S35-2, two females; DGoM Station MT2-2, twelve
females, four mancas; DGoM Station MT4-2, one manca; DGoM Station W2-3, one female.
Description
Based on holotype (Figs. 22A, B) and non-ovigerous female paratype. Body 1.3-1.6 mm,
3.3 times longer than broad.
Cephalothorax subequal to pereonites 1-3, subtriangular, becoming narrow anteriorly,
with slight rostrum and pair of antero-lateral setae, 18.2 percent total length; eyelobes absent.
Pereon 49.4 percent total length, pereonite 1 abbreviated; pereonites 3-4 with pair of
dorsolateral setae.
Pleon about 1.5 times longer than pereonites 5+6, 24.5 percent total length, all pleonites
with pair of dorso-lateral setae and additional pair of ventro-lateral setae. Pleotelson shorter than
pleonites 4—5, 7.3 percent total length, apex rounded, with pair of mediolateral setae.
Antennule (Fig. 23B) 33.3 percent of body length, with three articles; first article 55.1
percent of total length, 3.1 times longer than broad, with one long medial simple seta, three short
medial simple setae, one short distal simple seta, one long distal simple seta and three distal
broom setae; second article 1.6 times longer than broad, with one long distal simple seta, one
140
short distal simple seta and one distal broom seta; distal article 7.5 times longer than broad,
terminating with three simple setae, one aesthetasc and three bifid-tipped setae.
Antenna (Fig. 23C) with six articles, 95 percent length of Al; second article 1.2 times
longer than broad, about 2/3 length of third article, both articles distally with short, stout
spiniform seta (larger on article 2); fourth article 10.9 times longer than broad, with one short
distal simple seta, two long distal spiniform setae, one distal broom seta, and one sub-distal
broom seta; fifth article 3.5 times longer than broad, distally with one long simple seta; sixth
article small, terminating with one aesthetasc and four simple setae (one short, three long).
Mouthparts. Labrum (Fig. 24F) cap-shaped, distal central margin finely setose.
Mandibles (Figs. 24A, B) with distal margins coarsely denticulate (left) and finely denticulate
(right), right lacinia mobilis represented by short, pointed process, left lacinia mobilis well
developed and coarsely denticulate, pars molaris acuminate, with three subterminal denticles.
Labium (Fig. 24E) medially cleft with submarginal ridges. Maxillule (Fig. 24C) palp with two
equal setae, terminating with nine spiniform setae, two of which have a subterminal setule.
Maxilla not observed. Maxilliped (Fig. 24D) bases completely fused, with two medially located
setae; endites fused between 1/3 and 2/3 length, each with one short seta, with lateral margins
smooth; palp with four articles; article 1 without setae; article 2 with three unequal inner edge
simple setae and one outer-edge simple seta; article 3 with four unequal inner-edge simple setae;
article 4 with one small outer-edge simple seta and five inner-edge simple setae. Epignath not
observed.
Cheliped (Fig. 23A) strongly built, chela not forcipate; sclerite small; basis 1.5 times
longer than broad, without setae; merus nearly oval, with inferior seta; carpus 1.8 times longer
than broad, with two unequal medial inferior setae, one distal superior seta and one medial
superior seta; propodus 2.8 times longer than broad, about 1.9 times longer than carpus, palm
with four setulate comb setae (one long, three short) and two rows of small spinules and setules;
fixed finger with one inferior seta, three superior setae and one seta near articulation with
141
dactylus; dactylus 62.2 percent of propodus "length, width narrower than fixed finger, with
proximal spiniform seta.
Pereopod 1 (Fig. 25A) coxa with seta; basis 9.5 times longer than broad, with medial
inferior seta, proximal superior seta, and distal inferior seta; ischium with small seta; merus 3.3
times longer than broad, distally with long superior seta and short inferior seta; carpus 2.8 times
longer than broad, 1.1 times length of merus, distally with two short simple setae and one long
superior seta (subequal in length to that of merus); propodus 10.1 times longer than broad,
distally with short superior seta; dactylus without seta, length with unguis slightly shorter than
propodus.
Pereopod 2 (Fig. 25B) basis 6.7 times longer than broad, with proximal superior broom
seta, short medial inferior seta, short proximal inferior seta, and short distal inferior seta; ischium
with short inferior seta; merus 1.8 times longer than broad, distally with one long inferior serrate
spiniform seta and one long inferior simple seta; carpus 4.1 times longer than broad, about 1.4
times length of merus, distally with one short superior spiniform seta, one inferior blade-like seta
of about 75 percent length of propodus and one short spiniform seta with broad basal collar;
propodus 6.9 times longer than broad, with inferior marginal spinules, one long distal inferior
spiniform seta (near length of dactylus-unguis) and setulose terminal margin at emergence of
dactylus; dactylus without setae, length with unguis 2/3 length of propodus.
Pereopod 3 (Fig. 25C) basis lacking medial inferior seta; with superior marginal spinules;
otherwise similar to pereopod 2.
Pereopod 4 (Fig. 25D) basis 4.4 times longer than broad, tumid, with one medial broom
seta, one proximal broom seta, one proximal simple seta, and one medial inferior simple seta;
ischium with two unequal setae, longer seta about half length of merus; merus 2.2 times longer
than broad, distally with one long inferior spiniform seta; carpus 3.8 times longer than broad, 1.7
times longer than merus, distally with one short anterior simple seta, one long superior simple
seta of about half length of propodus, one short inferior spiniform seta and one inferior blade-like
142
spiniform seta of about 45 percent length of propodus; propodus 5.6 times longer than broad,
slightly longer than carpus, distaily with one long supero-terminal spiniform seta with heavily
setose distal half, two unequal inferior spiniform setae, one superior broom seta and superior and
inferior marginal spinules; dactylus fused with unguis to form claw, margins slightly setulose,
length with unguis less than half length of propodus.
Pereopod 5 (Fig. 25E) basis lacking proximal broom seta and lacking medial simple seta,
carpus distaily with short anterior spiniform seta (instead of simple seta), superior seta extending
beyond propodus, and inferior blade-like seta of about 60 percent of propodus length, propodus
4.3 times longer than broad, otherwise similar to P4.
Pereopod 6 (Fig. 25F) basis 5.1 times longer than broad, lacking medial simple seta, and
lacking medial broom seta, carpus distaily with superior seta extending beyond propodus, long
anterior spiniform seta (instead of simple seta), and inferior blade-like seta of about 65 percent of
propodus length, propodus with additional terminal spiniform seta, lacking small subterminal
inferior spiniform seta, and lacking broom seta, otherwise similar to peropod 4.
Pleopods (Fig. 23E) rami elongate, with terminal setae only; endopod slightly shorter
than exopod, with six terminal setae (innermost very short); exopod with nine terminal setae.
Uropods (Fig. 23D) exopod with two articles, 75 percent length of endopod, proximal
article slightly less than half length of ramus, with one distal simple seta, distal article with two
unequal simple setae; endopod with two articles, proximal article slightly greater than half length
of ramus, with one long distal simple seta and two broom setae, distal article with four long and
one short simple setae.
Remarks
Another species of the "affinis" group of Bird and Holdich (1989), this new species is
differentiated from those in the Gulf of Mexico by its combination of a long superior carpal seta
on pereopods 5 and 6 (shorter in species C and P), an additional long simple seta on the merus of
pereopods 2 and 3 (this seta is absent in species C and P), and a long superior seta on the merus
143
and carpus of the first pereopod (long only on the carpus of species P). In these respects, species
O is most similar to the northern Atlantic species Pseudotanais spatula and P. longispinus as
described by Bird and Holdich (1989) and P. nipponicus from the north Pacific, especially in the
nearly identical setation of the pereopods. Of these species, species O appears most closely allied
to P. spatula, differing primarily in the more acute, less denticulate mandible pars molaris, but
also in the general proportions and setation of the body segments. Species O has a considerably
larger pleon (25 percent of the total body length) than P. spatula (about 18 percent), has a broader
antennule article 1 (length 3.0 times width vs. 7.0 times width in P. spatula), and has more
segments bearing doso-lateral setae than illustrated for P. spatula by Bird and Holdich. The pars
molaris of species O is similar in appearance to that found in P. longispinus and also in P.
nipponicus but has fewer subterminal denticles. In addition, the maxiliped endites of species O
lack the short conical cusps present in P. longispinus and also in P. nipponicus.
144
Figure 22. Pseudotanais {Pseudotanais) sp. O. Holotype, female. A, Dorsal view. B, Lateral
view. Scale = 0.5 mm
145
Figure 23. Pseudotanais {Pseudotanais) sp. O. Paratype, female. A, Cheliped inner face. B,
Antennule. C, Antenna. D, Uropod. E, Pleopod. Scale = 0.2 mm
146
Figure 24. Pseudotanais {Pseudotanais) sp. O. Paratype, female. A, Left mandible. B, Right
mandible. C, Maxillule, inset shows detail of spiniform seta with setule (arrow). D,
Maxillipeds. E, Labium. F, Labrum. Scale = 0.2 mm
147
Figure 25. Pseudotanais {Pseudotanais) sp. O. Paratype, female. A - F, Pereopods 1 to 6,
respectively. Scale = 0.2 mm
148
Pseudoianais (Psendotanais) sp. P, n. sp. (Figures 26 - 29)
Diagnosis
As for the family, subfamily and subgenus except: Cephalothorax subequal in length to
pereonites 1 to 3. Eyelobes absent. Antennule first article 50 percent of total antennule length,
about 3.5 times longer than broad; distal article about 5.5 times longer than broad. Antenna 95
percent length of antennule; second article subequal in length to third article, both with short,
stout spiniform setae; fourth article about 9.5 times longer than broad and fifth article 3.5 times
longer than broad. Mandible pars molaris narrow, with three to five subterminal denticles.
Maxillule endite terminating with nine spiniform setae. Maxilliped bases with two medially
located setae; endites fused between 1/3 and 2/3 length, each with short seta. Cheliped with nonforcipate chelae; carpus 2.4 times longer than broad, with two inferior setae; propodus about 3.5
times longer than broad, fixed finger with one inferior seta; dactylus about 69 percent of propodus
length, narrower in width to fixed finger and possessing proximal seta. Pereopod 1 basis about
9.0 times longer than broad, with four simple seta; ischium with one seta; merus without setae;
carpus with three setae; propodus about 10.5 times longer than broad, with seta; dactylus with
exceptionally long seta, length with unguis subequal to propodus length. Pereopods 2 and 3 basis
with broom and simple setae; ischium and merus eachwith one seta, carpus with blade greater
than half length of propodus and dactylus with unguis 75 percent length of propodus. Pereopod
4 basis about 6.5 times longer than broad. Pereopods 4 to 6 basis with broom and simple setae,
ischium with two unequal setae, merus with two setae, carpus with additional bifid-tipped seta
and propodus with broom seta. Pleopods elongate with terminal setae only. Uropod exopods
with one article and endopods with one pseudo-articulated article; exopods about 80 percent
length ofendopods
Type Locality
North Atlantic: Gulf of Mexico, Texas A&M Deepwater Gulf of Mexico Station WC123 in the western Gulf of Mexico.
149
Depth Range
658-2450 m.
Distribution
Gulf of Mexico
Material Examined
Holotype, non-ovigerous female, 1.25 mm, DGoM Station WC12-3,27° 19.4320'N 91°
33.1625' W, western Gulf of Mexico, 5 May 2000, 1175 m. Paratypes: two females and three
mancas from type locality; DGoM Station S42-2, one manca; DGoM Station RW3-1, one nonovigerous female, dissected and slide mounted; DGoM Station RW4-1, one female; DGoM
Station S35-1, one female, three mancas; DGoM Station S36-1, one female; DGoM Station
AC 1-2, two females; DGoM Station C7-2, one female, three mancas; DGoM Station S35-2, one
female, two mancas; DGoM Station MT2-2, one male; DGoM Station S36-3, three females, one
dissected and slide mounted.
Description
Based on holotype (Figs. 26A, B) and non-ovigerous female paratype. Body 1-1.7 mm,
3.2 times longer than broad.
Cephalothorax subequal to pereonites 1-3, subtrapezoidal, narrowing anteriorly, about
equal in width to pereon, with pair of antero-lateral setae, 18.3 percent total length; eyelobes
absent.
Pereon 64.4 percent total length, pereonite 1 abreviated; pereonites 1 and 3-6 with dorsolateral setae, pereonites 3-6 with additional pair of ventro-lateral setae.
Pleon shorter than pereonites 5+6, 16.7 percent total length, all pleonites with pair of
dorso-lateral setae and additional pair of ventro-lateral setae.
Pleotelson (Fig. 27E) shorter than pleonites 3-5, 6.9 percent total length, apex slightly
produced, with pair of dorso-lateral setae and four apical setae (two simple, two spiniform).
150
Ant ennuie (Fig. 27A) 31.1 percent of body length, with three articles; first article 50.4
percent of total length, 3.7 times longer than broad, with three short medial simple setae, one
medial broom seta, one short distal simple seta, one long distal simple seta and three distal broom
setae; second article 2 times longer than broad, with one long distal simple seta, one short distal
simple seta and one distal broom seta; distal article 5.5 times longer than broad, terminating with
two simple setae, one aesthetasc, three bifid-tipped setae and one trifid-tipped seta.
Antenna (Fig. 27B) with six articles, 95.4 percent length of Al; second article 1.4 times
longer than broad, slightly shorter than third article, both articles distally with short, stout
spiniform seta (larger on article 2); fourth article 9.5 times longer than broad, with one short distal
simple seta, two long distal spiniform setae, four distal broom setae, and one sub-distal broom
seta; fifth article 3.3 times longer than broad, distally with one long simple seta; sixth article
small, terminating with two aesthetascs and four simple setae.
Mouthparts. Labrum (Fig. 28B) cap-shaped, distal central margin finely setose.
Mandibles (Figs. 28D, E) with distal margins coarsely denticulate (left) and finely denticulate
(right), right lacinia mobilis represented by short, pointed process, left lacinia mobilis well
developed and coarsely denticulate, pars molaris acuminate, with three to five subterminal
denticles. Labium (Fig. 28C) medially cleft with submarginal ridges. Maxillule (Fig. 28G) palp
with two unequal setae, terminating with nine spiniform setae, three of which have a subterminal
setule. Maxilla not observed. Maxilliped (Fig. 28F) bases completely fused, with two medially
located setae; endites fused between 1/3 and 2/3 length, each with one short seta, with lateral
margins smooth; palp with four articles; article 1 without setae; article 2 with three unequal inner
edge simple setae and one outer-edge simple seta; article 3 with four unequal inner-edge simple
setae; article 4 with one small outer-edge simple seta and five inner-edge simple setae. Epignath
(Fig. 28A) elongate, sausage-shaped, tip unarmored.
Cheliped (Fig. 27D) strongly built, chela not forcipate; sclerite well developed; basis 1.5
times longer than broad, with distal superior seta; merus triangular, with inferior seta; carpus 2.4
151
times longer than broad, with two unequal medial inferior setae, one distal superior seta and one
medial superior seta; propodus 3.7 times longer than broad, about 1.6 times longer than carpus,
palm with four setulate comb setae (one long, three short); fixed finger with one inferior seta,
three superior setae (two simple, one limbate) and one seta near articulation with dactylus;
dactylus 68.8 percent of propodus length, width narrower than fixed finger, with proximal
spiniform seta.
Pereopod 1 (Fig. 29A) coxa with seta; basis 9.1 times longer than broad, with medial
inferior seta, proximal superior seta, proximal inferior seta, and distal inferior seta; ischium with
small seta; merus 2.2 times longer than broad, distally with no setae; carpus 2.5 times longer than
broad, 1.3 times length of merus, distally with two short simple setae and one long superior seta;
propodus 10.5 times longer than broad, distally with short superior seta; dactylus with seta, length
with unguis subequal to length of propodus.
Pereopod 2 (Fig. 29B) coxa with seta; basis 7.1 times longer than broad, with medial
superior broom seta, short proximal inferior seta, and short distal inferior seta; ischium with short
inferior seta; merus 3.2 times longer than broad, distally with long inferior spiniform seta; carpus
4.4 times longer than broad, about 1.2 times length of merus, distally with one short superior
spiniform seta, one inferior blade-like seta of about 60 percent length of propodus and one short
spiniform seta with broad basal collar; propodus 8.1 times longer than broad, with inferior
marginal spinules, superior marginal spinules, one long distal inferior spiniform seta (near length
of dactylus-unguis) and setulose terminal margin at emergence of dactylus; dactylus without
setae, length with unguis 75 percent length of propodus.
Pereopod 3 (Fig. 29C) basis without broom seta; carpus with inferior blade-like seta of
about 85 percent length of propodus; propodus 5.7 times longer than broad; otherwise similar to
pereopod 2.
Pereopod 4 (Fig. 29D) coxa with seta; basis 6.3 times longer than broad, with-two medial
broom setae and one medial inferior simple seta; ischium with two short setae; merus 2.5 times
152
longer than broad, distally with one short inferior simple seta and one long inferior spiniform
seta: carpus 3.6 times longer than broad, 1.7 times longer than merus, distally with one bifidtipped simple seta of about half length of propodus, two short anterior simple setae, one short
inferior spiniform seta and one inferior blade-like spiniform seta of about 45 percent length of
propodus; propodus 5.4 times longer than broad, slightly longer than carpus, distally with one
long supero-terminal spiniform seta with heavily setose distal half, two unequal inferior spiniform
setae, one superior broom seta, superior marginal spinules and setulose terminal margin at
emergence of dactylus; dactylus fused with unguis to form claw, margins slightly setulose, length
with unguis less than half length of propodus.
Pereopod 5 (Fig. 29E) basis 5.1 times longer than broad, carpus distally with long
posterior spiniform seta and lacking anterior simple seta, otherwise similar to P4.
Pereopod 6 (Fig. 29F) basis lacking medial broom seta; ischium with two unequal simple
setae; merus lacking distal simple seta, carpus distally with long anterior spiniform seta (instead
of simple seta), propodus with additional terminal spiniform seta and lacking broom seta,
otherwise similar to pereopod 4.
Pleopods (Fig. 27C) rami elongate, with terminal setae only; endopod about 75 percent
length ofexopod, with six terminal setae (innermost very short); exopod with nine terminal setae.
Uropods (Fig. 27E) exopod with one article, about 80 percent length of endopod, with
one medial simple seta and two unequal terminal simple setae; endopod with one article pseudoarticulated about mid-length, with one long simple seta and two broom setae at pseudoarticulation and five long and one short terminal simple setae.
Remarks
This new species is another member of the Gulf of Mexico group which has prominent
carpal blade setae on pereopods 2 and 3 that exceed greater than half the length of the propodus
(see remarks for species O for differentiation among members of this group). In short, this new
species differs from species C by (1) having terminal spiniform setae on propodus of pereopods 2
153
and 3 shorter by half or 3/4 than the length of the dactylus-unguis (subequal length in species C);
(2) pereopods 2 and 3 lack the long spiniform superior carpus seta present on species C, and (3)
pereopod 1 has a long simple seta on the carpus (absent in species C). It is probably most closely
related to species O, both have the characteristic elongate dactylus-unguis of pereopod 1, but is
distinguished from that species by its lack of long superior simple setae on the carpus of
pereopods 5 and 6 and the merus of pereopods 2 and 3. Furthermore, species P differs from all
others of the group by its uropods which have uni-articulate exopods and pseudo-articulate
endopods. The uropods do, however have the usual setae at the location where these articulations
would be expected, which lends to the speculation that this lack of articulation is of recent
derivation. The uropods notwithstanding, species P falls outside of Bird and Holdich's (1989)
"affinis group" by virtue of its short terminal spiniform propodal setae on pereopods 2 and 3 and
may be somewhere near Pseudotanais colonus which also has uni-articulate, but very short,
uropod exopods. Species P, along with species O and P. spatula, are unique among members of
the subfamily Pseudotanainae in that they have three carpal setae on pereopod 1; all others have
two or less. One male specimen of species P was discovered that had the characteristic long
carpal blades on peropods 2 and 3, antennule with five articles possessing a typical abundance of
setae known for male tanaids, chelae with strong serations on margin of fixed finger, pleopods
present with long, natatory setae, and uni-articulate uropods characteristic of the species.
154
Figure 26. Pseudotanais {Pseudotanais) sp. P. Holotype, female. A, Dorsal view. B, Lateral
view. Scale = 0.5 mm
155
Figure 27. Pseudotanais {Pseudotanais) sp. P. Paratype, female. A, Antennule. B, Antenna.
C, Pleopod. D, Cheliped. E, Pleotelson with uropods, ventral view. Scale = 0.2 mm
156
Figure 28. Pseudotanais {Pseudotanais) sp. P. Paratype, female. A, Epignath. B, Labrum. C,
Labium. D, Left mandible. E, Right mandible. F, Maxillipeds. G, Maxillule, inset shows detail
of setulate spiniform seta (arrow). Scale = 0.2 mm
Figure 29. Pseudotanais {Pseudotanais) sp. P. Paratype, female. A - F, Pereopods 1 to 6,
respectively. D,E,F, - insets show detail of bifid-tipped setae (arrows). Scale = 0.2 mm
158
Taxonomy of Cryptocopidae
Family Cryptocopidae, n. fam.
Diagnosis.
Eyelobes pigmented, rudimentary or absent. Pereon with six free pereonites none of
which are reduced. Pleon with five free pleonites. Antennule with four articles. Antenna with
six articles, second and third articles with simple setae. Mandible pars molaris broad or narrow,
with or without terminal ring of setae. Maxillule palp terminating with two setae and endite
terminating with five to ten spiniform setae. Maxilla rudimentary. Maxilliped bases completely
or partially fused. Maxilliped endites completely separate; with simple setae, cusps, or naked.
Cheliped attached to body via sclerite. Chelae not forcipate. Cheliped carpus with one or two
inferior setae. Cheliped propodus less than 1.5 times longer than carpus. Cheliped fixed finger
with one or two inferior setae. Cheliped dactylus seta present or absent. Marsupium formed by
one or possibly four (Bird and Holdich 1989) pairs of oostegites. Pereopods 4 to 6 merus with
two setae and without fusion of dactylus and unguis to form claw. Pleopods well developed, with
terminal setae only, or rudimentary without setae. Uropod exopods with one or two articles.
Uropod endopods with two articles.
Remarks.
The diagnosis was modified after Sieg (1977), Larsen and Wilson (2002) and Larsen
(2005) from their diagnoses of the family Pseudotanaidae and the subfamily Cryptocopinae which
contained all the members of the new family Crytocopidae. The new family is currently
polyphyletic (Fig. 5) and includes the subfamilies Cryptocopinae, Cryptocopoidinae and
lungentitanainae.
Key to Subfamilies.
1.
Pleopods rudimentary, uropod exopods uni-articulate
Pleopods not rudimentary, uropod exopods bi-articulate
2.
Pleopods well developed, lateral setae present
Cryptocopinae
2
lungentitanainae
159
Pleopods elongate, without lateral setae
Cryptocopoidinae
Subfamily Cryptocopinae Sieg, 1977
Diagnosis.
As for family except: Eyelobes absent. Mandible pars molaris narrow, with bifid tip.
Maxilhile endite terminating with nine spiniform setae. Maxilliped endites with simple setae
only. Cheliped carpus with two inferior setae. Cheliped fixed finger with one inferior seta.
Cheliped dactylus seta absent. Pleopods rudimentary, without setae. Uropod exopods with one
article.
Type genus.
Cryptocope G.O. Sars, 1882
Genera included.
Cryptocope G.O. Sars, 1882
Remarks.
This subfamily was originally erected by Sieg (1977) to contain four mono-specific
genera of Pseudotanaidae with four antennular articles, completely separate maxilliped endites
and no claw-like pereopod dactylus - all plesiomorphic characters. Two genera, Iungentitanais
and Parahmgentitanais, were placed in the subfamily with reservations owing to the unknown
nature of the marsupium. Two additional mono-specific genera, Latitanais and Curtichelia, were
placed in Crytocopinae by Larsen and Wilson (2002) based on a cladistic analysis of the
Paratanaoidea. In the current revision, a single species, Crytocope abreviata, remains in the
subfamily.
160
Cryptocope abreviata (G.O. Sars, ! 882)
Synonymy.
Tanais abbreviatus G.O. Sars, 1886
Cryptocope abreviata Greve, 1965: 45
Cryptocope abreviata Sieg 1977: 22
Diagnosis.
As for subfamily with the following: Antennule first article twice as long as broad, third
article about half as long as broad, distal article about twice as long as broad. Antenna second
article subequal in length to third article, fourth article 3.5 times longer than broad and fifth
article nearly 3.0 times longer than broad. Cheliped propodus about twice longer than broad,
dactylus greater than half length of propodus and narrower than fixed finger. Pereopod 1 basis
6.0 times longer than broad, with simple setae; ischium and merus with one seta, carpus and
propodus with two setae and dactylus without setae; propodus about 5.0 times longer than broad
and dactylus with unguis subequal to length of propodus. Pereopods 2 and 3 basis without setae,
ischium with one seta and merus with two setae; dactylus with unguis subequal in length to
propodus. Pereopod 4 basis 6.0 times longer than broad. Pereopods 4 to 6 basis without setae
and ischium with two equal setae. Uropod exopods half length of endopods.
Type Locality.
North Atlantic off Norway.
Depth Range.
60-680 m.
Distribution.
Known only from the coasts of Norway.
Remarks.
161
Sieg examined material collected by Greve (!965: 45) from near the type locality
consisting of nine females of which were three ovigerous or preparatory. The original type
material of Sars is apparently lost.
References.
Sieg (1977): 22-26, Figs. 12-15.
Subfamily Crytocopoidinae, n. subfam.
Diagnosis.
As for family except: Eyelobes absent. Mandible pars molaris broad with terminal ring
of short denticles. Maxillule endite terminating with eight or nine spiniform setae. Maxilliped
bases not completely fused. Maxilliped endites with simple setae only. Cheliped carpus with two
inferior setae. Cheliped fixed finger with two inferior setae. Pleopod rami elongate with
terminal and/or subterminal setae. Uropod exopods with two articles.
Type genus.
Cryptocopoides Sieg (1976)
Genera included.
Cryptocopoides Sieg (1976)
Remarks.
The three species of this subfamily appear as paraphyletic on the cladogram in Fig. 1 and
in subsequent cladograms depicting character traces with some character states (e.g., pleopod
development) that lie intermediate between the more ancestral lungentititanainae and the
Crytocopinae (character 63, Fig. 87). In contrast, the subfamily shows an ancestral trait with
partial separation of the maxilliped bases (character 23, Fig. 49). The two most recently described
species, C. pacificus and C. sp. A form a well supported clade in a more ancestral position than
the third member of the genus, C. arcticus.
162
Genus Cryptocopoides Sieg (1976)
Diagnosis.
As for subfamily.
Species Included.
Cryptocopoides arcticus (Hansen, 1887) - type species
Cryptocopoides pacificus McLelland, 2007
Cryptocopoides sp. A
Cryptocopoides articus (Hansen, 1887)
Synonymy
Crytocope artica Hansen, 1887:209
Cryptocope antarctica Vanhoffen, 1914:482
Cryptocopoides arctica (= arcticus) Sieg, 1977: 17
Diagnosis.
As for subfamily with the following: Antennule first article about twice as long as broad,
third article about as long as broad, distal article 3.5 times as long as broad. Antenna second
article subequal in length to third article, fourth article 3.5 times longer than broad and fifth
article about 6.0 times longer than broad. Maxilliped bases fused except for distal notch.
Maxillule endite with eight terminal spiniform setae. Cheliped propodus about 2.5 times longer
than broad, dactylus greater than half length of propodus, subequal in width to fixed finger and
lacking a proximal seta. Pereopod 1 basis 7.0 times longer than broad, with simple setae; ischium
and merus with one seta, carpus and propodus with three setae and dactylus without setae;
propodus about 4.0 times longer than broad and dactylus with unguis distinctly shorter than
propodus. Pereopods 2 and 3 basis with broom setae, ischium and merus with one seta; dactylus
with unguis subequal in length to propodus. Pereopod 4 basis 6.0 times longer than broad.
163
Pereopods 4 to 6 basis without setae and ischium with one seta. Pleopod rami elongate with
terminal setae only. Uropod exopods subequal to length of endopods.
Type locality.
Kara Sea, Arctic Ocean
Depth Range.
103 m (North Atlantic); 4664-5631 m (Antarctica).
Distribution.
North Atlantic: North Sea, Arctic Ocean; South Atlantic: Antarctica; Pacific: Japan.
Remarks.
Sieg (1977) based his description and synonymy on a combination of specimens from
two largely separated areas: Cryptocope arctica collected by Hansen (1913) from the north
Atlantic and Cryptocope antarctica collected off Antarctica by Vanhoffen (1914). He determined
them to be identical except for some "small variations" and synomized them under one species,
Cryptocopoides arcticus. Hopefully, additional specimens from both polar regions will be reexamined in greater detail someday to determine if these "variations" could be interpreted as
representing two separate species.
References.
Sieg (1977): 17, Figs. 9-11, 15; Vanhoffen (1914): 482, Fig. 19a,b.
Cryptocopoides pacificus McLelland, 2007
Diagnosis.
As for subfamily and C. arcticus with the following: Antennule first article about twice
as long as broad, third article about as long as broad, distal article 3.5 times as long as broad.
164
Antenna second article longer than third article, fourth article 3.5 times longer than broad and
fifth article about twice as long as broad. Maxilliped bases about half fused. Maxillule endite
with eight terminal spiniform setae. Cheliped, dactylus possessing a proximal seta. Pereopod 1
basis 9.0 times longer than broad, with simple setae; ischium and merus with one seta, carpus
with four setae, propodus with three setae and dactylus without setae; propodus about 4.0 times
longer than broad and dactylus with unguis subequal in length to propodus. Pereopods 2 and 3
basis with simple setae, ischium and merus with one seta; dactylus with unguis subequal in length
to propodus. Pereopod 4 basis 5.0 times longer than broad. Pereopods 4 to 6 basis with simple
setae, ischium with two unequal setae, carpus with additional spatulate setae and propodus with
broom seta. Pleopod rami elongate with terminal and subterminal setae. Uropod exopods
subequal to length of endopods.
Type Locality.
Kamchatka Trench in the North Pacific off Japan.
Depth Range.
3145-5484 m.
Distribution.
North Pacific: Kamchatka Trench off Japan.
Remarks.
This species differs further from C. arcticus by (1) having antenna article 2 with two
distal setae instead of one, (2) having a pair of basal setae and pair of submarginal setae on the
maxilliped endites, (3) having a small proximal seta on cheliped dactylus, and (4) having two
ischium setae and spatulate carpal setae present on pereopods 4 and 5.
Reference.
McLelland (2007): 88-93, Figs. 1-3.
Cryptocopoides sp. A, n. sp. (Figs. 30 - 32)
165
Diagnosis.
As for subfamily and C. pacificus except: Pleopods with terminal setae only. Maxillule
endites with nine terminal spiniform setae. Pereopod dactylus with unguis distinctly shorter than
propodus. Four apical setae on pleotelson.
Type locality.
Off Kiawah Island, South Carolina, 31° 42.04N, 78° 49.42W, 03 August 2003, 554 m.
Depth Range.
All - 875 m.
Distribution.
North Atlantic: off east coast of U.S.
Material examined.
Holotype, non-ovigerous female, 1.65 mm., MRRI sample Tl 0031019 collected by the
Johnson Sea-Link submersible using a suction device. Paratypes: MRRI submersible collection
T10041060 off Jacksonville Beach, Florida, 30° 16.56N, 79° 20.38W, 836 m, 30 August 2004 - 2
non-ovigerous females, 1.4 mm; 1 non-ovigerous female, 1.8 mm, dissected and slide mounted.
Johnson Sea-Link submersible collection 2004-0315 (Dive no. 3470) off Jacksonville, Florida,
30° 63.2500N, 79° 73.4722 W, 875 m, 30 August 2004 - 1 female, fragmented. Museum
specimens listed as Cryptocope sp. A: 1 female (USNM 174730), off Cumberland Island, GA,
30° 54' 04 N, 79° 43' 50 W, 422 m, collected by Texas Instruments, 25 November 1977; 1
neuter/subadult female (USNM 174731), off Jacksonville, FL, 30° 23' N, 80° 10' W, 149 m,
collected by Texas Instruments, 1 March 1977.
Description
Based on non-ovigerous females. Body 1.4-1.8 mm in length, 4.4 times longer than
broad.
166
Cephalothorax (Fig. 30A) longer than pereonites 1-3, trapezoidal, becoming narrow
anteriorly, slightly wider than pereon, with pair of antero-lateral setae, 25.1 percent total length;
eyelobes absent.
Pereon 51.9 percent total length; no pereonites abreviated; each pereonite with pair of
dorso-lateral and ventro-lateral setae emerging from anterior margin (pereonite 1 with three
additional pairs of ventrolateral setae).
Pleon (Fig. 30B) shorter than pereonites 5+6, 15.2 percent total length, pleonites 3-5
with pair of dorso-lateral setae (pleonite 5 with additional pair of extremely long ventro-lateral
setae).
Pleotelson (Fig. 30F) shorter than pleonites 3-5, 7.8 percent total length, apex slightly
produced ventrally, with one pair of distolateral setae or four apical setae
Antennule (Fig. 30D) 9.5 percent of body length, with four articles; first article 47.3
percent of total length, 1.6 times longer than broad, with three short medial simple setae, one
short distal simple seta, one long distal simple seta and two distal broom setae; second article 0.9
times longer than broad, with one long distal simple seta and two distal broom setae; third article
0.9 times longer than broad, with two long distal simple setae and one distal broom seta; distal
article 3.8 times longer than broad, terminating with four simple setae and two aesthetascs.
Antenna (Fig. 30E) with six articles, 88.1 percent length of Al; second article 1.7 times
longer than broad, 1.6 times longer than third article, both articles distally with long, slender
spiniform seta; fourth article 3.6 times longer than broad, with two medial setules, six short distal
simple setae (one thick), and one long distal simple seta; fifth article 2.7 times longer than broad,
distally with one long simple seta; sixth article small, terminating with two aesthetascs and three
simple setae (one long, two short).
Mouthparts. Labrum (Fig. 3IE) cap-shaped, heavily setose with setulate lateral margins.
Mandibles (Figs. 31A,B) with distal margins denticulate, right lacinia mobilis absent, left lacinia
mobilis well developed and denticulate, pars molaris broad, with six blunt terminal teeth and
167
three short marginal setae. Labium not observed. Maxillule (Fig. 31D) endite terminating with 9
spiniform setae, including 1 with expanded, slightly bifid tip and two accessory setae. Maxilla
not observed. Maxilliped (Fig. 31C) bases about half fused, with seta near each palp articulation;
endites completly separate, each with two unequal setae; palp with four articles; article 1 without
setae; article 2 with one medio-distal extremely long simple seta, one inner-edge simple seta, one
inner-edge setulose spiniform seta, and one outer-edge simple seta; article 3 with one short distal
simple seta, two inner-edge setulose spiniform setae, and one inner-edge simple seta; article 4
with one small outer-edge simple seta, four inner-edge setulose spiniform setae, and one inneredge simple seta. Epignath not observed.
Cheliped (Fig. 30C) chela not forcipate; sclerite well developed; basis 1.7 times longer
than broad, with medial superior seta; merus triangular, with inferior seta; carpus 1.6 times longer
than broad, with two unequal medial inferior setae, one distal superior seta and one medial
superior seta; propodus 2.6 times longer than broad, slightly longer than carpus, palm with one
long and two short comb setae, plus row of several small setules; fixed finger with two unequal
inferior setae, three superior setae, terminal unguis and subterminal inner-margin notch which
receives tip of dactlyus; dactylus 45 percent of propodus length, width subequal to fixed finger,
with proximal spiniform seta and terminal unguis.
Pereopod 1 (Fig. 32A) coxa with seta; basis 8.9 times longer than broad, with proximal
superior seta; ischium with small seta; merus 2.2 times longer than broad, distally with minute
superior seta and short inferior seta; carpus 3.8 times longer than broad, 1.5 times length of
merus, distally with three short simple setae (one inferior, one superior, one anterior) and one
minute superior spiniform seta; propodus 5 times longer than broad, distally with long superior
spiniform seta and short inferior spiniform seta; dactylus without seta, length with unguis 60
percent length of propodus.
Pereopod! (Fig. 32B) coxa with seta; basis 8.8 times longer than broad, with proximal
superior broom seta and proximal superior seta; ischium with short inferior seta; merus 2.5 times
168
longer than broad, distally with long inferior spiniform seta; carpus 3.9 times longer than broad,
about 1.5 times length of merus, distally with one short superior simple seta, one long superior
spiniform seta and two long inferior spiniform setae; propodus 4.9 times longer than broad, with
one short distal superior spiniform seta and one long distal inferior spiniform seta; dactylus
without setae, length with unguis 70 percent length of propodus.
Pereopod 3 (Fig. 32C) basis with medial inferior broom seta (proximal superior broom
seta lacking); otherwise similar to P2.
Pereopod 4 (Fig. 32D) coxa with seta; basis 5.7 times longer than broad, with two medial
inferior broom setae; ischium with two short setae; merus 2.4 times longer than broad, distally
with two long inferior spiniform setae; carpus 5.1 times longer than broad, 1.2 times longer than
merus, distally with one short spatulate seta and four long spiniform setae; propodus 5.1 times
longer than broad, 1.4 times longer than carpus, distally with one long superior terminal
spiniform seta, two long inferior spiniform setae, one superior broom seta and setulose terminal
margin at emergence of dactylus; dactylus not fused with unguis, length with unguis about 75
percent length of propodus.
Pereopod 5 (Fig. 32E) propodus with shorter, stouter superior terminal spiniform seta,
otherwise similar to P4.
Pereopod 6 (Fig. 32F) basis with medial superior broom seta (two inferior broom setae
lacking); ischium with single short seta; carpus distally with short simple seta (instead of
spatulate seta), propodus with additional terminal spiniform seta and lacking broom seta,
otherwise similar to P4.
Pleopods (Fig. 30G) rami elongate, with terminal setae only; endopod about 2/3 length of
exopod, with four terminal setae; exopod with seven terminal setae.
Uropods (Fig. 30F) exopod with two articles, 75 percent length of endopod, proximal
article slightly less than half length of ramus, with one distal simple seta, distal article with one
long simple seta; endopod with two articles, proximal article about half length of ramus, with one
169
broom seta, distal article with five long simple setae (one subterminal, four terminal) and one
broom seta.
Remarks.
Cryptocopoides sp. A is the third species described for the genus. The first, C. arcticus
(Hansen, 1887), is well known from northern Atlantic waters and was considered by Sieg (1977)
to have a bi-polar distribution (see remarks for C. arcticus). Recently McLelland (2007)
described an additional species, C. pacijicus, from the north Pacific Ocean. Of the three species
of Cryptocopoides, C. sp. A has more features in common with C. pacificus than with C. arcticus,
since both have a greater degree of setation on body segments, mouthparts and pereopods and
have a similar cephalothorax length which is comparatively shorter than that of C. arcticus. In
addition, both have a distinctive spatulate carpal seta on pereopods 4 and 5, a small proximal seta
on the cheliped dactylus, a superior seta on the cheliped basis, and a pair of setae on the fused
maxilliped bases, all of which appear to be lacking in C. arcticus. Furthermore, both species
differ from C. arcticus by having two (vs. one) sub-marginal setae on the maxilliped endites, four
or five (vs. two) distal setae on pereopods 2 and 3, and a distinctive pair of long setae on the
ventro-lateral margins of pleonite 5 (lacking in C. arcticus). Cryptocopoides sp. A shares
similarities with C. arcticus that distinguish it from C. pacificus including: (1) antenna article 2
with a single spiniform seta rather than two, (2) pereopod dactylus-unguis lengths which are
distinctly shorter than the propodus (sub-equal in C. pacificus), and (3) the lack of a sub-terminal
endopod seta on the pleopods. The new species is unique among its congenitors by having four
rather than two apical setae on the pleotelson and nine terminal spiniform setae on the maxillule
endite rather than eight.
170
Figure 30. Cryptocopoides sp. A. Holotype female. A, Dorsal view. B, Lateral view. Paratype
female. C, cheliped, inner face; D, antennule. E, Antenna. F, Pleotelson and uropods, ventral
view. G, Pleopods. Scale bars: A-B = 0.5mm, C-G = 0.2mm
171
Vv;'"''1'';1'^
Figure 31. Cryptocopoides sp. A. Paratype, female. A, left mandible; B, right mandible; C,
maxilliped; D maxillule endite, inset shows spiniform seta with inflated tip; E, labrum. Scale
0.2mm
172
Figure 32. Cryptocopoides sp. A. Paratype, female. A-F, pereopods 1-6 respectively. Inset at D
shows detail of spatulate seta indicated by arrow. Scale = 0.2mm.
173
Subfamily lungentitanainae, n. subfam.
Diagnosis.
As for family except: Antennule third article 1.5 to 2.5 times longer than broad.
Antennule distal article and antenna fourth article both greater than 4.0 times longer than broad.
Mandible pars molaris broad, truncate, with terminal ring of denticles. Maxilliped bases
completely fused. Pleopods well developed, with terminal and lateral setae. Uropod exopods
with two articles.
Type Genus.
Iungentitanais Sieg, 1977
Genera Included.
Iungentitanais Sieg, 1977
Paraiungentitanais Sieg, 1977
Latitanais Kudinova-Pasternak, 1987
Curtichelia Kudinova-Pasternak, 1987
Remarks.
This subfamily is a provisional repository for four mono-specific genera with
questionable affinities. Nonetheless, the group formed a fairly well supported monophyletic
clade, ancestral to the other Cryptocopidae subfamilies, as seen in Figure 1 and in subsequent
character trace figures.
Iungentitanais primitivus Sieg, 1977
Synonymy
Mortenseniaprimitivus Sieg, 1973: 173-177; Kudinova-Pasternak, 1975: 223
Diagnosis
As for subfamily except: Eyelobes present, with pigment. Antenna fourth article six to
8.0 times longer than broad. Maxillule endite terminating with five spiniform setae. Maxilliped
174
endites with cusps and setae. Cheliped carpus greater than 4.0 times longer than broad, with one
inferior seta. Cheliped fixed finger with one inferior seta. Cheliped propodus greater than 4.0
times longer than broad. Cheliped dactylus less than half length of propodus, narrower than fixed
finger and bearing a proximal seta. Pereopod 1 basis without setae, ischium with 1 seta, merus
without setae, carpus with two setae, propodus with three setae, and dactylus with one short seta.
Pereopods 4 to 6 basis without setae and ischium with one seta. Uropod exopod half length of
endopod.
Type Locality
North Atlantic: Saint Thomas in the tropical West Indies.
Depth Range
18-33 m.
Distribution
Atlantic West Indies, Caribbean, tropical Gulf of Mexico.
Remarks
This species is characterized by its long, slender chelipeds, pigmented eyes, and five
terminal spines on the maxillule endite. The latter characters separate it from its closest relative
Paraiungentitanais longidigitatus. Mandibles were described from additional specimens collected
in the Florida Keys since those from the holotype were lost (Sieg and Heard 1988).
References
Sieg (1977): 12-15, Figs. 5-7, 15; Sieg and Heard (1988): 48, Figs. 2-4, 12.
Paraiungentitanais longidigitatus (Kudinova-Pasternak, 1975)
Synonymy
175
Mortensenia longidigitara Kudinova-pasternak, 1975; Paraiungenlitanais longidigitatus
Sieg, 1977:15
Diagnosis
As for subfamily and Iungentitanais except: Eyelobes rudimentary, without pigment.
Maxillule endite terminating with nine spiniform setae. Cheliped carpus less than 4.0 times
longer than broad, with two inferior seta. Cheliped fixed finger with two inferior setae. Cheliped
dactylus broader than fixed finger. Uropod exopod 1/3 length of endopod.
Type Locality
South Atlantic: off Falkland Islands.
Depth Range
1660 m.
Distribution
Southwest Atlantic Ocean off the Falkland Islands.
Remarks
The original description provided only limited information. Characters for periopods and
pleopods were assumed to be the same as those for Iungentitanais primitivus; indeed these
similarities are stated by both, Kudikova-Pasternak and Sieg. This genus, according to Sieg, is
distinguished from Iungentitanais mainly by virtue of its nine terminal maxillule endite spines as
opposed to five for Iungentitanais. Sieg further mentioned that since the marsupium is unknown,
placement of this species in Pseudotanaidae is questionable.
References
Kudinova-Pasternak(1975): 223-225, Fig. 16; Sieg (1977): 15-17, Figs. 8, 15.
Latitanais beklemishevi Kudinova-Pasternak, 1987
Diagnosis
176
As for subfamily except: Eyelobes rudimentary, without pigment. Antenna fourth article
between four and 6.0 times longer than broad. Maxillule endite terminating with nine spiniform
setae. Maxilliped endites without setae. Cheliped carpus less than 4.0 times longer than broad,
with two inferior setae. Cheliped fixed finger with one inferior seta. Cheliped propodus less than
2.5 times longer than broad. Cheliped dactylus greater than 60% length of propodus, subequal in
width to fixed finger and lacking proximal seta. Pereopod 1 basis without setae, ischium without
setae, merus with two setae, carpus with two setae, propodus with one seta, and dactylus without
setae. Pereopods 4 to 6 basis without setae and ischium with two equal setae. Uropod exopod
length subequal to that of endopod.
Type Locality
Indian Ocean: Madagascar Basin
Depth Range
5040-5060 m.
Distribution
Indian Ocean from the type locality.
Remarks
The original description in Russian was very cursory and several appendages were not
illustrated. This species, originally a member of the family Leptognathidae, was placed, along
with Curtichelia expressa, within the Cryptocopinae by Larsen and Wilson (2002) as a result of a
cladistic analysis conducted on genera of Paratanaoidea. It is notable among cryptocopids by
having well developed pleopods (terminal and lateral setae), a maxillule endite with nine terminal
spines, and subequal uropod rami.
References
Kudinova-Pasternak (1987): 29-30, Fig. 1.
177
Curtichelia expressa Kudinova-Pasternak, 1987
Diagnosis
As for the subfamily except: Eyelobes lacking. Antenna fourth article between four and
6.0 times longer than broad, fifth article nearly 7.0 times longer than broad. Maxillule endite
terminating with nine spiniform setae. Maxilliped endites without setae. Cheliped carpus less
than 2.0 times longer than broad, with two inferior seta. Cheliped fixed finger with two inferior
setae. Cheliped propodus less than 2.5 times longer than broad. Cheliped dactylus greater than
60% length of propodus, broader than fixed finger and lacking proximal seta. Pereopod 1 basis
with one simple seta, ischium with one seta, merus with two setae, carpus with two setae,
propodus with two setae, and dactylus with one short seta. Pereopods 4 to 6 basis with two
broom setae and ischium with two equal setae. Uropod exopod 75 percent length of endopod.
Type Locality
Indian Ocean: off Madagascar.
Depth Range
4987-5025 m.
Distribution
Indian Ocean off Madagascar.
Remarks
As for Latitanais beklemishevi, the original description in Russian was very cursory and
some appendages were incompletely rendered. This species, originally a member of the family
Leptognathidae, was placed, along with Latitanais beklemishevi, within the Cryptocopinae by
Larsen and Wilson (2002) as a result of a cladistic analysis conducted on genera of Paratanaoidea.
It is distinguished by its short, massive chelipeds and unusually long mandibles. An atypical
arrangement of the maxillule endite terminal setation is seen in the "10 or 11 terminal spines" of
178
which nine are thick and one has an accessory tooth. The author is likely referring to the usual
nine spines with accessory setae.
Reference
Kudinova-Pasternak (1987): 34-35, Fig. 4.
Key to Species of Cryptocopidae and Pseudotanaidae
1. Antennule with four articles; maxilliped endites completely separate
Family Cryptocopidae
2
— Antennule with three articles; maxilliped endites partially or completely fused
Family Pseudotanaidae
2.
Pleopods rudimentary; uropod exopods uni-articulate
Subfamily Cryptocopinae
3.
Cryptocope abreviata
Pleopods not rudimentary; uropod exopods bi-articulate
3
Pleopods well developed, lateral setae present
Subfamily iungentitanainae
4
Subfamily Cryptocopoidinae
7
Pleopods elongate, without lateral setae
4.
9
Setae present on basis of pereopods 1 and 6; maxillule endite terminating with ten spiniform
setae
Curtichelia expressa
Setae absent on basis of pereopods 1 and 6; maxillule endites with nine or less terminal
spiniform setae
5.
Pereopod 1 ischium and dactylus without setae
5
Latitanais beklemishevi
Pereopod 1 ischium and dactylus with setae
6.
6
Cheliped carpus and fixed finger each with one inferior seta, maxillule endite terminating ....
with five spiniform setae
Iungentitanais primitivus
Cheliped carpus and fixed finger each with two inferior setae; maxillule endite terminating
with nine spiniform setae
...Paraiungentitanais longidigitatus
179
Cheliped dactylus without proximal seta; pereopods 4 to 6 ischium with one seta
Cryptocopoides articus
Cheliped dactylus with proximal seta; pereopods 4 to 6 ischium with two setae
8
Pereopods 1 to 3 dactylus with unguis distinctly shorter than propodus; maxillule endites
terminating with nine spiniform setae
Cryptocopoides sp. A
Pereopods 1 to 3 dactylus with unguis subequal to length of propodus; maxillule endites
terminating with eight spiniform setae
Modified blade-like carpal setae absent on pereopods
Cryptocopoides pacificus
Subfamily Akanthinotanainae 10
Modified blade-like carpal setae present on pereopods
19
Pereopod 1 propodus with two setae; maxilliped endites with cusps and setae present
subgenus Guilleitanais
Akanlhinotanais (G) guillei
Pereopod 1 propodus with one or no setae; maxilliped endites with or without setae but no
cusps
subgenus Akanthinotanais, s.s
11
Eyes absent
12
Eyes present, with pigment
14
Pereopods 2 and 3 dactylus with unguis subequal to length of propodus; uropod exopods
with two articles
Akanthinotanais (A) gaussi
Pereopods 2 and 3 dactylus with unguis half to 2/3 length of propodus; uropod exopods with
one article
13
Pereopod 4 basis without setae; pereopod 1 basis about 10.5 times longer than broad
Akanthinotanais {A) longipes
Pereopod 4 basis with broom seta; pereopod 1 basis about 5.0 times longer than broad
Akanlhinotanais (A) similis
Uropod exopods and endopods each with one pseudo-articulated article
Akanthinotanais (A) makrothrix
180
Uropod exopods and endopods each with two articles
15
Pereopod 1 merus without setae
16
Pereopod 1 merus with one seta
18
Uropod exopods half length of endopods; cephalothorax subequal to pereonites 1 to 3
Akanthinotanais (A) malayensis
Uropod exopods 2/3 to subequal length of endopods; cephalothorax longer than pereonites 1
to 3
17
Cheliped dactylus with proximal seta; pereopod 1 ischium with two setae
Akanthinotanais (A) gerlachi
Cheliped dactylus without proximal seta; pereopod 1 ischium without setae
Akanthinotanais {A) scrappi
Antennule distal article about 9.5 times longer than broad; cheliped dactylus without
proximal seta
Akanthinotanais {A) siegi
Antennule distal article 6.0 times longer than broad; cheliped dactylus with proximal seta
Akanthinotanais (A) mortenseni
Blade-like carpal setae on pereopods 4 to 6; cheliped with two setae on fixed finger
Subfamily Parapseudotanainae
20
Blade like carpal setae on pereopods 2 to 6; cheliped with one or no setae on fixed finger
Subfamily Pseudotanainae
21
Uropod endopods with two articles; antenna article 2 longer than article 3
Parapseudotanais abyssalis
Uropod endopods with one article; antenna article 2 shorter than article 3
Parapseudotanais sp. A
Chela strongly forcipate with serrate incisive margins; spatulate-tipped setae present on
merus of pereopods 2 and 3 and carpus of pereopods 4 to 6
181
Subgenus Mystriocentrus
22
Chela forcipate or not but without serrate incisive margins; without spatulate-tipped setae on
pereopods
Subgenus Pseudotcmais
23
Pereonite 3 markedly shorter than pereonite 2; cheliped dactylus without proximal seta
Pseudotcmais (M) serratus
Pereonite 3 subequal in length to pereonite 2; cheliped dactylus with proximal seta
Pseudotcmais (M) sp. A
Pigmented eyelobes present (oculate)
24
Eyelobes absent or without pigment (non-oculate)
28
Pereopod 1 merus without setae; cheliped dactylus narrower than fixed finger
25
Pereopod 1 merus with one seta; cheliped dactylus width subequal to fixed finger
26
Pereopod 3 disproportionately smaller than pereopod 2; uropod rami bi-articulate
Pseudotanais (P) unicus
Pereopod 3 about same proportions as pereopod 2; uropod rami uni-articulate
Pseudotanais (P) borceai
Pereopods 1 to 3 ischium with two setae, pereopods 2 and 3 basis without setae
Pseudotanais (P) mediterraneus
Pereopods 1 to 3 ischium with one seta; pereopods 2 and 3 basis with broom setae
27
Cephalothorax subequal in length to pereonites 1 to 3; pereopod 1 basis with broom seta
Pseudotanais (P) oculatus
Cephalothorax longer than pereonites 1 to 3; pereopod 1 basis without setae
Pseudotanais (P) mexikolpos
Chelae strongly forcipate
29
Chelae not forcipate, or only weakly so
34
Pereopod 1 carpus with one seta; maxilliped endites fused except for notch on distal margin..
182
30
Pereopod 1 carpus without setae; maxiliiped endites completely fused, with smooth distal
margin
32
Cheliped carpus with two inferior setae; pereopods 4 to 6 ischium with two setae
31
Cheliped carpus with three inferior setae; pereopods 4 to 6 ischium with one seta
Pseudotanais (P) abyssi
Pereopods 2 and 3 carpus blade-like setae greater than half length of propodus; pereopods 4
and 5 propodus without broom seta
Pseudotanais (P) vulsella
Pereopods 2 and 3 carpus blade-like setae less than half length of propodus; pereopods 4 and
5 propodus with broom seta
Pseudotanais (P) falciculata
Pleopods present; pereopods 2 and 3 merus with one seta
33
Pleopods absent; pereopods 2 and 3 merus with two setae
Pseudotanais (P)forcipatus
Cheliped fixed finger without inferior seta; pereopod 1 merus with two setae
Pseudotanais (P)jonesi
Cheliped fixed finger with one inferior seta; pereopod 1 merus with one seta
Pseudotanais (P) californiensis
Antenna articles 2 and 3 with short, stout spiniform setae
38
Antenna articles 2 and 3 with setae otherwise (simple or spiniform)
35
Pereopods 2 and 3 carpus with blade-like seta greater than half the propodus length
36
Pereopods 2 and 3 carpus with blade-like setae less than half the propodus length
37
Uropod exopods with one article; pereopod 1 dactylus with unguis subequal to length of
propodus
Pseudotanais (P) longisetosus
Uropod exopods with two articles; pereopod 1 dactylus with unguis distinctly shorter than
propodus
Pseudotanais (P) corollatus
Pleopods absent; pereopod 1 dactylus with unguis distinctly longer than propodus
183
Pseudotanais (P) lilljeborgi
Pleopods present; pereopod 1 dactylus with unguis subequai to length of propodus
Pseudotanais (P) inflatus
Cephalothorax longer than pereonites 1 to 3
39
Cephalothorax subequai or shorter than pereonites 1 to 3
47
Maxilliped endites nearly fused except for notch on distal margin
40
Maxilliped endites fused 1/3 to 2/3 length
42
Uropod exopodswith one article; pereopod 1 dactylus without seta
Pseudotanais (P) colonus
Uropod exopods with two articles; pereopod 1 dactylus with short seta
41
Pereopod 1 merus with three setae and carpus with one seta; cheliped dactylus narrower than
fixed finger
.-. Pseudotanais (/') nordenskioldi
Pereopod 1 merus with one seta and carpus with two setae; cheliped dactylus subequai in
width to fixed finger
Pseudotanais (P) baresnauti
Antenna second article subequai in length to third article; pereopods 2 and 3 merus with two
setae
43
Antenna second article shorter than third article; peropods 2 and 3 merus with one seta
45
Cheliped dactylus narrower than fixed finger; pereopod 1 merus with one seta
Pseudotanais (P) denticulatus
Cheliped dactylus subequai in width to fixed finger; pereopod 1 merus without setae
44
Pereopod 1 carpus with four setae, propodus with one seta; pereopods 4 and 5 basis with
broom setae
Pseudotanais (P) spicatus
Pereopod 1 carpus with two setae, propodus without setae; pereopods 4 and 5 basis without
setae
Pseudotanais (P) vitjazzi
Pereopods 2 and 3 carpus with blade-like setae less than half length of propodus; pereopod 1
184
ischium without setae
Pseudotanais (P) sp. A
Pereopods 2 and 3 carpus with biade-like setae greater than half length of propodus;
pereopod 1 ischium with seta
46
Cheliped carpus with three inferior setae; pereopod 1 propodus without setae
Pseudotanais (P) macrocheles
Cheliped carpus with two inferior setae; pereopod 1 propodus with seta
Pseudotanais (P) sp. C
Uropod exopods with one article; pereopods 2 and 3 merus with one seta
Pseudotanais (P) sp. P
Uropod exopods with two articles; pereopods 2 and 3 merus with two setae
Pereopod 1 ischium without setae; pereopods 4 to 6 ischium with one seta
Pereopod 1 ischium with one seta; pereopods 4 to 6 ischium with two setae
48
49
50
Pereopod 1 carpus with three setae; pereopods 5 and 6 carpus each with abnormally long
superior seta, subequal to length of propodus
Pseudotanais (P) spatula
Pereopod 1 carpus with two setae; pereopods 5 and 6 without abnormally long setae
Pseudotanais (P) scalpellum
Pereopod 1 dactylus seta absent; pereopods 4 to 6 ischium with two unequal-length setae ..51
Pereopod 1 dactylus seta present; pereopods 4 to 6 ischium with two setae of equal length. 52
Antenna second article shorter than third article; pereopod 1 merus with two setae
Pseudotanais (P) sp. O
Antenna second article subequal in length to third article; pereopod 1 merus with one seta
Pseudotanais (P) nipponicus
Peropods 2 and 3 carpus with blade-like setae about half length of propodus; pereopod 1
propodus without setae, dactylus with unguis distinctly shorter than propodus
Pseudotanais (P) affinis
185
Pereopods 2 and 3 carpus with blade-like setae greater than half length of propodus;
pereopod 1 propodus with two setae, dactylus with unguis subequal in length to propodus
Pseudotanais (P) longispinus
APPENDIX A
Gulf of Mexico Station Data for new species of Pseudotanaidae*
Station
Postition
Date
AC 1-2
26° 23.2813 TM
94° 33.2633 'W
26°23.5394'N
94°33.4048'W
27M2.2050'N
91°24.1077'W
27°12.1525'N
9r24.3131'W
26°33.0774'N
92°12.9990'W
26°33.3750'N
92°13.4716'W
26°32.9491'N
92°13.1357'W
26°09.9750'N
91°43.9772'W
26°09.9750'N
91° 43.9772'W
27°47.8832'N
91°28.1670'W
28°03.6046'N
90°14.9329*W
28°03.5783'N
90°14.9483'W
26°22.9752'N
89°14.4854'W
27°27.5640'N
89'47.1391'W
27°27.6103'N
89°46.7720'W
27°27.1450'N
89°46.5588'W
27°43.6967'N
89°58.7782'W
27°44.1045'N
89°59.0927'W
27°43.9713'N
89°58.6211W
28°33.1285'N
88°34.2763'W
28°27.0646'N
89° 40.3563'W
28°27.0726'N
89°40.2206'W
28°13.4727'N
19 May
2000
19 May
2000
6 May
2000
6 May
2000
12 May
2000
12 May
2000
12 May
2000
10 May
2000
10 May
2000
18 June
2001
30 May
2000
30 May
2000
2 June
2000
31 May
2000
20 June
2000
27 July
2000
30 May
2000
16 June
2001
30 May
2000
5 June
2001
17 June
2000
17 June
2000
4 June
AC 1-3
Bl-1
Bl-5
B2-1
B2-2
B2-3
B3-2
B3-3
BH-5
Cl-1
CI-2
CI 2-2
C4-1
C4-2
C4-3
C7-1
C7-2
C7-2
HP-3
MT2-1
MT2-2
MT3-1
Depth
PspA
PspC
PspO
fll
2450
2
2480
2260
2
6
2254
2
1
2630
3
3
2630
4
1
2625
8
1
6
2580
2650
1
533
4
2
334
5
336
5
2920
2
1455
2
3
1452
2
1
1463
2
1080
4
1045
2
1
5
1
2
1070
1562
676
11
677
16
980
1
Station
MI3-1
MT3-2
MT3-5
MT4-2
MT5-3
MT6-4
NB2-2
NB3-3
NB5-2
NB5-3
RW1-2
RW1-3
RW2-1
RW2-2
RW2-3
RW3-1
RW3-3
RW3-5
RW4-1
RW4-2
RW5-1
RW5-2
RW5-3
RW6-1
Postition
Date
89'30.7579'W
28°13.2246'N
89°29.7679'W
28°13.1533'N
89°29.7860'W
28°13.2624'N
89°30.5530'W
27°49.7018'N
89°09.8829'W
27°20.0759'N
88°39.7337'W
26°59.6643'N
88°00.6912'W
27°08.2040'N
91°59.9207'W
26°32.9725'N
91°49.1853'W
26°15.0855'N
91°12.7524'W
26°14.8501'N
91M2.7500'W
27°29.9333'N
96°00.2164'W
27°30.0733'N
95°59.9273'W
27M5.2435TSI
95°44.6186'W
27°15.2852'N
95°44.6402'W
27°14.9757'N
95°44.8287'W
27° 00.2983'N
95°30.1920'W
27°00.1881'N
95°29.7639'W
27°00.5014'N
95°29.5417'W
26° 44.8861'N
95° 14.7680'W
26°44.9468'N
95° 14.6826'W
26°30.2819'N
95°00.1866'W
26° 30.0261 T^
95°00.1315'W
26°30.0206'N
95° 00.0512'W
25° 59.9222'N
94 29.7608'W
2001
16 June
2000
16 June
2000
4 June
2001
15June
2000
4 June
2000
13 June
2001
7 May
2000
8 May
2000
9 May
2000
9 May
2000
23 May
2000
23 May
2000
22 May
2000
22 May
2000
22 May
2000
22 May
2000
22 May
2000
22 May
2000
21 May
2000
21 May
2000
21 May
2000
21 May
2000
21 May
2000
18 May
2000
Depth
PspA
m
983
2
987
985
1401
14
2267
2737
1530
1875
2
2060
2060
213
212
950
4
950
950
6
1320
1340
1
1335
2
1570
3
1580
1620
2
1620
7
1620
1
3015
1
Station
Postition
Date
RW6-5
25°59.8382 'N
94 29.73478'W
29° 20.0500'N
87°03.3758'W
29°19.9897'N
87° 02.9021 "W
29°20.1091'N
87° 02.7818"W
28°55.1647'N
87°40.2232'W
28°55.0080'N
87°40.0627'W
28°55.1634'N
87°40.2154'W
28°33.4054'N
87°45.7357'W
28°33.4292'N
87° 45.644 l'W
28° 33.2345 'N
87° 45.8373 "W
28° 16.3144 'N
87° 19.5165'W
27° 29.9500'N
87° 00.0849'W
28° 00.8463'N
86°34.5587'W
28° 01.4243 'N
86°34.2753'W
28°15.1557'N
86° 25.0663'W
28°15.1070'N
86°25.0209'W
28° 15.2344'N
86° 24.7995'W
28°45.0632'N
85°44.8807'W
27°34.7791'N
93°32.8573'W
27°34.7791'N
93°32.8573'W
27°24.8008'N
93°20.2579'W
27°24.7019'N
93°20.3849'W
27°24.7019'N
93°20.3849'W
27° 24.8517 'N
93°20.4010'W
27° 24.8517'N
18 May
2000
11 June
2000
11 June
2000
6 June
2000
12 June
2000
12 June
2000
12 June
2000
13 June
2000
13 June
2000
13 June
2000
14 June
2000
6 June
2000
8 June
2000
8 June
2000
9 June
2000
7 June
2001
10 June
2000
11 June
2000
13 May
2000
5 Aug
2001
14 May
2000
14 May
2000
17 Feb
2000
15 Aug
2001
17 Feb
S35-1
S3 5-2
S35-5
S36-1
S36-2
S36-3
S37-1
S37-2
S37-3
S3 8-2
S39-1
S41-1
S41-2
S42-1
S42-2
S43-2
S44-3
Wl-1
Wl-1
W2-1
W2-2
W2-2
W2-3
W2-3
Depth
in
3010
658
667
666
1825
1832
1826
2388
2382
2387
2642
3007
2974
2974
767
773
360
212
379
389
625
625
625
625
625
189
Station
W3-1
W3-2
W4-1
W4-2
W4-5
W5-2
W6-2
WC121
WC122
WC123
WC5-1
WC5-2
WC5-5
Postilion
Date
93° 20.4010'W
27° 10.371 l'N
93° 19.3081 'W
27°10.3840'N
93°19.3315'W
26° 43.9027 'N
93°19.1708'W
26°43.8500'N
93° 19.1843 'W
26° 43.8494'N
93°19.1836'W
26° 16.5781'N
93° 21.7309'W
25° 59.7526 'N
93° 18.8003'W
27°19.3945'N
91°33.3486'W
27°19.11205'N
91°33.2314'W
27°19.4320'N
91°33.1625'W
27°46.2838'N
91°45.7293'W
27°47.1150'N
91°45.8502'W
27° 46.9893'N
91°45.8800'W
2000
14 May
2000
17 Aug
2001
15 May
2000
18 Aug
2001
15 May
2000
18 June
2001
17 May
2000
5 May
2000
9 Aug
2001
5 May
2000
4 May
2000
23 May
2001
4 May
2000
Depth
01
PspA
PspC
PspO
PspjP
MspA
PPsp
A
860
860
1420
1460
1460
2740
3145
1168
1125
1175
400
320
325
* PspA - Pseudotanais sp. A, PspC = Pseudotanais sp. C, PspO = Pseudotanais sp. Q, PspP :
Pseudotanais sp. P, MspA = Mystriocentrus sp. A. PPspA = Parapseudotanais sp. A
190
APPENDIX B
Description and diagnostics of most parsimonious tree from parsimony analysis
Tree rooted using four out-groups
Tree length = 630
Consistency index (CI) = 0.2540
Homoplasy index (HI) = 0.7460
CI excluding uninformative characters = 0.2432
HI excluding uninformative characters = 0.7568
Retention index (RI) = 0.5179
Rescaled consistency index (RC) = 0.1315
Character summary: 69 characters, all with equal weight and with unordered character states,
nine were parsimony-uninformative; gaps were treated as "missing"; character states were
optimized with Accelerated transformation (ACCTRAN)
Character diagnostics:
Character
1 Cephalothorax length vs. pereonites 1 -3
2 Eyelobes present or absent
3 Pereonite 1 abreviation +/4 Antennule biramous or uniramus
5 Antennule number of articles
6 Antennule article 1 % of ramus
7 Antennule article 1 length/width
8 Antennule article 3 length/width
9 Antennule distal article length/width
10 Antenna number of articles
11 Antenna length vs. antennule length
12 Antenna article 2 length vs article 3
13 Antenna articles 2 & 3 setation
14 Antenna article 4 length/width
15 Antenna article 5 length/width
16 Mandible pars molaris palp +/17 Mandible pars molaris width
18 Mandible pars molaris complexity
19 Maxillule number of endites
20 Maxillule palp number of terminal setae
21 Maxillule endite number of spiniform setae
22 Maxilla complexity
23 Maxilliped bases degree of fusion
24
25
26
27
28
29
30
31
Maxilliped endites degree of fusion
Maxilliped endite setation
Cheliped chela degree of forcipation
Cheliped carpus length/width
Cheliped carpus number of inferior setae
Cheliped propodus length/width
Cheliped propodus length vs. carpus
Cheliped fixed finger inferior setae
steps;
12
10
1
1
3
15
18
3
14
1
8
13
5
17
11
1
4
23
1
1
9
1
6
13
19
9
11
6
11
3
7
CI
0.167
0.200
1.000
1.000
0.667
0.200
0.167
0.667
0.286
1.000
0.250
0.154
0.200
0.235
0.182
1.000
0.250
0.391
1.000
1.000
0.556
1.000
0.500
0.231
0.263
0.222
0.182
0.500
0.182
0.333
0.429
RI
0.412
0.500
1.000
0/0
0.889
0.429
0.464
0.750
0.545
0/0
0.250
0.389
0.733
0.594
0.500
0/0
0.786
0.517
0/0
0/0
0.000
0/0
0.400
0.730
0.300
0.417
0.438
0.000
0.438
0.895
0.600
RC
0.069
0.100
1.000
0/0
0.593
0.086
0.077
0.500
0.156
0/0
0.062
0.060
0.147
0.140
0.091
0/0
0.196
0.202
0/0
0/0
0.000
0/0
0.200
0.168
0.079
0.093
0.080
0.000
0.080
0.298
0.257
HI
0.833
0.800
0.000
0.000
0.333
0.800
0.833
0.333
0.714
0.000
0.750
0.846
0.800
0.765
0.818
0.000
0.750
0.609
0.000
0.000
0.444
0.000
0.500
0.769
0.737
0.778
0.818
0.500
0.818
0.667
0.571
191
Character
32 Cheliped dactylus % length of propodus
33 Cheliped dactylus width vs. fixed finger
34 Cheliped dactylus proximal seta
35 Marsupium development
36 Pereopods ischium present or absent
37 Pereopod 1 basis length/width
38 Pereopod 1 basis setation
39 Pereopod 1 ischium setation
40 Pereopod 1 merus setation
41 Pereopod 1 carpus setation
42 Pereopod 1 propodus length/width
43 Pereopod 1 propodus setation
44 Pereopod I dactylus setae +/45 Pereopod 1 dactylus setae type
46 Pereopod 1 dactylus +unguis length
47 Pereopods 2 & 3 basis setation
48 Pereopods 2 & 3 ischium setation
49 Pereopods 2 & 3 merus setation
50 Pereopods 2 & 3 merus modified setae
51 Pereopods 2 & 3 carpus blades +/52 Pereopods 2 & 3 carpus blade length
53 Pereopods 2 & 3 dactylus + unguis length
54 Pereopod 3 size vs. pereopod 2
55 Pereopod 4 basis length/width
56 Pereopods 4-6 basis setation
57 Pereopods 4-6 ischium setation
58 Pereopods 4-6 merus setation
59 Pereopods 4-6 carpus modified setae +/60 Pereopods 4-6 carpus other mod. setae +/61 Pereopods 4-6 propodus broom setae +/62 Pereopods 4-6 dactylus fusion
63 Pleopods degree of development
64 Uropods uniramous or biramous
65 Uropod exopod number of articles
66 Uropod exopod length vs endopod length
67 Uropod exopod proximal article length
68 Uropod endopod number of articles
69 Uropod endopod proximal article length
steps,
10
18
10
2
1
14
18
11
17
16
9
18
11
3
16
25
6
15
1
1
6
22
1
16
19
16
9
2
3
9
2
6
1
12
9
6
11
1
CI
0.200
0.111
0.100
1.000
1.000
0.214
0.222
0.182
0.235
0.375
0.222
0.278
0.091
0.667
0.125
0.120
0.500
0.200
1.000
1.000
0.167
0.136
1.000
0.125
0.158
0.125
0.222
0.500
0.667
0.111
0.500
0.667
1.000
0.250
0.222
0.333
0.273
1.000
RI
0.333
0.360
0.250
1.000
0/0
0.577
0.481
0.182
0.235
0.677
0.682
0.480
0.474
0.500
0.417
0.312
0.250
0.333
1.000
1.000
0.643
0.406
0/0
0.462
0.448
0.481
0.417
0.957
0.750
0.500
0.889
0.778
0/0
0.182
0.417
0.000
0.273
1.000
RC
0.067
0.040
0.025
1.000
0/0
0.124
0.107
0.033
0.055
0.254
0.152
0.133
0.043
0.333
0.052
0.037
0.125
0.067
1.000
1.000
0.107
0.055
0/0
0.058
0.071
0.060
0.093
0.478
0.500
0.056
0.444
0.519
0/0
0.045
0.093
0.000
0.074
1.000
HI
0.800
0.889
0.900
0.000
0.000
0.786
0.778
0.818
0.765
0.625
0.778
0.722
0.909
0.333
0.875
0.880
0.500
0.800
0.000
0.000
0.833
0.864
0.000
0.875
0.842
0.875
0.778
0.500
0.333
0.889
0.500
0.333
0.000
0.750
0.778
0.667
0.727
0.000
APPENDIX C
Character Matrix for Phylogenelic Analysis of Pseudotanaidae
Note: A = character state 10; missing data indicated by "-" or "?"
1
Taxon
Carpoapseudes prospectnes
2
2
Sinelobus stanfordi
1
Hargeria rapax
1
Typhlotanais parvus
3
Cryptocope abreviata
3
Cryptocopoides arcticus
Cryptocopoides pacificus
2
3
Cryptocopoides sp. A
1
Curtichelia expressa
Iungentitanais primitivus
3
Paraiungentitanais
longidigitatus 3
Latitanais beklemishevi
1
Parapseudotanais abyssalis
3
Parapseudotanais sp. A
3
Mystriocentrus serratus
2
Mystriocentrus sp. A
_3j
Pseudotanais (A) gaussi
3
Pseudotanais (A) gerlachi
3
Pseudotanais (A) guillei
3
Pseudotanais (A) longipes
3
Pseudotanais (A) rnakrothrix
3
Pseudotanais (A) malayensis
2
Pseudotanais (A) mortenseni
3
Pseudotanais (A) scrappi
3
Pseudotanais (A) siegi
3
2
3
1
1
3
3
3
3
3
3
1
2
2
3
3
3
3
3
1
1
3
1
1
1
1
1
3
2
2
2
2
2
2
2
2
2
2
2
2
2
4
1
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
5 6
1 1
2 3
2 3
3 4
2 3
2 2
2 2
2 2
2 1
2 2
2 1
2 1
3 2
3 2
3 3
3 3
3 2
3 4
3 3
3 3
3 3
3 3
3 3
3 3
3 3
7
8
3
2
3
4
2
1
1 9
1 1
1 1
1 1
1 1
1 2
2 2
1 2
1 2
2 ?
2 9
3 9
2 9
3 9
2 9
2 9
4 9
3 9
2 9
2 9
2 9
3 9
10
9
1
1 2
1 2
2 2
2 2
2 2
2 2
2 2
3 2
3 2
3 2
4 2
5 2
4 2
2 2
2 2
3 2
3 2
3 2
4 2
3 ?
3 2
3 2
3 2
5 2
9
11
1
2
2
1
1
2
2
2
2
2
2
2
1
2
2
3
1
2
2
2
?
2
2
2
1
12
?
3
2
3
2
2
3
3
3
2
2
2
3
1
2
2
2
2
1
3
?
2
2
1
2
C laracters
13 14 15
9
9
2
2
1
2
1
1
2
2
1
2
2
2
1
2
1
2
2
3
3
2
2
3
2
2
2
2
2
3
2
3
3
2
3
4
2
1
2
1
3
4
2
1
1
3
2
2
2
1
2
2
5
9
9
2
2
1
2
2
2
3
2
2
1
2
3
5
16
1
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
17
2
2
2
2
2
2
2
2
2
2
2
2
2
2
18
1
2
2
3
8
4
4
4
4
4
4
4
8
9
A
8
A
A
A
7
A
A
A
A
A
19
2
20
2
?
9
9
?
9
9
9
9
21
1
1
1
3
3
4
4
3
2
6
3
3
3
3
3
3
3
3
3
3
?
3
3
5
3
22
2
23
1
3
3
3
4
2
3
3
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
24
25
1
o
j
2
2
4
4
2
3
4
2
4
4
4
4
5
5
5
5
5
6
3
3
6
4
5
6
4
5
6
1
5
5
5
6
5
5
1
C laracters
Taxon
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
1
(A) similis
(P) abyssi
(P) affinis
(P) baresnauti
(P) borceai
(P) califomiensis
(P) colonus
(P) corollatus
(P) denticulatus
(P)falciculata
(P) inflatus
(P) forcipatus
(P)jonesi
(P) liljeborgi
(P) longisetosus
(P) longispinus
(P) nipponicus
(P) macrocheles
(P) mediterraneus
(P) mexikolpos
(P) nordenskioldi
(P) oculatus
(P) scalpellwn
(P) spatula
(P) spicatus
(P) unicus
(P) vitjazi
(P) vulsella
(P) sp. A
(P) sp. C
3
3
2
3
3
3
3
2
3
1
3
3
3
3
3
2
1
3
3
3
3
2
2
1
3
3
3
3
3
3
2
3
3
3
3
1
3
3
3
3
3
3
3
3
2
3
3
3
3
1
1
3
1
3
3
3
1
3
3
3
3
3
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
4
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
5
3
3
3
3
3
3
3
3
3
3
3
3
3
3
3
3
3
3
3
3
3
3
3
3
3
3
3
3
3
6
7
8
9
10
11
12
13
14
15
16
17
18
4
4
3
4
3
4
3
2
4
4
4
4
2
3
2
2
2
3
2
2
3
3
4
3
2
3
2
2
3
2
2
2
2
3
3
3
3
2
3
3
3
4
?
?
?
?
?
3
2
2
2
2
2
?
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
1
2
2
2
2
?
2
2
2
3
3
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
?
2
1
2
2
1
2
2
1
1
2
2
1
2
1
2
1
2
2
2
2
2
2
1
1
2
2
1
1
2
?
1
2
3
4
2
2
?
2
2
4
3
5
2
2
2
2
5
4
3
3
2
2
2
5
4
3
2
4
3
3
4
1
3
2
1
1
?
1
1
2
1
2
1
1
1
1
2
2
1
2
1
1
1
2
2
2
1
2
1
1
1
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
1
1
2
1
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
1
2
A
A
7
8
A
A
A
4
4
9
4
A
A
7
5
5
5
8
A
A
5
7
7
6
6
A
6
9
4
5
^
j
3
3
3
3
3
3
3
3
3
4
->
j
3
3
4
3
?
?
?
?
?
?
?
?
?
?
?
?
?
?
?
?
?
?
?
?
->
j
3
2
2
2
2
3
3
2
5
2
2
2
3
3
3
3
3
3
3
3
3
3
3
9
?
?
?
?
3
3
2
3
1
2
2
2
2
2
2
1
!
1
2
2
1
2
1
1
1
2
1
2
1
1
0
-)
2
2
2
2
2
2
2
2
2
2
2
2
2
2
19
20
?
?
?
?
?
?
?
?
?
?
?
?
21
3
3
3
3
?
?
3
3
3
3
5
3
3
3
3
3
3
3
3
3
3
3
3
3
3
3
3
3
3
3
22
25
23
24
4
4
4
4
3
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
5
3
5
3
2
5
5
3
6
3
4
5
3
6
3
6
2
5
4
3
1 3
4
6
4
6
3
5
3
5
2
3
2
3
2
5
3
6
4
5
5
3
3
5
5
2
2
5
2 l~5
5
3
2
5
3 U,
2
3
2
5
Taxon
Pseudotanais (P) sp. 0
Pseudotanais (P) sp. P
Taxon
Carpoapsendes prospectnes
Sinelobus stanfordi
Hargeria rapax
Typhlotanais parvus
Cryptocope abreviata
Cryptocopoides arcticus
Cryptocopoides pacificus
Cryptocopoides sp. A
Curtichelia expressa
hmgentitanais primitivus
Paraiungentitanais
longidigitatus
Latitanais beklemishevi
Parapseudotanais abyssalis
Parapseudotanais sp. A
Mvstriocentrus serratus
Mystriocentrus sp. A
Pseudotanais (A) gaussi
Pseudotanais (A) gerlachi
Pseudotanais (A) guillei
Pseudotanais (A) longipes
Pseudotanais (A) makrothrix
Pseudotanais (A) malayensis
Pseudotanais (A) mortenseni
Pseudotanais (A) scrappi
Pseudotanais (A) siegi
1 2 1 3 4
~2^ 3 2 2
2 3 2 2
26
3
3
3
3
3
3
3
3
3
3
3
3
3
3
1
1
3
2
2
3
3
^
j
3
3
3
27
2
2
28
1
3
2
3
3
3
3
o
j
2
2
2
2
2
2
1
1
1
2
1
1
2
1
2
3
4
3
3
3
3
3
3
3
3
3
o
j
3
3
3
3
3
5
3
3
29
1
1
1
2
1
I
1
2
1
3
3
1
2
2
2
1
1
2
2
3
2
2
2
2
7
2
2
6
3
3
30
2
2
2
2
8
?
?
9
4
3
31
1
3
2
2
3
2
2
2
2
3
2
2
2
2
3
3
3
32
3
2
2
2
2
2
2
1
3
1
1
-»
j
2
3
3
3
3
3
3
3
2
2
3
3
2
2
T
2
2
1
2
I
2
10
2
2
33
1
2
1
2
1
2
2
2
3
1
3
2
2
2
2
2
1
1
2
2
1
2
1
2
1
11
2
2
34
2
2
1
2
2
2
2
2
2
2
2
12
1
2
35
1
3
1
1
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
Characters
13 14 15
1
1
5
4
1
1
Characters
36 37 38
4
1
2
2
2
1
2
1
1 4
4
2
4
2
4
3
4
3
4
3
5
3
3
5
3
5
3
5
4
2
3
5
5
2
3
4
2
4
2
4
5
7
?
2
5
5
3
5
2
5
3
17
2
2
16
2
2
39
2
2
2
2
2
2
2
2
2
2
3
2
3
3
2
2
1
2
2
?
2
2
o
j
2
18
7
6
19
1
1
20
1
1
21 22
3
1
3 |1
9
42
2
2
2
2
2
2
2
2
2
3
3
3
2
2
2
2
4
2
2
4
7
43
1
2
2
4
4
3
3
4
4
3
3
5
4
5
4
4
6
5
4
5
7
6
5
7
6
3
2
3
4
6
5
6
5
40
1
4
3
4
4
4
4
3
3
5
5
3
4
4
4
4
4
5
4
4
41
1
5
2
5
5
4
3
3
5
5
5
5
4
4
6
6
7
6
5
6
9
5
4
5
4
44
1
2
2
1
2
2
2
2
1
1
1
2
1
1
2
1
1
2
2
1
?
2
2
2
2
24
2
2
25
5
5
46
3
3
2
2
2
47
3
1
1
3
4
2
3
1
23
4
4
45
1
3
3
3
2
2
3
3
3
7
-
o
j
2
3
2
3
3
2
2
2
3
3
3
3
2
3
7
3
3
2
n
j
9
4
4
7
4
3
4
1
2
4
1
4
2
4
4
3
4
Taxon
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
(A) similis
(P) abyssi
(P) affinis
(P) baresnauti
(P) borceai
(P) californiensis
(P) colonus
(P) corollatus
(P) denticulatus
(P)falciculata
(P) inflatus
(P) forcipatus
(P) jonesi
(P) liljeborgi
(P) longisetosus
(P) longispinus
(P) nipponicus
(P) macrocheles
(P) mediterraneus
(P) mexikolpos
(P) nordenskioldi
(P) oculatus
(P) scalpellum
(P) spatula
(P) spicatus
(P) unicus
(P) vitjazi
(P) vulsella
(P) sp. A
(P) sp. C
26
3
1
3
3
3
1
3
3
3
1
2
1
27
1
2
2
]
3
3
3
3
3
3
2
3
2
3
3
3
3
2
1
3
3
2
2
2
28
3
2
3
3
3
3
3
2
3
3
3
3
3
3
3
3
3
2
3
3
3
3
3
3
3
3
3
3
3
3
29
2
2
1
2
1
1
1
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
30
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
31
3
4
3
3
3
3
3
3
3
3
7
3
4
3
3
3
3
3
3
3
3
3
3
3
3
3
3
3
3
3
32
2
2
2
2
2
2
2
2
2
2
2
2
2
1
1
2
2
2
2
2
2
2
2
2
2
9
2
2
2
33
1
2
2
2
1
2
2
2
1
2
2
2
3
2
2
1
1
1
2
2
1
2
1
1
2
1
2
1
1
1
34
2
2
2
35
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
"7
2
2
Characters
36 37 38
5
2
2
3
4
4
4
3
?
5
5
2
4
2
2
5
4
2
3
3
5
4
4
2
4
2
4
2
4
2
1
2
2
1
4
2
4
2
5
2
4
2
3
2
1
3
1
3
4
4
5
2
1
3
5
4
4
4
4
3
39
2
2
2
2
3
2
2
2
3
2
2
2
2
2
2
2
2
2
1
2
2
2
3
3
3
2
3
2
3
2
40
4
4
4
4
5
4
5
4
4
4
5
4
3
4
4
4
4
4
3
4
2
4
4
4
5
5
5
4
4
4
41
6
6
5
5
7
7
5
6
3
6
6
7
7
5
6
5
5
5
6
6
6
6
5
4
3
7
5
6
6
5
42
3
2
4
2
2
2
2
3
2
2
2
2
2
2
3
4
3
^
j
2
2
2
3
4
4
3
2
3
3
2
4
43
6
5
6
5
6
5
4
5
5
4
6
5
5
5
6
4
5
6
5
5
5
5
5
5
5
5
6
4
4
5
44
2
1
1
]
2
2
2
2
2
1
2
2
2
2
2
1
2
2
2
2
1
2
1
1
2.
2
2
1
2
2
45
3
3
3
3
3
3
3
3
3
-
46
3
2
3
2
3
3
3
2
3
47
2
1
2
4
9
7
2
4
3
4
9
3
3
1
2
2
2
3
2
2
2
2
2
2
1
2
2
2
2
3
1
2
3
1
1
1
3
4
2
1
2
1
3
4
4
4
2
2
1
32
2
3
33
1
1
34
1
1
35
2
2
31
3
3
2
Characters
36 37 38
4
1
1
1
3
52
-'
1
1
1
1
-
53
2
3
4
3
2
2
2
3
?
4
4
2
4
4
5
4
2
3
4
3
3
4
3
4
54
55
2
1
1
1
3
56
1
1
1
2
4
4
3
57
3
2
3
2
3
1
2
2
3
3
2
3
3
3
3
3
2
2
1
?
2
1
2
Characters
58 59 60
9
1
9
2
9
2
9
3
9
2
9
2
2
2
2
2
?
2
9
2
9
2
?
2
9
2
9
2
2
2
3
2
2
3
?
3
?
2
?
2
?
2
?
?
?
2
?
2
?
2
Taxon
Pseudotanais (P) sp. O
Pseudotanais (P) sp. P
26
3
3
27
1
2
28
3
3
29
2
2
30
2
Taxon
Carpoapsendes prospectnes
Sinelobus stanfordi
Hargeria rapax
Typhlotcmais parvus
Cryptocope abreviata
Cryptocopoides arcticus
Cryptocopoides pacificus
Cryptocopoides sp. A
Curtichelia expressa
lungentitanais primitivus
Paraiungentitanais
longidigitatus
Latitanais beklemishevi
Parapseudotanais abyssal is
Parapseudotanais sp. A
Mystriocentrus serratus
Mystriocentrus sp. A
Pseudotanais (A) gaussi
Pseudotanais (A) gerlachi
Pseudotanais (A) guillei
Pseudotanais (A) longipes
Pseudotanais (A) makrothrix
Pseudotanais (A) malayensis
Pseudotanais (A) mortenseni
Pseudotanais (A) scrappi
48
1
3
4
4
4
4
4
?
4
4
49
1
2
3
3
3
4
4
4
?
4
4
50
2
2
2
2
2
2
2
2
2
2
2
2
2
2
1
1
2
2
2
2
2
2
2
2
51
9
9
4
5
4
4
4
3
4
4
4
3
4
3
4
3
3
2
4
3
3
3
3
3
3
3
2
2
2
2
2
2
1
2
2
1
1
2
3
3
2
1
1
3
?
1
3
1
2
4
4
4
4
4
2
2
1
4
2
4
9
4
2
4
39
2
61
1
2
2
2
2
2
1
1
2
2
2
2
2
2
1
1
2
2
2
2
?
2
2
2
40
3
5
41
4
4
42
4
4
43
5
5
44
2
1
45
2
46
2
2
47
1
1
62
2
1
1
2
2
2
2
2
2
2
2
2
63
1
1
1
1
4
3
2
3
1
1
1
1
2
3
3
3
3
3
3
3
3
64
1
2
65
67
1
3
2
2
2
2
2
?
2
2
?
2
2
68
1
1
1
4
2
2
2
2
2
2
2
2
2
4
3
2
2
2
2
2
3
2
2
69
?
0
66
1
1
2
2
3
3
3
3
2
2
3
3
3
2
2
3
3
3
3
3
2
3
?
3
3
1
4
4
4
2
2
2
2
2
2
2
2
2
3
2
2
2
2
4
3
3
3
2
2
?
2
2
3
4
1
1
?
2
2
2
2
2
2
2
?
2
?
?
2
2
2
n
2
?
2
2
9
l
Taxon
Pseudotanais
Pseudotcmais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
(A) siegi
(A) similis
(P) abyssi
(P) affinis
(P) baresnauti
(P) borceai
(P) californiensis
(P) colonus
(P) corollatus
(P) denticulatus
(P)falciculata
(P) inflatus
(P) forcipatus
(P)jonesi
(P) liljeborgi
(P) longisetosus
(P) longispinus
(P) nipponicus
(P) macrocheles
(P) mediterraneus
(P) mexikolpos
(P) nordenskioldi
(P) oculatus
(P) scalpellum
(P) spatula
(P) spicatus
(P) unicus
(P) vitjazi
(P) vulsella
(P) sp. A
48
4
4
4
4
4
?
4
4
4
4
4
?
4
4
4
4
4
4
4
3
4
4
4
4
4
4
4
?
4
4
49
4
4
3
3
3
?
4
4
3
3
3
?
3
4
3
3
3
3
4
3
3
4
3
3
3
4
3
3
4
50
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
51
1
1
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
0
2
52
-
2
2
2
2
2
2
1
2
2
1
2
2
1
2
1
2
1
53
3
4
54
4
3
4
4
3
3
3
4
5
3
2
3
2
3
4
4
2
4
3
4
3
4
2
5
4
5
2
55
3
1
3
2
2
?
7
2
2
1
2
2
2
2
2
2
2
2
1
1
1
2
->
2
1
1
2
2
1
56
4
2
2
2
2
?
?
0
z-
4
2
2
7
2
2
4
1
1
1
3
1
1
2
1
2
1
2
2
4
4
2
Characters
57 58 59 60
?
2
1
3
?
1
2
3
?
2
3
3
?
2
3
2
7
2
3
3
7
?
7
2
?
7
?
2
?
2
2
3
7
2
2
2
?
2
2
2
?
2
2
1
?
?
?
2
?
2
2
3
7
2
3
3
7
2
2
3
2
3
3
2
3
3
2
2
2
3
2
1
i
?
2
2
?
2
2
2
7
2
2
2
7
2
2
3
7
2
2
2
7
2
3
3
2
3
3
3
?
2
2
2
?
2
2
2
7
?
2
3
7
2
2
2
?
2
2
L*
61
2
1
2
2
2
2
7
2
2
1
1
?
2
2
2
2
1
1
2
2
2
2
2
1
1
1
2
2
2
1
62
?
?
?
63
3
3
3
3
3
3
3
3
3
3
3
3
5
3
5
3
3
3
3
3
3
3
3
3
3
3
3
3
3
64
65
2
4
2
2
2
4
3
4
2
2
2
i
3
2
2
4
2
2
2
2
2
2
2
2
2
2
2
2
2
2
66
3
3
3
3
3
3
2
2
3
3
3
3
67
2
2
2
2
?
2
2
2
3
9
2
2
3
3
3
2
3
3
3
3
3
3
3
3
3
3
3
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
68
2
4
2
2
2
4
3
2
2
2
2
2
3
2
2
2
2
2
2
2
2
2
2
2
2
4
2
69
2
7
2
2
2
2
?
2
2
2
2
2
7
2
2
2
2
2
2
2
J
2
2
2
2
?
2
T
2
2
2
2
Taxon
Pseudotanais (P) sp. C
Pseudotanais (P) sp. O
Pseudotanais (P) sp. P
48
4
4
4
49
4
3
4
50
2
2
2
51
2
2
2
52
2
2
2
53
4
3
3
54
1
1
1
55
2
2
3
56
1
1
1
57
1
1
1
Characters
58 59 60
1
2
2
3
2
3
2
1
2
61
1
1
1
62
1
1
I
63
3
3
3
64
1
1
1
65
2
2
4
66
3
3
3
67
2
2
-
68
2
2
3
69
2
2
?
GO
199
APPENDIX D
Synopsis of Characters
The following is an annotated list of the 69 characters used to perform the heuristic
parsimony analyses detailed in chapter III; associated character states are in parentheses. An
abbreviated list with diagnostics for each character is presented in Appendix B and a complete
matrix of species with character states is presented in Appendix C. As mentioned earlier, scores
for character states were taken from both, original descriptions of new species discovered during
this research, and from published illustrations and text descriptions in the literature. The
characters are listed in order from the anterior to posterior ends of the body and characters states
are, in general, in the suspected order of primitive to derived. Figures depicting the tracing of all
characters are given except for characters 4, 10, 16, 19, 20, 22, 36, 54 and 64 which were
parsimony-uninformative. Figures in this appendix were produced using McClade from the
consensus of six most parsimonious trees (see Figure 1 in chapter III). A summary of results is
listed in Chapter IV.
1. Cephalothorax length vs. length of pereonites 1-3 (1 = shorter, 2 = subequal, 3 =
longer). A shorter cephalothorax of less than or subequal to the length of the first three pereonites
appears to be a plesiomorphic feature based on the out-group and one of the Cryptocopinae
clades. A longer cephalothorax is seen in all but twelve of the Pseudotanainae species, including
only one of the subgenus Akanthinotanais, and in six of the Cryptocopinae (Fig. 33).
2. Eyelobes present or absent (1 = present with pigment, 2 = rudimentary with no
pigment, 3 = absent). The species in this study comprise a mixture of eyed and eyeless
representatives including those of the out-group, thus this character has doubtful use in making
inferences on phylogeny, but is likely more an indication of bathymetric distribution (Larsen and
Wilson 2002, Guerrero- Kommritz and Brandt 2005). It is notable that seven of the ten
Akanthinotanais species and four of the Pseudotanainae possess eyes, reflective of their
comparatively shallow distribution (Fig. 34).
200
3. Pereonite abbreviation (1 = no pereonites abbreviated, 2 = at least the first pereonite
abbreviated). This character is one of the most defining features of the Pseudotanaidae and one
that sets apart the Pseudotanainae from the Cryptocopinae. However, Larsen and Wilson (2002)
noted with caution that this feature can also be present in some other tanaid families as an
ontogenetic variable. All out-group and Cryptocopinae species lack abbreviated pereonites (Fig.
35) while the species of the Pseudotanainae have the first and sometimes the second pereonites
abbreviated.
4. Antennule rami number (1 = biramous, 2 = uniramous). This character was included
to resolve out-group relationships - the antennule is biramous in apseudomorphs but uni-ramous
in tanaidomorphs (Sieg 1983).
5. Antennule number of articles (1 = more than four articles, 2 = four articles, 3 = three
articles). Another diagnostic feature of the Pseudotanainae which all have three antennular
articles, this character is also useful in resolving relationships among the out-group species (Fig.
36).
6. Antennule first article percent of total length of antennule (1 = up to 42 %, 2 = 42.1 to
49 %, 3 = 50 to 59.3%, 4 = 59.4 % or greater). The length of the first antennular article tended to
be shorter in the apseudomorph out-group and in seven of the Cryptocopinae (length of 49% or
less). An apomorphic tendency toward a longer first antennule article in comparison to other
antennular articles was apparent in the Pseudotanainae with the exception of four species,
including both Parapseudotanais species (Fig 9).
7. Antennule first article length vs. width (1 = up to 2.9 times longer than broad, 2 = 3.5
to 5.0 times longer than broad, 3 = 5.1 to 6.9 times longer than broad, 4 = 7.0 or greater times
longer than broad). The same apormorphic tendency toward a lengthening first antennular article
seen in character 6 was also manifested in this character. The first article width tended to be short
and broad in the out-group species (except for the apseudomorph), and in most of the
Cryptocopinae species, but was longer in all of the Pseudotanainae species (Fig.38).
201
8. Antennule third article length vs. width (1 = up to 1.0 times longer than broad, 2 = 1.5
to 2.5 times longer than broad, 3 = 4.0 or greater times longer than broad). This character was
scored only for those species among the out-group and Cryptocopinae having four antennular
articles. It was important in defining a strong clade within the Cryptocopinae (clade C) composed
of four monospecific genera, Iungentitcmais, Paraiungentitanais, Latitanais and Curtichelia, all
with character state 2 (Fig. 39).
9. Antennular distal article length vs. width (1 = up to 1.9 times longer than broad, 2 =
2.0 to 3.9 times longer than broad, 3 = 4.0 to 6.5 times longer than broad, 4 = 6.6 to 8.0 times
longer than broad, 5 = 8.1 or greater times longer than broad). This character showed no clear
patterns other than showing most in-group species with a longer distal antennular article than
those of the out-groups (Fig. 40). Apparently a longer distal antennular article arose
independently among several pseudotanaid species.
10. Antenna number of articles (1 = more than six articles, 2 = six articles). This outgroup resolving character sets apart the apseudomorphs from all other species in the study that
have only six antenna articles.
11. Antenna length vs. length of antennule (1 = up to 80 % length of antennule, 2 = 80.1
to 108.0 % length of antennule, 3 = 108.1 % or greater length of antennule). An antenna that is
shorter than 80% of the antennule is possibly an ancestral trait since it was a feature of the
apseudomorph out-group but was most likely independently evolved since it appears in five
scattered in-group species. Likewise, an abnormally long antenna, as compared to the antennule
length, was only found in three Pseudotanainae species (Fig. 41).
12. Antenna second article length compared to that of third article (1 = shorter than third
article, 2 = subequal to third article, 3 = longer than third article). A long third article inthe
antenna was another possible ancestral trait since it appeared in three of the out-group species.
This character was more variable in the in-group species and showed a slight tendency to become
subequal or shorter in more derived species (Fig. 42).
202
13. Antenna second and third articles type of seta (1 = short, stout spiniform seta. 2 =
seta otherwise - simple or spiniform). Sixteen species of Pseudotanainae have the stout,
spiniform version of these antennular setae, twelve of which appeared in clade A, furthest away
from the ancestral end of the tree (Fig.43). This seemingly contradicts the notion that spiniform
setae are plesiomorphic and are replaced by simple setae. The state of this character was found to
be diagnostic by Bird and Holdich (1989) in distinguishing Pseudotanais corollatus from the redescribed P. affinis.
14. Antenna fourth article length vs. width (1 = up to 4.0 times longer than broad, 2 = 4.1
to 5.9 times longer than broad, 3 = 6.0 to 8.0 times longer than broad, 4 = 8.1 to 10.2 times longer
than broad, 5 = 10.3 or more times longer than broad). A shorter antennal fourth article (state 1)
was characteristic of the out-groups and half of the Cryptocopinae species and is, thus considered
to be ancestral. Varying lengths (states 2-4) are found in one group of Cryptocopinae (clade C)
and in all Pseudotanainae but no particular pattern was evident (Fig. 44).
15. Antenna fifth article length vs. width (1 = up to 3.9 times longer than broad, 2 = 4.0
to 5.9 times longer than broad, 3 = 6.0 or more times longer than broad). An elongated fifth
article of greater than 4.0 times longer than broad (state 2) was found in only 15 Pseudotanainae
species and of these, only three had lengths greater than 6.0 times longer than broad (state 3).
The extreme length of the fifth article (state 3) is a defining feature of the genus
Parapseudotanais (Fig. 45).
16. Pars molaris palp present or absent on mandibles (1 = absent, 2 = present). This
character was included to resolve out-group relationships; a palp is present on the mandible pars
molaris of apseudomorphs (Sieg 1983) but absent in tanaidomorphs.
17. Width of the mandible pars molaris (1 = broad and truncated, 2 = narrow with
acuminate tip). The broad, truncate aspect of this character is apparently an ancestral state given
that it is predominantly found in the out-group species and all but one of the Cryptocopinae.
203
Except for four Pseudotanainae species, the pars inolaris is acuminate or tapered to a point (Fig.
46).
18. Mandible pars molaris complexity (1 = unarmored, 2 = with pad-like grinding
surface, 3 = with non-setose crest, 4 = with ring of setae and/or short denticles, 5 = with spinelike process and five or more denticles, 6 = with five or more denticles but no long processes, 7 =
with three to four blunt terminal/subterminal denticles, 8 = with bifid tip, 9 = with some marginal
setules, 10 = simple pointed). The wide variety of pars molaris tip secondary characters, together
with the nature of its width as seen in the previous character, is reflective of independently
evolved feeding strategies among widely dispersed species of tanaids. Although highly plastic
when seen in the context of 57 species, the decrease in complexity of the pars molaris structure
was viewed as a transition from primitive to a more derived state (Sieg 1977). This trend was
seen to some degree in the phyologeny presented in this research (Fig. 47). In general, the broad,
truncated pars molaris found in the out-group and Cryptocopinae species had the more complex
tips, whereas in most of the Pseudotanainae species with a narrower pars molaris, the tips were
reduced in complexity. Four species of Pseudotanainae (Pseudotanais denticulatus, P.
corollatus, P. inflatus, and P. sp. A) retained the ancestral broad pars molaris with a terminal ring
of spines and setae. In contrast, in all but one species of the subgenus Akanthinotanais, the pars
molaris was reduced to a simple or whip-like point.
19. Number of maxillule endites (1 = two endites, 2 = one endite). This character was
included to resolve out-group relationships; the maxillule is biramous in apseudomorphs and
uniramous in all tanaidomorphs (Sieg 1983).
20. Number of terminal setae on the maxillule palp (1 = more than two setae, 2 = two
setae). This character was included to resolve out-group relationships; aspseudomorphs and some
members of the Tanaeoidea have more than two maxillule palp setae (Sieg 1983). It must be
noted here that Sinelobus stanfordi with its two terminal maxillule palp setae differs in that
respect from others of the Tanaeoidea and was incorrectly coded by Larsen and Wilson (2002).
204
Twenty in-group species were not coded for this character in the analysis because of literature
deficiencies (not described or illustrated), but these species presumably have only two setae since
they are members of the Paratanaioidea.
21. Number of terminal spiniform setae on the maxillule endite (1 = eleven, 2 =
ten, 3 = nine, 4 = eight, 5 = seven, 6 = five). This character was assumed by Sieg (1977) to be
diagnostic in separating genera of the Pseudotanaidae, especially among the Cryptocopinae
whose members have setae numbering from five to ten (Fig. 48). In fact, the chief distinguishing
character between Iungentitanais and Paraiungentitanais is that the latter has nine maxillule
endite spines and the former has five. Nine setae is universal among the Pseudotanainae except
for Pseudotnais (A) scrappi and P. (P) inflatus, both with seven. The reliability of this character
is questionable because the small structural size of the endite, and subsequent difficulty in
orientation, renders it subject to misinterpretation.
22. Maxilla complexity (1 = well developed, 2 = rudimentary). This character was
included to resolve out-group relationships; apsedomorphs have a maxilla with multiple lobes and
specialized setae whereas in tanaidomorphs, the maxilla is small, oval and featureless (Sieg
1983).
23. Maxilliped basis degree of fusion (1 = completely separate, 2 = about half fused, 3 =
fused except for a distal notch, 4 = completely fused). This character shows transition between
the primitive (apseudomorph) and apomorphic (tanaidomorph) states and is used here to resolve
relationships among the four out-groups (Fig. 49). Some degree of separation is seen in the three
species of Cryptocopoides and in one species (Pseudolanais borceai) of the Pseudotanainae,
although the coding of the latter was based on a questionable illustration (Bacescu 1960).
24. Maxilliped endites degree of fusion (1 = completely separate, 2 = fused between 1/3
and 2/3 length, 3 = fused except for notch on the distal margin, 4 = completely fused). As in the
previous character, the degree of fusion of the maxilliped endites is considered to indicate a
transition between primitive and derived states (Sieg 1983). The endites are completely separate
205
in the out-groups and all the Cryptocopinae thus; this character is diagnostic for the
Pseudotanainae whose members display varying degrees of endite fusion (Fig. 50). Species with
the more derived degrees of fusion (states 2 and 3) were found in the Akanthinotanais clade and
in clade B while those with a lesser degree of fusion were predominantly in clade A. States 4 and
2 were diagnostic on a generic level for Mystriocentrus and Parapseudotanais respectively.
Pseudotanais (P) inflatus was coded as state 1 based on a questionable illustration (KudinovaPasternak 1973).
25. Maxilliped endite setation (1 = numerous complex setae and coupling hooks present,
2 = two plumose setae, 3 = cusps plus setae, 4 = cusps only, 5 = setae only, 6 = without setation).
As in the mandible pars molaris complexity, this character is highly plastic and probably reflects
individually evolved feeding strategies. The first two highly complex character states are found
only in the out-groups and less complex states 3 to 6 varied widely among the Pseudotanaidae ingroup species with no apparent trends discernable (Fig. 51).
26. Cheliped degree of forcipation (1 = strongly forcipate, 2 = weakly forcipate - small
gap present, 3 = not forcipate). Cheliped forcipation is a derived condition probably related to
feeding mode and is likely evidence of an independently evolved response to food type or
availability. The seven species with strongly forcipate chelae (state 1), including the genus
Mystriocentrus for which this feature is diagnostic, were all in clade B (Fig. 52). Six other
Pseudotanainae species with lesser degrees of forcipation (state 2) were scattered among the subfamily.
27. Cheliped carpus length vs. width (1 = up to 1.9 times longer than broad, 2 = from 2.0
to 3.9 times longer than broad, 3 = greater than 4.0 times longer than broad). The trait for a
shorter cheliped carpus (state 1) was the most common condition among the species tested and
was dispersed equally among all clades. No apparent trends for a longer carpus (state 2) were
evident and only one species, Iungentitanaisprimitivus, had a carpus of 4.0 times longer than .
broad (Fig. 53).
206
28. Cheliped carpus number of inferior setae (1 = ten or more. 2 = three. 3 = two, 4 =
one). The number of inferior carpal setae was useful for resolving relationships among the outgroups (ten or more in apseudomorphs) but showed only slight variation among the pseudotanaid
species with nearly all having the normal two inferior setae condition (Fig. 54).
29. Cheliped propodus length vs. width (1 = up to 2.4 times longer than broad, 2 = 2.5 to
4.0 times longer than broad, 3 = greater than 4.1 times longer than broad). Even though the
overall consistency index was low for this character (0.182), the propodus length appeared to be
diagnostic in that the shorter length was prevalent in the out-group and in five of the eight
Cryptocopinae species (Fig. 55). All but five of the Pseudotanainae had a propodus length of
greater than 2.5 times longer than broad (character state 2) and two of these, both of the subgenus
Akanthinlanais, had a propodus length of greater than 4.0 times longer than broad; this may
indicate that an elongated propodus is a derived condition among pseudotanaids.
30. Cheliped propodus length vs. carpus length (1 = less than or equal to 1.5 times length
of carpus, 2 = greater than 1.5 times length of carpus). A lengthened propodus in comparison to
the carpus length (state 2) is apparently a derived feature seen in most of the Pseudotanainae, with
the notable exception of eight species of the subgenus Akanthinotanais (Fig. 56). All species of
Pseudotanais sensu stricto as well as those of the Mystriocentrus and Parapseudotanais had a
cheliped propodus of greater than 1.5 the length of the carpus, whereas all out-group and
Cryptocopinae species characteristically have shorter, more compact chelae (state 1).
31. Cheliped fixed finger number of inferior setae (1 = numerous, 2 = two, 3 = one, 4 =
none). The number of inferior setae on the cheliped fixed finger was useful in resolving
relationships among the out-group species as well as among some in-group species of
Pseudotanainae. Apseudomorphs generally have numerous inferior setae (six to seven in
Carpoapseudes) while all but two of theCryptocopinae species have two (Fig. 57). The normal
presumably derived condition among Pseudotanainae is one inferior seta; however, one of the
diagnostic characters for the genus Parapseudotanais is the ancestral state of two inferior setae.
207
This character was excluded from Larsen and Wilson's (2003) analysis of the Paratanaidoidea
since it was earlier shown to be unstable in a molecular analysis of Paralanais (Larsen 2001) but
was used as a diagnostic character by Guererro-Komritz and Brandt (2005) in an analysis of the
Akanthophoreinae group of tanaids.
32. Cheliped dactylus percent of propodus length (1 = up to 49.9 %, 2 = 50.0 to 63.0 %,
3 = greater than 63.1%). The dactylus percentage of propodus length is likely reflective of
independently evolved feeding modes, similar to that exhibited by degree of forcipation. There
were no clear patterns among the clades for dactylus percentage length except that a percentage
longer than 63.1% (state 3) appeared to be diagnostic for the genus Mystriocentrus (Fig. 58).
33. Cheliped dactylus width vs. fixed finger width (1 = narrower, 2 = subequal, 3 =
broader). The comparative widths of the dactylus vs. the fixed finger appeared to be a highly
variable among all species tested with no clear trends exhibited. It is noticeable that six of the ten
species of the subgenus Akanthinotanais have a dactylus that is narrower than the fixed finger,
but this fact is probably non-diagnostic given that the trait also appeared in 13 other species of
Pseudotanainae as well as in two out-group and two Cryptocopinae species (Fig. 59). The three
species that had broad dactylus widths (state 3), Pseudotanais (P)jonesi, Curtichelia expressa,
and Paraiungentitanais longidigitatus, all had chelipeds that were drastically dissimilar, further
lessening the importance of this character.
34. Cheliped dactylus proximal seta (1 = present, 2 = absent). A proximal seta on the
cheliped dactylus was present in all but six of the Pseudotanainae species but absent in half of the
Cryptocopinae and three of the four out-group species (Fig. 60). The fact that this seta was
mostly absent in the out-groups and present in the in-groups is counter-intuitive to the assumption
that loss of setae is evidence of a derived character state. Thus, this character with its low CI
value (0.100) was less essential in establishing polarity between the out-group and in-group
species tested.
208
35. Marsupium formation (1 = by four pairs of oostegites, 2 = by one pair of oostegites,
3 = by ovisacs). The formation of the marsupium has long been the most definitive character
separating the Pseudotanaidae from other tanaidomorphs, all of which posses four pairs of
oostegites. In spite of Bird and Holdich's (1989) discovery of some specimens of
Cryptocopoides arcticus possessing four oostegites which made questionable the placement of
the genus in Pseudotanaidae, it was decided to study the phylogeny using the original description
of only a single pair for that species. In reality, most species descriptions of pseudotanaids lack
mention of the ovigerous state so in this research a single pair of oostegites was assumed and was
coded as such (Fig. 61). The ovisac character state found only in the Tanaidae, was represented
here by Sinelobus stanfordi. A separate search for the most parsimonious tree was conducted to
test the effect of omitting this character, the results of which appear elsewhere.
36. Pereopod ischium presence on all pereopods (1 = present, 2 = absent). The character
for presence or absence of the pereopod ischium was included to resolve relationships among the
out-groups. Sinelobus stanfordi of the family Tanaidae lacks an ischium article on all pereopods,
whereas the article is present in all other species being tested, both out-group and in-group.
37. Pereopod 1 basis length vs. width (1 = up to 5.0 times longer than broad, 2 = 5.1 to
7.9 times longer than broad, 3 = 8.0 to 9.9 times longer than broad, 4 = greater than 10 times
longer than broad). A pereopod basis length of less than 5.0 times greater than broad, seen in the
four out-group species, is apparently a derived condition given that ratios of greater length,
though variable, were present in all in-group species (Fig. 62). Extreme basis lengths of greater
than 10.0 times longer than broad were seen in only six Pseudotanainae species.
38. Pereopod 1 basis setation (1 = four or more setae, 2 = one to three simple plus broom
setae, 3 = broom setae only, 4 = one to three simple setae only, 5 = setae lacking). Evidence of
transition from primitive to derived as demonstrated by pereopod 1 basis setation was only
sparingly indicated for some clades and was considered a weak character since two of the four
out-group species showed sparse setation (state 4), thus polarity was not firmly established. Most
209
notably, six of the ten Akanthinotanais species lacked setae (state 5), somewhat strengthening this
clade, but in more than half of the remaining pseudotanaid species, setation complexity equaled
or surpassed that of the out-groups (Fig. 63). Crytocopinae species had shorter propodus lengths
(state 1); but with the large number of Pseudotanainae species also having short propodus lengths,
this character had dubious phylogenetic value.
39. Pereopod 1 ischium number of setae (1 = two setae, 2 = one seta, 3 = lacking setae).
The setation of pereopod 1 ischium showed no systematic change from primitive to derived (Fig.
64). All out-group species and 44 of the 57 in-group species had a pereopod 1 ischium with a
single seta. Of the others, 11 species from four different genera lacked setae and only two,
Pseudotanais (A) gerlachi and P. (P) mediterraneus had two setae on the ischium.
40. Pereopod 1 merus setation (1 = numerous including several marginal setae, 2 = three
setae, 3 = two setae, 4 = one seta, 5 = setae lacking). The merus setation of pereopod 1 showed
various degrees of reduction from plesiomorphic seen in the apseudomorph out-group (state 1) to
the more apomorphic condition present in the other out-group species as well as the in-group
species of Pseudotanaidae (Fig. 65). The typical state was either one or two setae, with setae
lacking in 12 species, and three setae being present in only one, Pseudotanais (P) nodenskioldi.
No clear-cut trends were apparent for this character.
41. Pereopod 1 carpus setation (1 = numerous setae including several simple and
spiniform, 2 = five setae, 3 = four setae, 4 = three setae, 5 = two setae, 6 = one seta, 7 = setae
lacking). As in the previous character, the varying degrees of reduced pereopod 1 carpal setation
showed no clear-cut transition among the in-group species (Fig. 66). The apomorphic state with
numerous setae exhibited by the apseudomorph out-group appeared to be distant from the
reduced number present in the other out-group and the in-group species, the latter having two or
less setae (states 5-7) as the most common condition. In one case, the presence of four carpal
setae appeared to help define a clade consisting of Cryptocopoides pacificus and C. sp. A as
distinguished from the type species of the genus, C. arcticus.
210
42. Pereopod 1 propodus length vs. width (1 = less than 6.0 times longer than broad, 2 =
6.1 to 8.0 times longer than broad, 3 = greater than 8.1 times longer than broad). A trend toward
increasing length of the pereopod 1 propodus appeared to be evident, especially among species in
clade A (Fig. 67). Six of the 10 species of the subgenus Akanthinotanais had a propodus
exceeding 6.0 times longer than broad (states 2 and 3) and seven of the Pseudotanais sensu
stricto species had a propodus length exceeding 8.1 times longer than broad (state 3).
Meanwhile, all out-group species and five of the eight Crytocopinae species had shorter propodus
lengths (state 1); but with the large number of Pseudotanainae species also having short propodus
lengths, this character had dubious phylogenetic value.
43. Peropod 1 propodus setation (1= numerous setae including several distal simple
setae, 2 = four setae, 3 = three setae, 4 = two setae, 5 = one seta, 6 = setae lacking). The number
of propodal setae on pereopod 1 showed a general trend of reduction in number from out-group
species with four or more to in-group species having three or less (Fig. 68). The trend in
reduction was also manifested in species of the in-group subfamilies Cryptocopinae and
Pseudotanainae. Seven of the eight Cryptocopinae species exhibited two to three propodal setae
(states 3 and 4) while only nine of the 49 Pseudotanainae species had two setae (state 4), the rest
showing a reduction in setae from zero to one (states 5 and 6).
44. Pereopod 1 dactylus setation presence or absence (1 = setae present, 2 = lacking
setae). Even though the dactylus seta was present on two of the out-group species including the
apseudomorph with its highly setose dactylus, its importance in establishing polarity was lessened
by its sporadic occurrence among in-group species (Fig. 69).
45. Pereopod 1 dactylus setal type (1 = numerous or complex, 2 = single and
exceptionally long, 3 = single and short). The character describing the dactylus seta on pereopod
1 was included to make distinctions among those species with a setose dactylus (Fig. 70). It
served mainly to further define the genus Parapseudotanais, both species of which have a single
elongate dactyl seta on the first pereopod.
211
46. Pereopod 1 length of dactylus-plus-unguis compared to length of propodus (1 =
distinctly longer, 2 == subequal, 3 - distinctly shorter). The comparative length of the terminal
article vs. the propodus of pereopod 1 had diagnostic value for the Pseudotanais subgenus
Akanthinotanais (Fig. 71) as all but two species bore a short dactylus (state 3). This character
also further defined the genus Mystriocentrus, both species of which were state 3. The elongate
condition (state 1) of the dactylus was found only in two species, Pseudotanais (P) lilljeborgi and
P. (P) spicatus.
Al. Pereopods 2 and 3 basis setation (1 = broom plus simple setae, 2 = broom setae only,
3 = simple setae only, 4 = setae lacking). The basis setation of the second and third pereopods
displayed no definite trends (Fig. 72). All four character states appeared to be evenly distributed
among the clades and no polarity could be established from the out-groups. A complete loss of
setae was evident in only 16 in- group species and four others were not scored due to lack of
information from the original descriptions.
48. Pereopods 2 and 3 ischium setation (1 = three setae, 2 = two setae, 3 = one seta, 4 =
setae lacking). The number of ischium setae on the second and third pereopods, with a few
exceptions, showed a reduction from three in the apseudomorph out-group to one in the majority
(43) of in-group species (Fig. 73). Two ischium setae were present in three species of the
subgenus Akanthinotanais, in Pseudotanais (P) mediterraneus, and in the out-group species,
Hargeria rapax. Only in one species, Parapseudotanais sp. A, was the ischium setae lacking on
the second and third pereopods.
49. Pereopods 2 and 3 merus setation (1 = numerous marginal setae, 2 = three setae, 3 =
two setae, 4 = one seta). The number of merus setae on the second and third pereopods showed a
reduction from two or more in the out-groups to one in only a few of the in-group species (Fig.
74), the majority retained the ancestral two setae on this article and one species, Mystriocentrus
sp. A, had three setae. Reduction was more evident in the Cryptocopinae. Of the eight species of
that sub-family, five had only one seta, two were unknown due inadequate description and only
212
one, Crylocope abreviata, was with two meral setae. In the Pseudotanainae, reduction to one seta
was seen in only eleven scattered species.
50. Pereopods 2 and 3 merus spatulate setae presence or absence (1 = present, 2 =
absent). The character for spatulate setae on the merus of the second and third pereopods was
included as diagnostic for the genus Mystriocentrus; all other species lacked these specialized
setae on that particular article (Fig. 75).
51. Pereopods 2 and 3 carpus blade-like setae presence or absence (1 = absent - all
simple or spiniform, 2 = present). The presence of specialized blade-like setae on the carpus of
the second and third pereopods was the chief diagnostic character used by Sieg (1977) to separate
species of the subgenus Pseudotanais, senso stricto, from the subgenus Akanthinotanais which
lack such setae, having simple non-modified setae instead. Accordingly, in this research
specialized blade-like setae are clearly seen as a distinct apomorphic character defining those
species that have them as separate from species more ancestral including the Akanthinotanais as
well as the Cryptocopinae and species of the out-groups (Fig. 76).
52. Pereopods 2 and 3 carpal blade-like setae length vs. length of propodus (1 = half or
less the length of the propodus, 2 = greater than half the length). Of the 36 species having bladelike setae, 21 have blades shorter than half the length of the propodus (state 1) and 15 have longer
blades (state 2). These two types were separated mostly into two groups designated as clades A
and B with 19 and 16 species respectively (Fig. 77). Clade A was composed mostly of longbladed types (13 to 6) while short-bladed types predominated in clade B (14 to 2). The 31 species
of Pseudotanais sensu stricto, were almost equally divided into 15 with long blade-like setae (13
in clade A, two in clade B) and 16 short-bladed types (ten in clade B, six in clade A).
53. Pereopods 2 and 3 length of dactylus-plus-unguis compared to length of propodus (1
= subequal, 2 = 2/3 to 3/4 length, 3 = about half length, 4 = distinctly less than half length).
Even though a longer dactylus length (states 1 and 2) was a feature of three of the out-group
species, no clear trends toward modification of this trait could be seen among any of the in-group
213
clades (Fig. 78); thus no phylogenetic importance could be placed on this highly inconsistent
character.
54. Pereopod 3 size comparison to pereopod 2 ( 1 = similar in proportion, 2 =
disproportionately smaller). Pseudotanais (P) unicus is distinct from its congenitors in that the
third pereopod is smaller than the second in all proportions.
55. Pereopod 4 basis length vs. width (1 = up to 4.0 times longer than broad, 2 = 4.1 to
5.9 times longer than broad, 3 = greater than 6.0 times longer than broad). The length of the
fourth pereopod basis appeared to have little diagnostic value among the species tested even
though three of the four out-group species had the shortened form of this character (state 1). The
two other character states for greater lengths were evenly spread among the in-group species. An
elongate basis (state 3) was characteristic of the genus Mystriocentrus but was found in only six
species of Pseudotanainae and two species of Cryptocopinae (Fig. 79).
56. Pereopods 4 to 6 basis setation (1 = simple and broom setae, 2 = broom setae only, 3
= simple setae only, 4 = setae lacking). Simple and broom setae on the basis of the fourth
through sixth pereopods were present in both combinations (states 1 and 2) for a majority of
species tested with only a slight trend toward reduction seen in the Akanthinotanais group (Fig.
80). It is worth noting that the highest number of species with broom setae only (state 2) was in
Pseudotnainae clade B where 63% shared this feature and the majority of those with a
combination of broom and simple setae (state 1) were in clade A. Of the 16 species lacking basis
setae (state 4), five each were in the Akanthinotanais and Cryptocopinae groups.
57. Pereopods 4 to 6 ischium setation (1 = two unequal setae, 2 = two equal setae, 3 =
one seta). Ischium setae on pereopods 4 to 6 did not demonstrate polarity in that reduction in
setae between the out-group and in-group species was not apparent, thus phylogenetic importance
attributed to that character is limited in this case. However, Pseudotanainae clades A and B were
distinctive for this character, with few exceptions, in that the species in them possessed two and
one setae respectively (Fig. 81).
214
58. Pereopods 4 to 6 merus setation (1 = numerous setae, 2 = two setae, 3 = one seta).
The number of meral setae on pereopods 4 to 6 ranged from numerous in the apseudomorph outgroup to one or two in all other species including the other three out-groups, thus limiting its
value as a polarizing feature (Fig. 82). The character had no major diagnostic value among the
in-groups except that it helped define the genera Parapseudotanais (state 2) and Mystriocentrus
(state 3). The dominant condition for this character was two setae (state 2) which occurred in all
but twelve of the in-group species.
59. Pereopods 4 to 6 carpus blade-like setae presence or absence (1 = absent, 2 =
present). As for the carpal blade-like setae on pereopods 2 and 3 (character 51), the presence of
these modified setae was likewise used by Sieg (1977) to define the subgenus Pseudotanais,
senso stricto, as separate from the subgenus Akanthinotanais (Fig. 83). An exception is that
species of the genus Parapseudotanais have blade-like setae only on pereopod 2, thus the absence
of these specialized setea on pereopods 4 to 6 (state 1) is diagnostic for that genus.
60. Pereopods 4 to 6 carpus presence of additional specialized setae (1 = with additional
bifid-tipped setae, 2 = with additional spatulate setae, 3 = with abnormally long superior simple
seta equaling or exceeding length of propodus). The presence of these additional specialized
setae served to resolve or define certain clades among the Pseudotanaidae and also in the genus
Crytpocopoides. Specifically, both species of Mystriocentrus and two species of Cryptocopoides
have spatulate setae. Seven species in Pseudotanainae clade A were equipped either with
additional superior elongate or bifid-tipped setae, thus adding strength to that particular grouping
(Fig. 84).
61. Pereopods 4 to 6 propodus broom setae presence or absence (1 = present, 2 =
absent). Broom setae on the propodus of pereopods 4 to 6, an apparent apomorphic character,
occurred in 13 Pseudotanainae species, nine in clade A and four in clade B (Fig. 85). A single
Akanthinotanais species, Pseudotanais (A) similis, and two species of Cryptocopoides, C.
pacificus and C. sp. A, also possessed these broom setae. The quality of this character is
215
somewhat in doubt because of its sporadic appearance in the literature, possibly resulting from
the study of poor quality specimens - broom setae are easily lost or overlooked.
62. Pereopods 4 to 6 dactylus fusion with unguis (1 = fused with unguis to form a claw,
2 = not fused with unguis). The fusion of the pereopod dactylus and unguis forming a "claw" is
considered apomorphic primarily in tanaidomorphs (Larsen 2005) and is generally associated
with shallow water species (Larsen and Wilson 2002). In the Pseudotanaidae, Sieg (1977) used
this character to distinguish members of the subfamily Pseudotaninae (fusion present) from the
Crytocopinae (no fusion). In accordance with Sieg's hypothesis, the fused dactylus (state 1) was
strongly diagnostic for separating the Pseudotanainae from Cryptocopinae in this research (Fig.
86).
63. Pleopod complexity (1 = rami well developed with terminal and lateral setae, 2 =
rami elongate with terminal and subterminal setae, 3 = rami elongate with terminal setae only, 4 =
rami vestigial without setae, 5 = pleopods lacking). The reduction in complexity of pleopods
from well-developed to elongate with terminal setae clearly separates the subfamily
Pseudotanainae from the out-groups as well as from most of the Cryptocopinae (Fig. 87). The
presence of well-developed pleopods helped define a clade within the Cryptocopinae (clade C)
composed of four monospecific genera {Curtichelia, Iungentitanais, Paraiungentitanais, and
Latitanais) as separate from four others with less complex or vestigial pleopods.
64. Uropod complexity (1 = biramous, 2 = uniramous). The character for uropod
complexity was included to resolve out-group relationships; The Tanaidae representative,
Sinelobus stanfordi, lacks uropod exopods.
65. Uropod exopods number of articles (1 = more than two articles, 2 = two articles, 3 =
one article with pseudo-articulation, 4 = one article). A reduction in the number of exopods
articles occurred in all species tested other than the apseudomorph out-group (Fig. 88). The large
majority of Pseudotanaidae species had two exopods articles but there were seven occurrences of
a single article and four species with a "pseudo-articulated" article.
216
66. Uropod cxopods length vs. endopod (1 = less than 1/6 length of endopod, 2 = 1/3 to
half length of endopod, 3 = from 2/3 to subequal length of endopod). Comparative lengths of the
endopod and exopods appeared to be diagnostic at the family level (Fig. 89) as most had an
exopods length of at least 2/3 the length of the endopod (state 3) whereas two of the out-group
species {Carpoapseudes prospectnes and Typhlotanais parvus) had much shorter exopods (state
1). Of the eight species of Pseudotanainae having a short exopod (state 2), six were present in
clade B. Among these, character state 2 helped to define the genus Mystriocentrus.
67. Uropod exopods proximal article proportion of ramus (1 = about 1/4 length of ramus,
2 = 1/3 to 1/2 length of ramus, 3 = about 2/3 length of ramus). The character for the uropodal
exopod proximal article length proportion had little value within the family Pseudotanaidae other
than to enhance the description of individual species and in some cases distinguish between
otherwise closely related species (Fig. 90). For example, character state 3 is diagnostic in
separating Cryptocopoides arcticus from its congenitors, C. pacificus and C. sp. A, both of which
are state 2. This character was apomorphic (state 1) only in the apseudomorph out-group and
state 3 occurred only in five in-group species.
68. Uropod endopod number of articles (1 = more than two articles, 2 = two articles, 3 =
one article with pseudo-articulation, 4 = one article). A reduction in number of endopod articles,
similar to that of the exopod, occurred for all in-group species (Fig. 91). Three of the out-group
species had three or more endopod articles (state 1) and all of the in-group species had two or
less. In the Pseudotanainae there were five species with pseudo-articulated endopods and five
with a single article (states 3 and 4). No particular distributional pattern was noticed for this
character among in-group species.
69. Uropod endopod proximal article proportion of ramus (1 = 1/10 to 1/4 length of
ramus, 2 = 1/2 to 1/3 length of ramus). The short proximal article of the endopod (state l)was
seen only in two of the tanaidomorph out-group species (Sinelobus stanfordi and Hargeria
rapax), whereas in all in-group species the length of this article was at least half the ramus length
217
(state 2). Thus, no diagnostic value other than out-group polarity could be attributed to this
character (Fig. 92).
218
Oephslothorax length
length of pereonites 1
CH3 shsrlfe*
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P s e u d o t a n a i s (P) unicus
P s e u d o t a n a i s ( P ) nt e d i t e r r a n e w s
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P a ra p s e u d o t a n a is s p . A
P s e u d o t a n a i s (.A) l o n g i p e s
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Crytoeopoides arcticus
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C r y p t o c p o i d e s s p. A
l u n g e n t r t a n a is p r i m r t w i s
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Figure 33. Trace for character 1 - cephalothorax length
219
Eyeiobes present or absent
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"O P'.«•!!<}« L i tiJS>. ( A ) 0 4MlaiClU
P i , <»«<loLi j u s - , ( A ) g < i u s s «
P';,<»!!<!« SJ «js>, ( A ) \ W i l l i *
S'* ••', <* I! d HV I J tl ^ S <, ( A ) SIS ^1 L i y <»!»<• I:",
Pj> ewdntamai* (AJ s o . a p p i
ajPsftnidotanais (A) g u i l l e i
Cryptocop^ abreviata
Crylocopoides aietious
Crytocopoides pacificus
Cryptocpoides s p . A
lungentifaiiats primitwus
Para iuti gentitan a is i o n g i d i g i t a t u s
Lafitanais belt le mis hew
Curtiehelia e g r e s s a
_, r i S i n « l o b u * ^ t a n l o t d i
"j'—p H . i n i e i i a l i p a i t
fypliloLdiiJIS
J*JI¥«J
C J SJM* .![»':, I? 51 <!«?':, J i l l l ' , | ( ( ' i ; h l l " .
Figure 34. Trace for character 2 - eyeiobes present or absent
220
P^reonlte abbreviation
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotasia^,
P v e « d o tiisi^i'i
P i eu«lo i»i i i j f ,
P1!. « t x i o L i n <tii
P% c u d o l i i n A i t
P<4 e u d o l a n j r v
Ps e u d o t a n a i s
Pseudotanais
Pseudotonais
Pseudotanais
P s e u d o t a nais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Ps*udoianais
P« * u d « i a « a i s
Jr
(P)spatula
(P) s p. O
(P s p . P
(P| a Ifinis
(P] s c a l p e l l u m
(P lortgi>> pif> us
(I*
(P
(P d e i l t l C u l a i t l *
(P
(P
(P' •I ipp<»niCII%
(P' s p C
(P[ I on n is el© s us
(P] ©orollatos
(P| o c u l a t u s
(P) m
b oer xc iekaoil p o s
(P u n i e u s
(P m e d i t e r r a n e u s
(P hi Hates
(P s p. A
(P v u l s «l la
P i t u d o t a f t a t S ( P 1ai«i<;«itai.<i
Nty\ t i i o c e n t r t i t '» n a l i i i
ft
My% littMM* nltws %
P < t # u d o l * n ; m (P Abyt «. i
P & f i i < l o t i n < i K (P
Ps ewdoM n*»••"*. (P) c a l r f o m i * ms is
P s e u d o t a n a i s (P' f o r e i p a t u s
B Jr ™ I * "H " RjJS 88 sis S3)- <L H^
joues i
P s e u d o t a n a i s (P' eolonms
P s e u d o t a n a i s (P' i i l l j e b o r g i
P s e u d o t a n a i s (P' n o d e n s k i o l d i i
Para p s e u d o t a n a i s a b ^ s s a l i s
Parapseudotanais sp. A
P s e u d o t a n a i s (A) i o n g i p e s
P s e u d o t a n a i s (.AJsieys
P s * t » d o t a n a i s (A) makr«*Ilkiix
P-s « u d » t a n a i i (A) tiii o i t v n v e n i
P t e u d o t a n a i i (A) g e i l j e h i
Pv eud«L«n<*it ( A ) g d » ' . s i
P M - m l o y i i - n i , (A) * mi ilis
P$ # w d o i a n A i * (A) m a l j y e n ^ n
P s f t u d o t a i u u s (A) s «r j i p p i
P s e u d o t a n a i s (A) guiSiet
• Cryptaeopie a b r « v i a t a
j Crytoeopoides arcticus
Crytocopoides paclficws
JU C r y p t o c p o i d e s s p. A
I
o lungentitanais primitais
I 1 '~*n P a r a i u n g e n t i t a i i a i s Songidigitafus
_zu L a t i t a n a i s bek le mis ft e«i
"O C t i r t i e h e l i a e x p i t s s a
U Siiteiobus stanfordi
t f a r § « r i a fapaK
33 Typhi «ta MIS' parvus
33 C a r p « t l t p S » » l d « p!«>£p*$tn£S
Figure 35. Trace for character 3 - pereonite abreviation
Antennule number
of articles
C m ttt*e "*i£"i fc JI a?l(£ $ *
• H I -in.- ir.-.'r?s
Pseudotaisats
Pseudotanais
Pseudotauais
Pseudotanais
Pseudotanais
P-seud«>lanars
P *, e w i i o L m JI»,
(Pjspatula
( P ) s p. O
( P ) s p. P
(PJ a f l i n t s
( P ) s ©alp«SSum
( P ) l u n y k pin us
( P ) d «n»tsc uS^iis <i
P •, «MM!<> t j si•.«!', ( P ) »« }>!«: J I I E * .
P"i «MHlO Ll !i,»I«, ( P ) Wilj^lJfl
P<i **lnlttLl !>,ll", ( P ) n !pp ( H I K I H
P i *MH1*» I J njit't ( P } ass J C i « cSn>l**'.
P 5 4»u4«tia S*.»!S ( P ) 5 p C
P s e u d o t a nails
Pseudotanais
Pseudotanais
Pseudotamais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudolanaic
(P)longkei«sus
(P) corollatus
(P) ocuiatus
(P) mexikolpos
(P) borceai
(P) unicus
(PJ i n e d i t e r r a n e u s
(P) inflates
(P) s p A
(P)¥utselia
(P") t j l t i t u S j t j
Myilnocriiliut
•. p A
P<k <MS<I« l j ! 5S J ! ' ,
(P)*tb¥<,%i
Pm>iliiitMn<)i'i ( P ) b-t s *n, it-J ss h
P s «• u d 0 1 * n .* ss ( P ) «*ilrfo»«*i« n s i s
Ps eud»ti«n.sis ( P ) f o r e i p a t u s
Pseudotanais (P)jonesi
P s e u d o t a n a i s (P) colonws
Pseudotanais (P) iilljeborgi
Pseudotanais (P) nodenskioldii
P ara ps e u d o t a n a i s a b y s s alas
Para pseud otanais s p . A
P s e u d o t a n a i s (A) l o n g i p e s
P s e u d o t a n a i s (A) s ie g i
Ps e u d o t a n a i s (A) en isk»*> tins':.:
P v e u d o t a n a i s (A) m o r t e n s e n i
P •» »*««i<> !*i i»,ta':.
P * tMido I J !i*ii",
P >, *Mjrf<» i j ll/tl',
P<, **«nlo l„i I»,IS'.
(A)
(A)
(A)
(A)
g <-I!.H:!II
<| J I I < . ' , e
Sim ill*
m a l a y e n t 1*
Ps e u d o t a n j i i s ( A ) i c s * i p p i
P s c u d o t a n a i s (A) g u i II c i
Cryptoeope abreviata
C r y t o c o p o i d e s a reti cus
Crytocopoides pacificus
C r y p t o c p o i d e s s p. A
lufigentitanais primitvus
P ara iun g c n t i t a n a is t o n gid i g i t a t u s
Latitanais beklemishewi
Curtitshelia e x p r e s s a
Sinei<*bu<s; S t a n f o r d *
I! *i«K*- M j ««i|> J:X
£)(• d i p Q j p t e u d e t ps «"» p m i n e s
Figure 36. Character 5 trace - antennule, number of articles
Antennule article
percent length of
en <42
E = | 42.1 to 49.9
Psenidotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Ps v u d u i a n i i v
(P) s p a t u l a
( P ) s p. O
( P ) s p, P
( P ) affinis
( P ) s ©alpellwm
(P) lunyispinus
( P ) dentieulafaii.
P •> v u d o Id I I A I * ( P ) s p i c a l » i s
P •. * » « d o Li i i ^ ! ' , ( P ) *s!|>ijn
P* <Mssioij n<Mi\ ( P ) n ippoitMiU*
P^ imdotanais. ( P ) m j « i # « l i « l » %
Ps^Mdotamais (P) $ p, t
P s e u d o t a n a i s ( P ) I on 9 k e t o s u s
Pseudotanais (P) corollatus
Pseudotanais (P) oeulatus
Pseudotanais (P) mexikolpos
Pseudotanais (P) borceai
Pseudotanais (P) unicus
Pseudotanais (P) med(terranews
P s e u d o t a n a i s (P) i n f l a t e s
Pseudotanais ( P ) s p . A
Pseudotanais (P) vulsella
Pseudotanais (P) lalciculata
Mjciiioceitlmi
•. e i u l m
P i 4*UfIotalt*l* ( F ) j l l f , M
r s f u d o tamais ( P ) b a r » * n j u b
P s e u d o t a n a i s (PJ c a l i f o m i e ns is
Pseiidatanais (P) forcipatus
Pseudotanais (P)Jomesi
Pseudotanais (P) colonus
Pseudotanais (P) liiljeborgi
Pseudotanais (P) nodenskioldii
_npmaPara p s e u d o t a n a i s a b y s s a l i s
li_ajPara pseudotanais s p . A
Pseudotanais (A)Iongipes
Pseudotanais ( A ) s i e g i
P s e u d o t a n a i s (A) m a k i o t t i i i x
P s e u d o t a n a i s (A) m v i t e n S e n i
P s e u d o t a n a i s (A) ( j c i l . n : l i i
Ps t>«ds ta «»*us (A) (| ,m«, •, 1
P «. i>«dg ta nais ( A ) s i m i l i s
P t e u d o t a n a i s (A) m A la. yen*, m
P s e u d o t a n a i s (A) %. ^ r a p p i
s P s e u d o t a n a i s (jA}-§uillei
C r f p t o ^ o p e abreviata
rctit
SCriflocopoides
j J—:acrytocopoides paeifficus
l—:uCrypt»cpoides s p . A
__J"p:Dlusigentitanais primitvus
I 1 ~ n P a r a i u t i i g e n t i t a i ) a i s l o n g id ig Status
:oLatitartats b*sklemish«¥i
DCurtichelia express a
S i n s lob tis s t a n t f e i d i
H a r g e r i a rap ax
TyphIota M i s parwtif
DC a tp» a p t #»«*«* p f o s p t e t w t s
Figure 37. Character 6 trace - antennule, article 1 percent of total length of antennule
223
Antennule artlele 1
length x width
P s e u d o t a nais
Pseudotanais
Pseudstanais
Pseudotanais
Pse«d«ljtiai*;
P*» ttudota isais,
P.V e u d e f a n a i t
Pt,«udaljnjii
(P)
(P)
(P)
(P)
(P)
(P)
(P)
(f)
s patuia
s p. O
s p. P
a iff in is
s calpellum
ivnyi&iiinu^
d »Mih<:«! ji«,s
', pic j t i n
P", <M£ <J o Si! si ,i i', (f*) ii ij>|* in>n:«',
P5 eudo la »> Jis (P) in J<: i« •cSt«»l<»<.
Ps ewdotanaiss (P) 5 p. t
PseMdotainais (P) long is etesws
P s e u d o t a n a i s (P) c o r o l l a t u s
P s e u d o t a n a i s £P]i oeuIatus
P s e u d o t a n a i s (P) m e x i k o l p o s
P s e u d o t a n a i s (P) b o r c e a i
P s e u d o t a n a i s (P) unieus ._
P s e u d o t a n a i s (P) m e d i t e r r a n e u s
P s e u d o t a n a i s (P) inflatus
P f e u d o t a i i a r s (P) s p. A
P i e u d o t a n a i i (P) v u l s e l l a
P •, t'u<l« Jj 11 JI-, (P) I.ilctctiLita
My4 Inoce n l i t i t •» r is ,:iSii',
My«> l»io«f* nlius ". |» A
P<k p u d o t a n a i ^ (P)*iby«,*i«
Ps r v d « i » n j i i s ( P ) b j i ^ R 4 « f i
Ps ewilofcanjMs.(P) i t j i l r t o i n i e n s i s
Ps e u d o t a n j i s (P) *<*rci|»at*i«s
P s e u d o t a n a i s (P) j o n e s i
P s e u d o t a n a i s (P) eolonus
P s e u d o t a n a i s (P) l i l l j e b o r g i
P s e u d o t a n a i s (P) n o d e n s k i o l d i i '
Para p s e u d o t a n a i s abyss alis
Para p s e u d o t a n a i s s p . A
P s e u d o t a n a i s (A) l o n g i p e s
Pseudotanais (Ai)siegi
P s e u d o t a n a i s (A) m a k r o l h r t x
P', t'«d« t j l U l s (A) M o l t e n * * m
P«. 4 > u d « l j l t 4 ^ (A) 0 * l l . * c l i i
Hs « u 4 « b i M n (A) 11 J!?i', 1, 5
Pv#u<i«t<i»4»% (A) % i m i l i *
P<s eudotri ndis (A) m ,atayiMi% K
P s e w d o t a n a i s (A) s ©r^ppi
P s e u d o t a n a i s (A) g u i l l e i
Cryptseope abreviata
C r y t o c o p o i d e s arctieus
Crytoeopoides paeifietis
Cryptoepoides s p. A
lungesntitanais primHwus
Paraiungentitam ais l o n g i d i g i t a t u s
Latitanais
taeklemishevlC u r t i c h e l i a express a
S«nel»bu« S t a n f o r d !
II .tl(|VIlJ lilpilK
Ey[>hlftij».ii«,
p.nvti',
V. jt i p o Apt, v u d r \ |mt"i p «* e S n «* *.
Figure 38. Character 7 trace - antennule, article 1 length vs. width
224
Antennule article 3
length vs. width
£=li»t»1.«
SMI ULteSLS
fssg
Pseudotanais
Pseudotanais
Pseudotanais
Pse«d«tanais
Pseudatanais
(P)
(P)
(P;
(P
(P
spatula
s p. O
sp.P
a If in is
s calpellum
P', t ' S i t l o l j ii,ji«; ( P iortfl is pin «•»
P >, « M M I « t j i i a i ' , ( P d eikitc tilths \
!*•> e m d a t r f » a i % ( P ». pit: Ain't
!*«•*>« i f « i j 11 j i', ( P
P <. ins d o I:4| »»a i«« ( P n i p p »;»!<:?!•,
I* >, <» u <1 <> !,i »i ..n •, £Pin Jt c i « « h«»! v-',
P a «udot«» n < I I ^ ^P «* p C
P s «si do ton ,i is
Pseudotanais
P s e u d o t a tiais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudtttanais
Pseudotanai-s
(P
(P^
(P'
(P\
(P;
(P)
(P)
(P)
(P)
(P)
l o n g is e t a s us
oculatus
mexikolpos
borceai
unions
m e d i t e r r a t t eus
Sofia t u s
s p. A
vulsella
!•*•:, «*««!« I j «!,:«!'» ( P ) i& i c • e u I «i I A
U f . UlOCt>nttUt 5
A
P i *Mid*>M«*ai<> ( P )
P * e u d o t j n , * ! * ( P ) h aw, n<i«!«
P ^ e v d o t A n a i * (P) c ^ l r f w i n i ^ n s i s
Psewdiatamjiis (P) f a r c i p a t u s
Pseudotamais (P) j o n e s i
Pseud»tamais (P) colonus
P s e u d f t t s n a i s (P) l i i l j e b o r ^ i P s e u d o t a n a i s (P) n o d e n s k i o l d i i
P a r a p s e u d o t a n a is a b y s s a i i s
Parapseudotaitais sp. A
P s e u d o t a n a i s (A) i o n g i p e s
Pseud*»ta«ai<» ( A ) s i e g i
P s e u d o t a n a i ? (A) i n a k r o t h r i x
P> e t l d o L m a i v (A) to a i t e l t * « si a
P * w u d o l a ii»«i-» (A) « e i l j c h i
P •> <»«d« J j »,!!», ( A ) 1| . H i ' . ' . •
P * «*nd# t a n a i - i ( A ) •• >m l i e .
P ^ t > i l d « U n ^ n ( A ) t n a M y P I T V Pi
li^wrewwmrtt-n-,wffBrffmfWM^
P M H i r i f t t a n a i * (A) * e»^ppi
P s e M d w t a n a i s (A) g u • II c •
Cryptncope abrewiata
C r y t o c o p o i d e s artstieus
C rytocopoides paeifieus
r y p t o c p o i d e s s p. A
l u n g e n t i t a n a i s primHhras
P a r a i u n g e n t i t a n a i s longidigjitattts
L a t i t a t s a is b t k Its i n i s l i ewi
Ctirtichelia e s f i e s s a
Siiielubiis Stanford!
H a r g « i i a rap ax
lyphioljsi.u:', parvus
CMfp9»f>s««4ms
Figure 39. Character 8 trace - antennule, article 3 length vs. width
pf«sp«s«ln#s
Antennuie distal article
length vs. width
CZJ 2to&9JC lonpH* Willi broMl
ti_J 4 t o 5 . 5 x
Psewdota nais
wPseudoiasiais
Pseudoianais
Ps e«id»tanais
Ps eudota nais
P i eudoianart
(PJ
(PJ
(P)
(PJ
(PJ
(P)
spatula
s JJ. O
s p. P
affinis
s ealp*Ilum
Buiiyi;, p i n u f
Pi» eutlo ta II.it1* ( P ) d t?»iticu! jh» >,
P * etstfo l a » aiv ( P ) ». !>n:jf«<>
P>, < M l l l l l I j l l , l l ' . ( ( ' ) » l l | , I / l
P'i <USt!o f j IIJI', ( P ) n ipp Oil I *!!«'.
P», <»*irfo I J ii-Ii», ( P ) HI JIC i « ch*l<"<
PsriiiltUnjiis
Ps e u d o U n a K
Pseudatanais
Pseudotanais
Pseudotanais
Pseudotaoais
Pseudoianais
p»ip»pwwB*Pseu(l(iijnais
Pseudoianais
Ps«swd«taiiiais
Pseud<»tanart
P-» e«d« Lt JI JE«,
My^tnoeeittiU'S
My, !IIO(.'<MIIHC,
(r)jp C
(P) long w- eios MS
(PJ eorollatu s
(PJ o c u l a l u s
(P) m e x i k o l p o s
(P) b o r c e a i
(PJ utticus
(PJ m « d i t e r r a n e u s
(P) i n f l a t u s
(PJsp.A
(P) v u l s e l l a
(P) f a l c i c u l a t a
ieiialui
: p
A
P% r u d o l a i t a i * ( P J ^ b f ^ ^ i
P* p u d o t a i i a i s (PJ b at**'* nauM
Ps « w d » l a n a « (P) calrfwrniensis
Pse<*d«»tatia«s (PJ for«ipatu&
Pseudotanais (PJjonesi
P s e u d o t a n a i s (P) eolonus
P s e u d o t a n a i s (PJ l i i l j e b o r g i
P s e u d o t a n a i s (PJ n o d e n s k i o l d i i
r?n!WTi ,|P—P a t a p s e u d otanais a b y s s a l i s
IfegBBiparapseudotanais s p . A
P s e u d o t a n a i s (A) longgipes
Pstssidotanais ( A J s i f c g i
Ps «Md«ta nais (A) isi aksoth six
P s t t u d o t a n a r t (A) m « i t e f t ? e » i
P', t-5id» Lt n^as (A) g « i l a e h i
P % * u d a M » a i s (A) g j i n s i
P,s * u d o lanai<> (A) * mi ilis
P* *»<ido l a n a i t (A) m a l a y p i t * K
Ps eud«*l»n«««& ( A J & e t a p p i
PseMdwtanais. (A) g w i l l e i
C r y p t o c o p e abreviata
C iyt©copt>ides arcticus
Crytocopoides pacificus
Cryptocpoides s p . A
l u n g e n t r t a n a i s primitwus
Paraiungentitanais longidigitatus
MHsLalitan'ais b e k l e m i s h e w
Cu rticttelia esiHcss a
IT n s i n e lob us s t a n i u i d i
l i a iflt" l i a lap ax
S «p l i i « ! j JlillS
|>4ll*«".
t'. A «pt> jp>. 0 ude% p« o^ j» eclntfS:
Figure 40. Character 9 trace - antennuie, distal article length vs. width
226
Antenna length vs.
antennule length
r ] * #0 pwnwtnt (mtgth of
H H SOtol&Bparoafftof
• M > 108 percent of
Pseudotanais
Pseudotanais
Pseydotanais
Pseudotanais
t*s t ' u i i « i j n jis,
P i «Mitl«!j IIJI«,
(P)S|
( PP) )Ss| p . O
(P)sp. P
(P) affinis
(P) s c a i p e l l u m
(P) i u n g k p i i i u *
(P) tleitticui^ilu^
P,<r t t l l d o Li II ,11',
( P ) >, [>Sfc ,,IllI':,
Pv «Mid« tit INI>, (P) wsipj^i
P % <>tido la ii jti-, (!*) » i p p « n i c t i ^
P<> <>udttUnJti-i (P) m ^ c i o c h p l c *
P » « v i l « U n 4 i 5 (P) s p C
Ps«udi>tanais (P) longis etosus
Pseudotastais (P) eoroiiatus
P s e u d o t a n a i s (P) oeulatus
P s e u d o t a n a i s (P) ntexikoipos
P s e u d o t a n a i s (P) b o r e e a i
P s e u d o t a n a i s (P) unicus
P s e u d o t a n a i s (P) meditisrraneus
P s e u d o t a n a i s (P) inflatus
P s e u d o t a n a i s (P) s p. A
P s e u d a t a n a i v (P) wuS-vella
Ps « u d « t a n a i s (P) laleicul<ita
My% tno«entiMv t e t r j l u i
My * t i i o e e n l i u * *. p A
P t f u d n t j i u i ^ (P)«iby^^i
EP<, <Midol* M*I«% (P) b a I * * it * u 11
Ps *>Md«t*n«n& (P) cairtornHtnsis
P s e u d o t a n a i s (P) foreipatus
Pssudotanais ( P ) j o n e s i
P s e u d o t a n a i s (P) eoionus
P s e u d o t a n a i s (P) lillj«bcurgi
P s e u d o t a n a i s (P) n o d e n s k i o l d i i
Para p s e u d o t a n a i s abyss alis
Para p s e u d o t a n a i s s p . A
P s e u d o t a n a i s (A) l o n g i p e s
Pseudotanais ( A ) s i e g i
P s e u d o t a n a i s (A) en a k re Bt > ix
P s e u d o t a n a i s (A) m o i t e n s e n i
P *. t ' u t l o l j ti j n , (A) g«it«iclti
j —nP t r u d o ta n a i t (A) g j u ^ ^ i
P«> euriola nai^. (A) »> atay vn?, w.
Ps <>udotainai« (A)sci<npp>
P s e u d o t a n a i s (A) n u i l l e i
3DC ryptocope abreviata
mmmmmCijtOGQpoi&ss
a<rcticus
Crytocopoides pacsificos
Cryptoepoides s p. A
i u n g e n t i t a n a t s primrhws
P a r a i u n g e n t i t a n a i s l o n g id ig Status
Latitanais beklemishewi
C u rtiehefia e g r e s s a
Sirtelobus S t a n f o r d !
H a r g t ria rap ax
auTyphletanais p j i w n
DCilijIO j l p $ i t t i d « |*K»SpMtjlt»
Figure 41. Character 11 trace - antenna, length vs. antennule length
Antenna article 2 length
vs. article 3 length
Pseudotanats
Pseudotamais
Pseydotanais
Pseud#tanais
Pseudotanais
P*»ud«ta«ais
P» e i i d u t i nai*~
P*
P%
P%
J-—-QP*
lirTlPs
e t t d o l i HJIIS
VUdolailAI*
»udol«Mt*l<»
»Udotai»aiS
eudflfcllttai'Su
(P) spatula
( P ) s p. O
( P ) s p. P
( P ) affirtis
( P ) s cagpeliuim
( P ) l o n g e , pin us
( P ) d e u t i c ulafcu*
(P)
(P)
(P)
(P)
(P)
% picafu*.
KsSf^I^l
tt ipp ttlllCIIS
maciOChel**
S p. C
i n P s e u d o f a n a i l s (PJ l o n g i s e t o s u s
P s e u d n t a n a i s (P) eor&Iiatus
Psfiydstanais (P) oculatus
igPseudotartais (P) mexikolpos
Pseudotanais (P) borceai
Pseudotanais (P) unieus
Pseudotanais (P) mediterraneus
Ps t>ud<ttaitais ( P ) i n f l a t e s
P s e u d u t a n a i s ( P ) & p. A
Pseudotanais (P) vulsella
P.S e u d o i a suit. ( P ) fjIcicuE^itii
My* I i i a c e n t i u * V e i i a t u *
M y t l l i o c c n t i u t •> p A
P* e u d o l a n a i * ( P ) aby*. <k i
P¥4»wd«1aiiais> ( P ) b a t * v n a u H
Ps e w d o f a i t a i s ( P ) «.alrfotns^nsis
Psftddstanais (P) forcipatus
Pseudotanais (P)jonesi
P s e u d o t a n a i s (P) colonws
Pseudotanais (P) lilljeborgi
Pseudotanais (P) nodenskioldii
. jpBBParapseudotanais abvssalis
nlczso P a r a p s e u d o t a n a i s s p . A
P s e u d o t a n a i s (A) l o n g i p e s
Pseudotanais (A)siegi
P s t f i i d u f a i W i ' , (A) isi afesttlltlsx
P v t u d o t a n a i f (A) m o r i e n t e n i
P s r u d o y i i d i t i (A) g e i i a c l t a
PN ewdola I M I ' I
(A)gaii<>ti
Ps »u4ol4lMi<k (A) s int i l i t
Ps » u d o l a n « m (A) lit a la y »its i t
Pseudola»ats (A)&Giappi
j u P s e u d o t a n a i s (A) g u i l l e i
C r y p t n e o p e ataiewiata
Crytaeopoides aretieus
C r f t o e o p o i d e s pacifisms
C i v p t o c p o i d e s s p. A
l u n g e n t H t a n a i s prim'rtvus
P a r a i u n g e n t i t a n ais l o n g i d i g i t a t u s
Latitanais beklemisttevi
Curtictielia express a
S n i t f i o b MS S t a n f o r d !
H a i g e i i a rap ax
fyp b i o t a l i j c j paiwu*
V. J <|M> j p . \ « Bile^ JHOi p<M:ilH",
Figure 42. Character 12 trace - antenna, article 2 length vs. article 3 length
-i-'PssudotsiMis (P) spaisiia
.L ""npsesidotaisais ( P ) s p . O
L u P i e u d o t a i i a i s (P) s p P
Antenna articles 2 & 3
i
r
• i
^. "ir, ^(simpleorspiniform)
x P s e a d a i a w a i s (P) dffmis
''—HP'S «sud«ta«ia<« (P) i cxlpaiUtm
__J uP% «-ii(l«Uiuli*»
. rJlZCP^ * u d o l * n j i %
^Ji .
r>P^ c u d o l j i i i i i *
~
riPs <Midol,ut4i'.
nP". »udotiin<m
Pseudstartais
Pseudotanais
Pseudotaitais
Pseudotanais
Pseudotanais
Pseudetanais
Pseudetanais
Pseudetanais
Pseudetanais
Psettdotaftak
(P) deniH.-uS.lfu>.,
(P)spicJlt^
(P) *ii|.ivi
(P) imppomcus
(P) <i p f;
(P) l o n g i s c t o s u s
(P) corollatus
(P) oeulatus
(P) mexikolpos
(P) borceai
(P)unicus
(P) medrterraneus
(P) inflates
(P)sp.A
(P) v u k e H a
}'*, *Miti«r;iii»i!», ( F ) S j i c i t i i l J i J
My, !not;eiil(ti>. '. c i u S u ' ,
My, i«itM;tM»li!c, ', p A
™nP«, f iiiioLiiioii', (P) b d i p ' , is-ifih
Ps oiido l^wijis (P) «*lrlomii* "MS
Pseudstsinais (P) fftfcipatus
Pseudotanais ( P ) j o n e s i
« P s e u d o t a n a i s (P) colonus
Pseudotanais (P) l i l l j e b o r g i
U P s e u d o i a n a i s (P) n o d e n s k i o l d i i
Para pseudotanais abyssaiis
Para pseudetanais s p . A
Pseudotanais (A) larigipes
Pseudotanais ( A ) s i e g i
P*. f u r f « l j i i i i i » , (A) maiauUtux
V% ifUiUilaa.it:
P', «M! J « Lit! JC,
P>, <MI<io LlISJB',
P", <*<nl«y I I J I ' .
P', S-iKfoUllitl',
(A)
(A)
(A)
(A)
(ft)
nioaipii-, t-tsl
gttiijCIII
jj,!<!<,':, E
•, is:iiis>,
!!» ^ijJiPlll', 1$
P & e u d o l a n a i * (A) s ciapp i
Psei<detsit<ai!> (A) fluiiiei
Crfptocepe abrewiata
Crytocopoides arcticus
Cryiocopoides pacificus
Crfptoepoides s p. A
lurtgerstitanais primitwis
Paraiungentitanais longidigitatus
Latltanais b e k k m i s h e v i
Curtieltelia o f t e s s a
Sinelobtis Stanford!
IV)>l>ioLiii,ia', p j i v t i * .
Figure 43. Character 13 trace - antenna ,articles 2 and 3 setal type
229
Antenna article 4
length x width
i
t <4x
CD 4.1 to 0,f M longwr
$ to 8 x longer than
8.1 to 10.2x loriflef
> 10.2 x longer than
P s e u d o t a nais (PJ s p a t u i a
P s e u d o t a n a i s ( P ) s p. O
P s e u d o t a n a i s ( P ) s p. P
m S L G i m o P s e u d o t a f t a i s (P) affinis
( r ^ u B i i i P & * t i 4 l « t a i n d i s tP) s calpeEium
P-s e u d o t a n a i f <[P) t u n y i i , piiunis
!•'•> tMs<fo!jii».n', ( P ) d e n l i i - u l a t t i S
P •. « u d o Ls it J I S ( P ) •. pit: ,ii»i'.
P *> (Midota n<n*> ( P ) w f i p j / j
P «, <* ii«! o I J I » J I «; ( P ) ii IJ»|> <ntn:uv
P\fHd«Unjin
P* e u d o t j i n ^ i i
Pseudofanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Ps eudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseodotartais
P\ e u d o l j i i j i i ^
(P)
(P)
(P)
(P)
(P)
(P)
(P)
(P)
(P)
(P)
(P)
(P)
(P)
«»4»fi«ttchtl«S
^ p. 0
Song»^ *>tos»is
corollatus
oculatus
iriexikolpos
borceai
urticus
mediterraneus
inflatus
s p A
vuf« etla
laieiculata
M y , l i t t t c e jilMfs »« p A
P', <*mi« I j !&,m, {$*} Jlhy:, *, i
V% «Midof<«»ai5 ( P ) b a i e M i a U f i
P>i v i i d o l . * n<ii^ ( P ) c a l r i o i m e n s i s
Ps»eudotAii<iis ( P ) f o r c - i p a t u s
Pseudotanais (P)|ouesi
Pseudotanais (P) colonus
P s e u d o t a n a i s (P) liifjeborgi
P s e u d o t a n a i s (P) n o d e n s k i o l d i i
P araps eudotanais abyss alls
Parapseudotanais sp. A
P s e u d o t a n a i s (A) i o n g i p e s
Pseudotanais (A)siejji
Ps e u d o t a naiv (A) frtakiwfh tix
P<> e u d o i a n a H ( A ) i n o i t e i i f e lit
Pit «?id« Li A JI'< ( A ) ^ •uliM.'Iss
P t e u d o l a i t a i s ( A ) 8 aiis •• I
P •. <»inio l j ii JI>, ( A ) >> isiiHr.
P •, 4*!HfOl,l )»JS', ( A ) 5!»-li-l¥<>If, l».
P >, *Mnin l j i» J I S ( A ) $ « M p p i
P s e u d o t a n a i s (A) g u i i i e i
30 C t y p t o c o p e a b r e i f i a t a i
l^SDCrytooopoides aroticus
rp^s^C r f t o c o p o i d e s p a c i f i c u s
1—oCi'Tp-ptocpoides s p . A
iungentrlanais primitvus
P a r a i u n g e n t i t a i s a i s Ion gid i g i t a t u s
L a t i t a t ) a is b e k l e m i s h e w i
CurtioheSia e g r e s s a
3 S i n e l o b us S t a n f o r d !
l ±= > n H a F g e r i j i t a p a x
iDfyp biota M i s
putrms
3 CarpttJ.iistud.ts p»*spt«tnt*
Figure 44. Character 14 trace - antenna, article 4 length vs. width
Antenna article 5
length vs. width
t i * 3,0 X longer than bmad
WW 4 to 8*0 x longer than broad
* * > @ x longer thart broad
Pseudotanais
Pseudota nais
Ps e « d » t a rials
Pseudtttamais
(P)s patula
(P)sp. O
( P ) s p. P
(P)
P s «imi«iai«.:»ai ( P )
P«#UdotanJl£ (P)
P * <Mitfu L i ilea's ( P )
P'i *M(«l«5j SiJf, ( P )
a If in is
s ealpellum
longtg plAU*
deitticiilatu*
% psc,iS«*,
P1, e i i d o f a i i i t i s ( P )
vitja,n
P>, < M I < i » l £ tt J ! % ( P ) i t i p p o m c u i i
P <>i (* »j d <* i a «»<m ( P ) i n » i * ; t « e h * » l p 5 >
Pseudotanais
try * p. c
Pseudotanais
( P ) I o n gits e t o s u s
Pseudotanais
Pseudotanais
(P) coroSlatus
(P)oeulatus
Pseudotanais
(P)
Pseudotanais
Pseudotanais
(P) borceai
(P) urtieus
Pseudotanais
(P)
mediterraneus
Pseudotanais
(P)
inflalus
mexikolpos
P s e u d o t a n a i s ( P ) S |»- A
P«.fe«d<*t<i n a i s ( P ) v u l s « l l a
P ' i t ' u d o i j is^s'i ( P )
ialciculata
M y * i n n c i ; n t i t t ^ 'i * IS j f l l ' i
M f . 1JS«:M:<* » t i u « t \ p
A
P". < M I d o t a l » , i r . ( P ) <aby% * •
P \ * it d o i i i n ^ I * ( P ) S» -it<*t, t t j i u t i
P<s*»udoian.ais
(P)
calrforniensis
Pseudotanais
(PJ fotcipatus
s^ s ss $1 u o R»3I as a i s ( P ) J o n e s i
Pseudotanais
Pseudotanais
Pseudota
(P)
(P)
nais (P)
eolonus
lilljeborgi
nodenskioldii
abyssaiis
P a r a p s e u d o t a nais
P a r a p s e u d o t a nais s p . A
Pseudotanais
( A ) 1 o it g i p e s
Pseudotanais
(A)s
iegi
P s e u d o t a n a i s (A) mak-rotfirix
P « e u d « t a n a i * (A) in « i l e n v « n i
P ' , e u d o t i i at J!«. ( A ) a. e i l a e h i
P "s «• ! i d o i^i n j • ^ ( A ) ^ -«!«•. •» I
P'i. * u d o l a n a i < » ( A ) % i m i l i *
P«, 4»«id«L*n.«i<» ( A ) i n -«l-i y <>««;, »',
Ps e u d o i i n j i s (A) s c i a p p i
P s e u d o t a n a i s (A) nutilei
Crfptocwpe
abreviata
aretieus
Crftoeopoides
Crytocopoides pacificus
C r y p t o c p o i d e s s p. A
lurtgentlanais primHwus
P a r a i n n § e miliars a i s l o n g i d i i g i i t a t u s
Latitanais
Curfiehelia
bekiernishewi
eifressa
Sin«Iobus stanfordi
I I J : 0 « i i J s j } > J>:
1 ^I> I i i o S j rt J»<» }* «i I win'.
(' »* S j><* .Ij»> V U dt f ,
Figure 45. Character 15 trace - antenna, article 5 length vs. width
j»« <>•, p « M : I I M ? " >
231
Mandible pars mularia width
P s e u d o t a n a i s (P)spatuSa
P s e u d o t a n a i s (P) s p. O
P s e u d o t a n a i s (P) s p. P
P s e u d o t a n a i s (P) afrinis
P s e u d o t a n a i s (P) s c j l p e l l u m
Ps#ud<ttan*rv <P) lunge; put us
P^ t u d a t l f t l i f (P) il*nlicul4lu%
P i *««l»!*iitjm (P) * pr«.*lu%
P •, <*«idu Li iui<, (P) vrtfiiJti
P•, «»«doijitJI<,
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
—rnaPseudotanais
I —ip s e U 4 j 0 tastais
1*5 eutluLarkaiS
P«, «Miii«tfciit JSV
Ufi,
i l i o c e ItlluK
(P)
(P)
(P)
(P)
(P)
(P)
(P)
(P)
(P)
(P)
(P)
(P)
(P)
miicrovhvl^jt
sp.t
long is d » s u s
eorollatus
oculafus
mexikoipos
borceai
unicus
mediterraneus
inflatus
s p. A
* « is fella
IdlciculaUi
', 4 * ! J j f « ' ,
M y . f !•*»<:<» itliit', ', ft A
f* |. e-«»lo ! J i>4i!', ( P ) ^ b f , M
P$ e u d o t a n j i * (P) b*i«>^ nJMH
P»«udotamir$ (P> ojilrii»iiii«nists
P s e u d o t a n a i s (P) foreipatus
Pseudotanais ( P ) j o n e s i
P s e u d o t a n a i s (P) colonus
P s e u d o t a n a i s (P) l i l l j e b o r g j
P s e u d o t a n a i s (P) n o d e n s k i o l d i i
Para p s e u d o t a n a i s abyss aiis
Para p s e u d o t a n a i s s p . A
P s e u d o t a n a i s (A) l o n g i p e s
P s e u d o t a n a i s (A)si*sj}i
P s a u d o t a n a i s (A) malcrttlhrix
F-, *«nl« Li Si j s s (A) m art «• IW * n i
!*>, *'«<!<> Li it JS«,
(A)g«iljchi
P t, <Mnl«t I J r»iia«, (A) gjui*.«, •
P s ^ u d « y i t J H (A) A innlit
Ps <>-u«l«i«in.ai<v (ft) m*it«i)p*r»s «s
Ps eudotaitJis ( A ) s e t a p p i
P$«vd«fainJii5 (A) g u i l l e i
Cr^ptocope abreviata
n C r f t o c o p o i d e s ardticus
.rr=nCrytocopoides pacificus
"1 uCryptocpoides s p, A
^ r p z o l u n g e n t i t a n a i s primitOTis
_ i r 1'—nParaiungentiianais longidigttatus
•fi
"ti Latitat! a is b e k i e m i s h e ¥ i
DC«rtiehelia express a
,. ~Ti£in*l4»bus s t a n f a r d i
""Lrzollargt n» tap am
1
u typhi a i m * is PMIWS
ci;j»fp® jtpstwdts piMSpt«tfi*s
Figure 46. Character 17 trace - mandible, pars molaris width
232
Mandible pars molaris complexity
trtoffiff
•hwttfMltgiM
M l m •#(«««:* pfttema
wm mmbiuup
Pseudotanais
Pseudotanais
Pseudotanais
Pseud»ta«tais
Pstmdotanais
(P)
(P)
(P)
(P)
(P)
s patula
sp. O
s p. P
affmis
s calpeltum
P*h e u d o t d iui»» { ! ' ) loitgu» p m u %
P •, (Mid» Li ss«Ji'i ( P ) •, ( n t j l i n
P', <* (id O S J »1 .1!'. ( P ) "flijJ-EI
P', ( ' I M S O U t t j l ' , ( P ) itSj>p«[iS<;U>,
P% rudotelams (P) HIHICKM: is <'!<>•»
PseMdsisna)*
Pstudotartais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
(P) longK etos us
(P) coroliatus
(P) oculatus
(P) mexikoipos
(P) borceai
(P) unicus
(P) medrlerianeus
(P) inflates
(P)sp.A
P<, «MS<i<»Ll «UiS:
P'»«•«<!« Ll HJi'i
M|f!,!iiin'f iiliu',
M y . !(!«<;<'illni'.
(P) VuKella
(P) Ijicieuljtta
'. f f f [ j f t i ' >
•» p A
P', I M M I O L H M I ' . ( P )
Jfcf.u
P^ 4»udoia n*fB% ( P ) b;ti<»^ struts
Ps*>Md»1Un^(s (P) calilomi«n£i5
Pseudotanais (P) torcipatus
Pseudotanais ( P ) j o n e s i
Pseudotanais (P) colonus
Pseudotanais (P) lilljeborgi
Pseudotanais (P) nodenskioldii
Parapseudotanais abyssalis
Para pseudotanais s p . A
Pseudotanais (A) iongipes
Pseudtfta «ais ( A ) s k g i
P.', eutiuSji'tj»<, (A) iss^fciolli fix.
P>. «-f(«i»8ii j t j r . (A) in«ileiii eni
P'»«'«*!« t i 11 j r .
P<, <M!*!o Li n,«!«,
P«. <MS«t« Li n,u!.
Ps (Hid«i,i(i»ii'.
( A ) fl r i l . n ; hi
( A ) flj««,«. i
( A ) •• ii<t tir.
(A) n u U f c n ; K
P s r u d n M n i K (A)sc»>?pp>
Ps«»«dofci raais (A) guiifei
Crjptoeope abreviata
Crytocopoides arcticus
Crytocopoides pacificus
Ctyptocpoides s p . A
iungentitanats: primrtwus
P ara iun gentitan ais ton gid igitatus
Latitanais bekSemishewi
Curtieheiia esepress a
Sine lob us startfordi
Harg^ria tapaK
i?}> it!<>i.i i>,ii•. parws
CjMpWXp$feU4l«$
Figure 47. Character 18 trace - mandible, pars molaris complexity
ptO£p«et»*$
233
MaxiMuIe endite
terminal setation
d 3 •*« sphifomr s e a *
E S S Mirip?4ini5ii.-nj---<t'*i3
I H •iQMsj.nrfcrnsel**
i n *_ intu'm-i *, ?b-f
P s e u d o t a n a i s (P) s p a t u l a
Pseudotastais (P) s p. O
P s e u d o t a n a i s (P) s p . P
P s e u d o t a n a i s ( P ) affinis
Ps#ii<l»tanais (P) s ealp#llum
P s e u d o t a n a i v ( P ) Sony is pinuv
P4 e u d o t a »iS»», ( P ) tl eikh*:u!alts',
Ps e n d o la nai^ ( P ) >, jm: „il»s-,
P •, «*«<J» Li n.f i<, ( P ) *«l|»t^.j
P », <MH!«> M si-is*. ( P ) it ipp itnitii',
P •; eu«i« Li IUIV ( P ) in J C I O ch^lim
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
P^eudotiiiiai^
Ps e u d o l a n a i i
Myt !ri«me ulns',
My>tfnac« nlm*
(P) s p . C
( P ) longis e t o s u s
(P) corollatus
(P) oculatus
(P) mexikolpos
(P) boreeai
(P) u n i e u s
(P) medrterraneus
( P ) inflatus
( P ) s p. A
$P) vitH el la
( P ) iaicieul j t J
', f i i j t « ! i
*. p A
P % <*urfa ta nai-. ( P ) b a««KV naiili
P s e u d o f a n a i * ( P ) calrfpimeit$i$
P s e u d o t a n a i s ( P ) forcipatws
Pseudotanais (P)jonesi
P s e u d o t a n a i s ( P ) ©olonus
P s e u d o t a n a i s ( P ) lilljeborgi
P s e u d o t a n a i s (P) nodenskioldii
Parapseudotanais abyssalis
Para pseudotanais s p . A
P s e u d o t a n a i s (A) i o n g i p e s
P s e u d o t a n a i s (A)sieg.i
P<s eisdota naif (A) m a k t o t h u ' *
P & e u d o t a n a i t (A) m o i t e n ^ e m
P >, iMidolj naiv (A) a. «il jcSti
P \ f i i d « la Sidis ( A ) ( | J i r , - , s
P •,<*!!rfO 1,1 !k ^ J «, ( A ) <• I Jli I II'i
P ", « ' ! l d » M J'«^I>, ( A ) !ll J I j f P D S |i,
(Bq
P*. «urio ta nai& (A) *» ci-«pp»
P s e u d o t a n a i s (A) auiIIei
Cryptoeope abreviata
Crytocopoides areticus
C r y t o e o p o i d e s paoificus
Cryptoepoides s p. A
l u n g e n t f t a n a i s printitvus
p a r a i u n g e n t i t a n a i s Iongidig Status
Lalitanais beklemisttevi
Curtiehelia e g r e s s a
i^PBSinelobus stanfoidi
TtzxiHa»«eria t a p a x
fyp hiota nai% p<n¥«v
BIT .I»|M> ,l[»'. V U l I t t
plot p e d n * *
Figure 48. Character 21 trace - maxillule, number of endite terminal spiniform setae
Maxilliped basis
degree of fusion
1.._.! run pie hsf* sap wz*&
Pseudotanais
(P)spatula
Pseudotanais
Pseudotanais
Pseudotanais
Pseudlotanajs
(P)
(P)
(P)
(P)
sgt. O
s p. P
affinis
n- ealpeilum
P^^Udttt^iiaiS
(P) lottjjts pirms
P ' i e i i d o L i i i j i v (P) denticulafu*
8*'»t?ss«i« Li «t,se', ( P ) % pi«:jUi>,
P * » I I d O l»t It <l • ^ ( P ) w»tjj|/i
P *, «-udo la «»*««*> ( P ) (s i p j M n i s c w ,
P i » u d « la n * * * ( P ) I M , I « I O « h * I * * s
P^^uilafan^ij (P)sp C
Pseudotamais
Pssudotanais
Pseydotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
P-, euiiitLi nais
(P)
(P)
(P)
(P)
(P)
(P)
(P)
(P)
(P)
(P)
l o n g i * «*tos«is
GoroIIatus
oculatus
mexikolpos
borceai
unicus
medrterraneus
inflates
« p. A
wulselldi
P'. «-4i<iuSit iljtii'i ( P ) l a l e i « w l j l ^ l
M f < I B S « « ; » BSIMS\
NeliJltus
M y v l i i o c e n l i i K •» J* A
P •« «*««lo lii i t ^ i s ( P ) j b y n ' t i
P\ (MidoiaitJ!^ ( [ ' l l t J H " , ! ! , ! ! ! *
P %. e-snHwta via**, ( P J <;j»lrf«ntir»i« »*s. i s
Pseiidotsnais (P) f o n j i p a t w
Pseydotanais (P)Jonesi
Pseydotanais (P) colours
Pseudotanais (P) iiiijeborgi
Pseudotanais (P) nodenskioSdii
Para pseudotanais abyss alis
Para pseudotanais s p . A
Pseydotanais (A) fongipes
Pseudotanais (A)s»eoJ
P*f e-tidotanai^ (A) niaktfothiix
t*i«u<lot«nai« (A) m«rt«ft« e i>i
P". « > « < l o L t l i , l f i
( A ) fl «-il,i<i l i i
P '• »•• !!«{ « L i Jk J S >, ( A ) fl ;« «S <, 'J !
P <« « M l d « * ( U » a ( " i ( A ) t t i l l l i l t
P»» « m t J o i f i n ^ i i
Pivewdotanais
(A)maUfM)^|i.
(A) 5 c r a p p i
Pseudotanais (A) fl«iHci
Cryptoeope abreviata
^"^nCrytoeopoides aicticys
Crytoeopoides paeiffieus
Cryptocpoides s p. A
luitgetititanais primrbras
P a ra iu n g e ntita r» a is iongidigitatus
Laiitanais beklemishevi
C ai rtic h e I ia express a
^ ^ ^ J c n o S i n e t o b u s Stanford!
Tfpltlotafiitis parvus
Figure 49. Character 23 trace - maxilliped, basis degree of fusion
235
MaxtiHped endite degree
of fusion
C 3
completely;
1/3-2/3 length
SPseudotanais
]pPse«d©tamais
Pse«d#tanais
.oPseudofanais
•pP* e u d o t a n a i s
:*™rr."™nP^et**iota«tia»<s
j tjPiiruilotiii^it
,'p t ^ n P ^ , e a i l i t l j i i j i v
sPteudobinjit
j-^aQPs f m d o t a n a i *
in f s t*i«d« fainars
Pse«d»t»nais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pvcudotanait
MytlllOCeitllUt
( P ) s p atula
(P) s p. O
(P) s p. P
( F ) affinis
(P) s caipeliMm
(P) Ioftgi& pij»u;>
(P) denliculafai v
(P) \ g>i<:,iBu-.
(P)
(P)
(P)
(P)
(P)
(P)
(F)
(P)
(P)
(P)
(P)
(P)
(P)
(P)
nippaniew*
mAciachvle.t
& p It
long is etosws
corollatus
ocuiatus
mexikolpos
boreeai
unions
medtterraneus
inflatus
s p. A
vulsella
lalciculata
',4»IIjili<,
Ps e u d o t a i t j ^ ( P ) a b y t s i
Ps> eudota »<••% (P) b A t * * t»*i»i b
Ps eudwtanaix (P) c<drtorni«nsi5
P s e u d o t a n a i s (P) f o r c i p a t u s
Pseudotanais ( P ) j o n e s i
P s e u d o t a n a i s (P) colonus
P s e u d o t a n a i s (P) l i l l j e b o r g i
P s e u d o t a n a i s (P) nodenskioldis
rpza Para p s e u d o t a n a i s abyss alis
li_^Li Para p s e u d o t a n a i s sp„ A
P s e u d o t a n a i s (A) l o n g i p e s
Pseudotanais (A)siejji
P&feudotanais (A) m a k i o t h l i x
P« eudo latitat, (A) l i i o i l i M n « i>>
Pv e u d o t a n a i t ( A ) a ? i l 4 c i H
P^ « - u d » l * i i u n (A)fl»«««.',E
P ' , < * ! ! 4 S 0 l , l t* J J ' i
f A ) >, HIS «l8V
P<« ciido ta nJis (A) ma lay*n% p>
P* «Md<»ta«4»5 (A) 5. er^r»p«
P s e u d o t a n # i » (A) g«IHei
U D C r f p t o c o p e abreviata
"OCrftocsopoides arcticus
1=13 Cryto cop sides paeificus
1 u C r y p t o e p o i d e s s p. A
_ j — D iungentrtanais pilmrrwus
_ | t ,n P a r a i u n g e n t i t a n a i s l o n g i d i g i t a t u s
'• *- _;u L a t i t a n a i s b e k l e m i s h e m
D Curtierselia express a
S3 Sine tab us S t a n f o r d !
Ha riper! a rapax
oTyphletanais parws
C a rp a a ps « u d e$ p i os p « et« *ss
Figure 50. Character 24 trace - maxilliped, endite degree of fusion
iVlaxiliiped endtte setaticn
d ) ta«3( jnosesets*
r j«.;i'. aitr- >*,»*w
St«laif tfjlhi
F s & d d o t a n a i s (P) s p a t u l a
Psewdetartais (P) s p. O
Pseudotaraais (P) s p„ P
P s e u d e t a n a i s (P) aifiRis
p 5 « « d # t a wais (P) s e a i p e l l u m
P ? e u d o t a n a i ¥ (P) lungfe |>i!itj',
P». <* u <I« Li «i .1 i •» ( I f ) <l e it tic <«i j l w ,
P s e-«<i« 1»« is J! 1 , (I") viifjjn
P', r « d « i j i i * i f , (P) itipp omens
P«, <M«<i<>t<l «,i»«. (P) iii,i*:i« nj»**l«"i
Pse-Kdotanais (P) x p. V.
P s e w d o M o a i s (P) iotigts eto*u?
P s e u d o t a n a i s (P) corollatus
P s e u d n t a n a i s (P) osulatus
P s e u d o t a n a i s (P) m e x i k o l p o s
P s e u d o t a n a i s (P) b o r e e a i
P s e u d o t a n a i s (P) unicus
P s e u d o t a n a i s (P) m e d i t e r r a n e u s
P s e u d o t a n a i s (P) i n f l a t u s
P s e u d o t a n a i s (P) s p.. A
P?ttudotnnai? (P) *uls«;lta
P V«MJ»I<> 1,111.1!', ( P ) litStH-ttS J t j
Ps r a d o y i i . i : ' , ( P ) - i itif, *. i
P*. <nsd« M n-i!'. (P) b j m n n j u t i
Ps>eMflaMnj(i& (P) c * l r f o r n i « n s i s
P s e i i d o t a i t a i s (P) torcipatw*:
Pseudotanais ( P ) j o n e s i
P s e u d o t a n a i s (P) colonus
P s e u d o t a n a i s (P) l i l l j e b o r g i
Pseudotanais: (P) n o d e n s k i o l d i i
Para p s e u d o t a n a i s a b y s s a l i s
Para p s e u d o t a n a i s s p . A
P s e u d o t a n a i s (A) i o n g i p e s
PseudotaitJiis ( A J s i e g i
P s e u d o t a n a i s (A) m a k r o f t i r i x
P s t i t d o t a n a i s (A) m 0 r t * n s * n i
P s m d a t a i s d i s (A) g « r » « h i
P s t u d o f t a n a i s (A) g a u s s i
Psfm*«toii*Ss ( A ) i i m l l i *
P s e u d o t a n a i s (A) ma lay*ns is
Pseudotanais: (A) s «rappi
P s e u d o t a i i i i i s (A) g u i l i e i
Crypto«»pe abiewiata
C ryto c o p * i d es a f d i c u s
Crytocopoides paoificus
Cryptocpoides s p. A
lungentitanate primitvus
Paraiungentitanais longidigitatus
L a t i t a n a i s b e k l e m i s heiri
Curtichelia e g r e s s a
Bljl
Batg«tia U|»JX
Typ it i <*to M »$ p M rums
C a ip# *ps * ud«« pi»« p *«tn«s"
Figure 51. Character 25 trace - maxilliped, endite setation
Degree of cheliped
forcipation
C D 5trcrtcv^t£3«s
•*'ii«"i» f '.'t"irtiji. , jit*»
rtotfaicoate
J
PseudotanaJs
Pseudotamais
Pseudetanais
Pseudoianais
Pseudotanais
PveudotanaiV
p-t e u d o t a i i a i *
P«s * u d o L i n j i i %
P-» e u d o t a n a i t
P* HMidatanaii
Ps e u d f t t a n ^ i ^
Pseudotanais.
Pseud^fanais
Pseudotanais
Pseudotanais
pPseudotanais
Pseudatanais
Pseudtttarials
Pseudotanais
Pse-jdmtat&ais
Pseudotanais
n_bmz:DPs*ud«laiiais
|!
:nP* e u d o t a n a i s
My* I r i o e e n t m v
(P) s patula
( P ) s p. O
( P ) s p. P
( P ) affinis
(P) s eaipetium
( P ) i o i t g t£ pinitA
(P) de»ti«uljlu-*
( P ) *> p i c a t u s
( P ) vil|a.a
( P ) tk ipp o m e n *
(P) maeimehele^
( P ) * p. <?
( P ) l o n g is e t o s u s
(P) eorollatus
(P) oculatus
( P ) m exile o l p o s
(P) borceai
(PJ a n i e n s
( P ) m e d iter ran ens
( P ) inflatais
( P ) s p. A
(P) vulsella
(P) talciculala
* e M ^ilni*.
My<i IISOCP ratine '. p A
"Zr7"pP<t o u d o t a n a i * ( P ) a b ^ t i
Ps e u d o t a n a i * ( P ) b J M * S n a i i t i
P s e u d o t a n a i s ( P ) ijalrtoriiiieiiisis
(P) forcipatus
nJlZD P s e II d ofa
. n P s e u d o t a n a i s ( P ) Jones i
Pseudatanais (P) eolonus
P s e u d o t a n a i s ( P ) lill|«b*»f«»
Pseudotanais (P) nodenskloldii
P a r a p s e u d o t a n a i s abyssaSis
Para pseudotanais s p . A
P s e u d o t a n a i s (A) l o n g i p e s
Pseudotanais (A)siegi
P f e u d o t a n a i s (A) m a k r u t t i i i x
Ps* e u d o t a nais (A) A i o i t e i t s * mi
Ps v u d t t t t i t a i s ( A ) g e r l a c h i
Ps e u d o t a n a i * (A) g a u s s 1
P«i r u d o t a n a i s ( A ) % i m i l i t
Ps e u d o t a n a i s ( A ) m a l a y e n v i t
P s e u d o t a n a i ' " . ( A ) * r.rappi
^ P s e u d o t a n a i s (A) g u i l l e i
Cryptoeope abi€iriala
Crytoeopoides areticus
Crytoeopoides pacificus
C r y p t o c p o i d e s s p. A
l u n g e n t i t a n a i s pritnitwus
Paraiungentitanais longidigiiatus
L a t i t a r t a i s b e k l c m i s henri
C u r t i e h e l i a esrpress a
S i n e lob us S t a n f o r d !
H a i g e f i a rap a x
Tfp It i o tan a is p j r w u #
C JiipWiiipsf n d t f pr<t*f i»««*f*«i
Figure 52. Character 26 trace - cheliped, degree of forcipation
238
Cheliped carpus
length vs. width
l~J
! •
• I
t—y
< 2 x longer than broad
2 to 3.9 x longer than broad
> 4 x longer than broad
eQuSvocaii
Pseudetanais
Ps e u d o t a n a i s
Pseudetanais
Pseydetanais
PseudoLjnais
(P) s patula
( P ) s p. O
( P ) s p. P
( P ) a if in is
( P ) $ ftalpfctlwim
' P « tMKlo tiS (Wits ( P ) i o i t y i S pift!l<>
P s e i i i i o L H u i ' , ( P ) <d etsln: u l ^ i u s
titni»Ltti*is<:,
nt* * f i i d a Li HJII:
bPifuddUnJik
P<« «>iid»ijii^ii',
P s eu d a t * n a is
—DPS e w d o t a n a i s
trjuPseudotanais
rzJOPssudotanais
—oPseudotanais
psoPseudotanais
i^joPseudotanais
. _,._i€3Pseudotaitais
^j—zinPseudiotanais
I—"n
uPs*>ud«tanais
. —ZZ^DP^^udotauais
( P ) «• jut; j i t s ' ,
( P ) w si | »«.<•«
( P ) it ip p «ss i c u"i
( P ) m j c i n ch«|*i>
( P ) s p. C
( P ) ionflis; c t o s w s
(P) eoioliatus
(P) oculatus
(P) mexikolpos
(P) boreeai
(P) unicus
(P) mediterraneus
(PJ i n f l a t e s
( P ) s p. A
(P) vulsella
Mys t i i a e e n l i u s « * u . i t u *
Myv t t i o e e n l i w * % p A
f**, * » u d » l a IIJII* ( P ) j h y t , ' , i
D P •, «MBrf« S<l I* ^ I'4 ( P ) It ^IJ**', II J<! h
I, ,L)Ps?«idotait*is ( P ) «alrtoi»i<tn<MS
r^ n _ z z x i P s e u d « t a wais ( P ) f a r c p p atus
;
.nPsewdotanais ( P ) j © n e s i
r—M .
; u P s e « d K » t a n a i s ( P ) C=»IOI»MS
QPseudotanais (P) iilljeborgi
OPseudotanais (P) nodertskioldii
Para p s e u d e t a n a i s a b y s s a l i s
P a r a p s e u d ef.arsais s p . A
P s e u d e t a n a i s (A) l e t i g i p e s
Pseudetanais (AJsi^jji
P s f e u d o t a i t i i c (A) m a hero U n i x
Hs * * u d e t a i i a i « (A) m o i t # n s e n i
'•P •» <Msd« Li S8as'i ( A ) g e i l i f t C l i i
np;np* fn4oUniii*i
(A)g<iuN^i
s 4»ud«»Liin«m (A) % m u l l *
P<i < » u d o l 4 SiJs*, (A) m a lay*n<t K
P s * » w d e t » l » 3 » 5 (A) s « r * p p i
P s e u d f t l a n a i s (A) « u i II e i
HCryptocope abreviata
" n n c r f t o c o p o i d e s areticus
fp^nCrytocopoides paeificus
T—liUCrwptocpoides s p. A
fungentrtanais primitvus
P ara iun gerttita n a Is Eon 3 id i g i t a t u s
L a f i t a n a i s belt ie mis hewi
Curtiehelia express a
j~~aoSinel»b«s stanfardi
T_~io H a IS * ria rapjiM
iTyphletatiisis pjtrvy$
I iC «S f p ft M pS* * II4 <** p I *S p * «tft «
Figure 53. Character 27 trace - cheliped, carpus length vs. width
Cheiiped impm
inferior setae
L.I
munrO'JM'', V Ts5f«s
E 3 line
(•• Mi
Pseudetanais
Pseudotanats
Pseudotanais
Pseudotatsais
Pseudatanais
Pfeudolanai*
Pt eudutaiiai*
( P ) s patmla
(P)sp. O
( P ) s p. P
( P ) a if in is
(P) * c a l p e l l u m
(P ) S o n g ^ pin I K
( P ) <l •tulle siLthsv
P ' . t ' l i i l o l j i i j i 1 , ( ! ' ) '. ( i i c i h i i
(*', t ' l i i l n l j i i i j i ' ,
Ps e u d o l a nais
Pseudotanai*
Ps a u d i t a n a i s
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pset*d*>tanais
Ps feudolanais
P s c u d o t a riais
Ps
ttudttljnjK
MyS lllOCIMlllUS
(P )
(P)
(P)
(P)
(P)
(P)
(P)
(P )
(P )
(P)
(P)
(P )
(P)
(P )
n ippotiicus
«i a«sr» C!I*»I«K.
* p. I',
Songis e f o s y s
eorollatus
oculatus
mexikolpos
borceai
untie us
mediterrraneus
its f l a t u s
* p. A
vulvella
laifciculdia
teiljllll'j
My*, t n o c i ' n t i u * *> p A
_. . n P * ** M d o 1 A •) J i -i (P)<»by«isi
Ps «MMIO)-« Hjtis ( P ) It «m» s itiimh
Pseudofcanai* (P) c a l r f o m i ^ n s i s
Psfcudotunais. (P) torcipatu.*
Pseudotanais (P)jonesi
P s e u d o t a n a i s ( P ) coSonus
P s e u d o t a n a i s (P) i i l l j e b o r g i
P s e y d o t a n a i s (P) n o d e n s k i o l d i i
Parapseudotanais abyssaiis
Parapseudotanais sp. A
P s e u d o t a n a i s (A) i o n g i p e s
Pseudolanais ( A ) s i * g i
P***Udtfian.aiiis (A) m a If re 111 f i x
P* e u d o t a n a i * (A) t t i o i l « n « e n i
P ^ e u d n l j E U i ^ (A) « * i i . i c ! t !
P>< e i i d o t d ILIIS, (/t) f j u s ' . I
P', «MMJ o f <i H „i««, (A) % m i i l i s
Ps *MMJ«»1J n^n,
(A) m a l a f 4*n* tv
Ps * u d o f a n a i s (A) s c» a p p i
P s « u d « t a n a i s (A)
ftuillei
Crfptocope abreviata
Crytocopoides arclicus
Crytocopoidfcs paeifieus
C r y p t o c p o i d e s s p. A
l u n g e n i i t a n a t e printrtums
P a r a i u n g e n t i t a n a i s l o n g id ig Status
Latitaitais bekiemishevi
Curtiehelia express a
Siiiel#bus stafifordi
Hasfleis;t r a p a x
T y p h l e t m a i s pjtivus
C <MrpUJp$«ii4ts pra$p-««t»#s
Figure 54. Character 28 trace - cheiiped, carpus number inferior setae
240
Cheiiped propodus
length vs* width
r~""i up to 2 4 x longer
iiini 2.S*4»0 x
I K mora than 4,0 x
broad
Pseudotanais (P) spatula
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Ps *;wti(* Li ii^i'i
P i i-tsiluljndi'-,
(P) s p. O
(P)sp..P
(P) affimis
(P) s caipelium
(P) long is |)iiiuv
(P) d«-»iicutdlus
P', <:Mltl» ! j Bl.,111 ( P ) 4 piC j t u t
P*l «MI*it» L« tt JI«. ( P )
P i rudnijtstxi*,
P<s *Mid» S,m,i!s ( P )
P<i e u d o l a n a i s
VttfA'M
( P ) ss gpp o m e n s
mjci«chflff
( P ) "V p
C
Pseudotanjiis (P) Inngfe etwsus
Pseudotanais (P) corollatus
Pseudotanais (P) oculatus
Pseudotanais (P) mexikolpos
Pseudotanais (P) borceai
Pseudotanais (P) unicus
Pseudotanais (P) medrterraneus
Pseudotanais (P) inflates
Pseudotanais (P) s p. A
Pseudotanais (P) vulsella
Pseudotanais (P) falciculatai
My<, iiioceittiu', •, <?i[,t{»i«,
M y t lii<M : :*Miti«<,
<, jt
A
P', *M!dn Li JWII<> ( P ) >thf>> <, t
!* «, P U d o f i l U J I ' , ( P ) It JIM>% l i J M i l
Ps e v d » t * i t ; m (P) calrfomiensis
Psewdnlana** (P) forcip^tus
Pseudotanais ( P ) j o n e s i
asPseudotanais (P) eolonus
Pseudotanais (P) i i l i j e b o r g i
Pseudotanais (P) n o d e n s k i o l d i i
Para pseudotanais abyss alis
Para pseudotanais s p . A
Pseudotanais (A) loitgipes
Pseudotanais ( A ) s i e g i
Pseudotanais (A) m atciotfarsx
P>, t * « * i « S j t i ^ i - j
F i Miiiitljitji'.
(A) m o t l e i K t h i
(A) g « « I j « h i
P •. i>t»<ilf Li s» ja«, ( A ) g «iu% * •
P t t M i d o t a i i i m ( A ) <, im «Ii<;
P<i iMirfnljiHAit (A) tmalay*!*;. P»
P^ewdstan^is- (A) s erappi
Pseudotanais (A) g u i l l e i
H-XSCryptoeope abreviata
OCrytocopoides arcticus
Crytocopoides pacificus
SCryptocpotdes
s p. A
lung entitanais piimrhms
Paraiungentitanais longidigitatus
SOLatrtanais bekiemishevi
IQCurtiehelia express a
j"p3SSin«l«bus Stanford!
l i z 3 ] H a r j | t ria rapJix
'fptiltttjnais parvus
DCtrp»Mps*wd*s prosp*mtn«
Figure 55. Character 29 trace - cheiiped, propodus length vs. width
Chslipfd propodus
vs. mrpm length
P s e u d o t a n a i s (P) s patuia
P s e u d o t a n a i s (P) s p. O
P s e u d o t a n a i s (P) s p. P
P s e u d o t a n a i s (P) a f f i n s
Psfc«d«>ta«ti««s(P) s c a l p e l l u m
•
S U ttme» tengtti
Wk > 1,5 times length
P v e u d o t a i i a i l i ( P ) lOtigUt p i n i l *
P ^ «MH!«ijr !i»:ItS ( P ) d e t t t l C U i j l t i t ,
Ps e u d c L i n J K
P s e u d u L i !ui> a
r^fiidoUiuti.
P*, ( M l d o l j JJ,t!5
P«> e u d o t a satis
(P) \
ptvAla:
( P ) wtlfj^i
(I*) n ip j> t n i c n ' i
( P ) >H JCUMlil^if".
(P) s p. C
Pseudotanais ( P ) ! » n j B ? f « s « 5
P s e u d o t a n a i s (P) corollatus
P s e u d o t a n a i s (P) oswlatus
P s e u d o t a n a i s (P) m e x i k o l p o s
P s e u d o t a n a i s (P) b o r e e a i
P s e u d o t a n a i s (P) unicus
P s e u d o t a n a i s (P) m e d i t e i r a n e u s
P s e u d o t a n a i s (P) inflatus
Pseudetamais (P) s |> A
Ps*ud4»tAnai« (P) v u l s e l l a
?*««••<« do 1,1 it j«', (P) t a l c i c u l a l d
My* ttioeenlfu'i. ^ e i u t u t
Mys l i n t c f silKfi, •» j> A
P-, <*wdol..« ts,i5'i (!*) j b f , >, i
f* >• *»ud<> t-a i» j i ' . ( P ) b j n * i «i j ! l l i
Ps«?udoIanais (P) c a l i f m n i e n s i s
P s e u d o t a n a i s (P) f o i c i p a t u s
Pseudotanais ( P ) j o n e s i
P s e u d o t a n a i s (P) eolonus
P s e u d o t a n a i s (P) l i l i j e b o r g i
P s e u d o t a n a i s (P) n o d e n s k i o l d i i
Para p s e u d o t a n a i s a b y s s a l i s
Parapseudotanais s p . A
tZlOPseudotamais (A) i e n g i p e s
UbziPPsetidotanais ( A ) s i « g s
^ P s e u d o t a n a i s (A) m a if to 1ft i ix
P* eudulaiiai^; (A) inoit*»ir4 e i t i
'CPs. e u d o f c n d l s ,
P t eudoLtitJis
(A)
geilaclil
(A)JJ,HIV>I
P% < » « d « { j Ii.H', (ft) % 1!« llEl»>
P'• *nid«1J »*«i'i (A) m<iMy<>;r. i'.
Ps e n d o t a n a i * (A) s ««.*j>|>»
J-Psewdotanais (A) jj«»Slei
QCryptocope abreviata
JCrftocopoides a i c t i c u s
jrp^DCiftocopoides paeificus
T—iJCrfptncpoides s p. A
. . j y Z D l u m g e n t i t a n a i s primrtvtis
I 1 — n P a i a i u n g e n t i t a n a i s l o n g id io, Status
saLaiitanais b e k i e m i s h e v i
IDCurtieheiia e ' x j t t t s s i i
SjueiolMis S t a n f o r d !
;rt:i H j fyt.* n i t i.t}> j > :
~33t: a i p o j p t e u d e t
pf»sp««tl*tS
Figure 56. Character 30 trace - cheliped, propodus length vs. length of carpus
242
CheJipod fixed finger
number of inferior seta®
• • one lnT#i-!C" sM*
• •
,
:M! i!|ft' fiur*it;:ir
P5elJd©tatlai5
Psetidotanais
Pseudotasiais
Pseiidotaisais
P^euiiotiiiais
P', t'tstioLi s,;t»'i
(P)
(P)
(P)
(P)
(P)
(P)
s palula
s p. O
s p. P
affinis
* C4ip*>iium
l o t i g i s piiau*
P l , e i a d o t a stai^
I'«• tMnl o i»t si a i >,
P i «**nioi«i i u i > .
P «. P i i d o f i i SSJS%
( P ) d eiktic ailata^
(!-*)«. JIH: ^ t!!«i
( P ) vila.j.a
( P ) H i p p oatico*.
FstudofanJiis
Fseudotanais
Pseudotanais
Pseudotanais
Fseudotauais
Pseudotanais
Pseudotanais
Pseudotanais
Psemdotanais
Psetidolanais
Pseudotaatai?
I's *M(doLi iiiii'i
( P ) s p. C
( P ) Iwngis e t o s u s
(P) cnrollatus
(P) oeulatus
(P) mexikoipos
(P) b o r c e a i
(P) unicus
(P) medrterraneus
(P) i r t f l a t u s
(P)sp.A
( P ) «ruK elSa
(P) laleaculala
M y , t!!»M::e I S I K ! ' , •» «• BS j h s ^
M y * i i i f t c c <&«.»<»>• «, |> A
P •» •»(id«{-« n J«<. ( P ) b JI:«»S »sj«slt
P s o-iedolji ii-iii, ( P ) t; .itsittt smut", is
Ps«*udolassai5 ( P ) f o r c i p a t u s
Pseudotanais (P)Jonesi
Pseodotanais (P) colonus
P s e n d o t a n a i s (P) l i l l j e b o r g i
P s e u d o t a n a i s (P) n o d e n s k i o i d i i
j~=DParapseudotanais abyssalis
l i ;uParapseiid©tanaIs s p . A
P s e u d o t a n a i s (A) l o n g i p e s
Pseudotartais ( A ) s i e y i
Ps e u d o t a n a i s (A) m a k i o t t t else
Ps e u d o l a n a r * (A) fnoitt»ta£«tni
P •, «Mttio Li si j i ' i
P '« t»aado Li lijit',
P*<>udolaii*i%
P s *tit<{oi*i n J J S
P5«jwdol<M»j>$
(A)
(A)
(A)
(A)
(A)
g e i l a v ha
1J -m<. •» i
•, i!fi i!n.
in a lay#ai<» is
s <: J -i |> |* s
P s e w d « t a n a ? s (A) g u i l l e i
Cryptoeope abreviata
S C r y t o c o p o i d e s aretieus
_ ... . j r p ^ P C r y t o c o p o i d e s p a c i f i c u s
1 — - i i C i f p t o c p o i d e s s p. A
printitvus
i_JTra»Iungeiititattais
J| 1 ~H3 P ara iun g e n t i t a n a is I o n gid i g i t a t u s
^-—"•"•"'Latitanais b e k i e m i s t t e i r i
BCurtiehelia express a
S iin
n eel li «
t bbuus S t a n f o r d !
Haws'S
"TJbaDHaifjt jri*
.-li Yyp li I <tti it a Is pa r wti$
~DC 4 i i p < U p S (tM<t«:f: p*'«S |i*clll(f;'»
Figure 57. Character 31 trace - cheliped, fixed finger number inferior setae
Cheiiped dactyius
percent of propodus
Pseudotanais
Pseudotanais
Psetidotanats
Pseudotanais
(P)
(P)
(P)
(P)
s patula
s p. O
s p. P
affinis
! . 1 < 50 percent
H I 50 to S3 perewnf
ma > 63.1 percent
P.SMnl«falHlS
(P) « Caipftllum
P s e u d o t a n a i s (H) l o n g is pin uv
P v e u d v b i t t < i » ( P ) denffarulatut
P.1*. #udoLanai.t> ( P ) <v p i « 4 l u i
P* « u d » U n < i H (S*) *SIJ«I-PI
md
P% c u d o l a n a i N (I*) n ipp o n i c u *
P<t * u d o t a n a i s ( P ) ma«io«hel<»$
P s « M d « t a n a i s ( P ) s p. C
P s e u d o t a n a i s ( P ) l o n g is e t o s u s
P s e u d o t a n a i s (P) corellaius
Pseudotanais (P) oculatus
Pseudotanais (P) mexikolpos
Pseudotanais (P) boroeai
Pseudotanais (P) unieus
Pseudotanais (P) medtterraneus
P s e u d o t a n a i s (P) infiatus
P s e u d o t a n a i s ( P ) s p. A
Ps«udotanai& (P) vulsella
P« e i i d o t a n a r ^ ( p ) l a l c i « i i l a t i «
My4 l n o c e n1iuv «> <'ti *iS?f,
My% l u o c e n t H i 1 ! ». |* A
P', 4MJ ll «!..l «<>.« 3'. ( P ) j b f l SI
P5 e n d o t a n J15 ( P ) b a n i : n a u Si
P & « u d o t a n a i s ( P ) c a l r f o i n i o 1151s
P s e u d o t a n a i s (P) forsipatus
P s e u d o t a n a i s (P) j o n e s i
P s e u d o t a n a i s ( P ) colomus
P s e u d o t a n a i s (P) l i l l j e b o r g i
Pseudotanais (P) n o d e n s k i o l d i i
Parapseudotanais abyssalis
Para p s e u d o t a n a i s s p . A
P s e u d o t a n a i s (AJ l o n g s p e s
Pseudotanais (A)siegi
P s e u d o t a n a i s (A) m a k i o t t m x
P* # u d o t a n a i f (A) m u f l t t n ^ e t t i
Pa e u d o l a naiv (A) g e i l a c l m
f*"k t u d v t j n a i \ (A) HJ j?is % j
P t # M d o L * i i * i s (A)*inuli<>
r — O P 5 « u d o l a n a i s (A) m a l a y e n s «
•-• n P s e u d o t * n a i s (A) s s i a p p *
P s e u d o t a n a i s (A) g u i l l e i
Cryptoeope abreviata
Crytoeopoides areticus
Crytocopoides paeifieus
C r y p t o c p o i d e s s p. A
lungentltaitais primitvus
Paraiumg&ttiitaitais longidigitatus
L a f i t a n a i s beklemishe-wi
Curtiehelia express a
Sin globus s t a n f o r d i
Ifargetia rapax
iypiilotdi ssji!«. |»,isw<('«
t ^ t i p o j p s e u d e ^ oi 01 p o c l i i e *
Figure 58. Character 32 trace - cheiiped, dactylus percent of propodus length
244
Cheliped dactylus width
L... J is-€•"**» "Milft***dtngar
WKM firr * r « nivs Siuf! %£••??
tais (P) s patula
Pseudotanais
*ais
P s e u d o t a n<
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
P* «-udot»rt.a«s
(P)
(P)
(P)
(P)
(P)
(P)
s p. O
s p. P
affinis
s eatpeliMm
I o u t f i t pin «•>
dtsi* hew l a t a *
P% <*«nl» I d I M l t i ( P ) 4 piC j l l u t
P"» e u d o l d i i A i t ( P ) wtfj«i.£i
Pit ew4ot*n«t«"i
Psewdoyitais
Ps «u 4 a t a n * is
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseiidotanais
Pseudotanais
Myvtuocentiuv
(P) n i p p o n i c u v
(tP) mi <f«io«h*l«s
(P) s p. C
(P) longi? et»s«s
(P) coroflatu s
(P) o c u l a t u s
(P) m e x i k o l p o s
(P) b o r e e a i
(P) u n i c u s
(P) m ed rterraneus
(P) i n f l a t e s
(P) s p. A
(P) v u l s e l l a
(P) taleieul.at,a
v e i l alius
M y i t i i f t c e i i i u i " . «, jt
P s aMldo L« H J ! ! \ ( P )
A
jib^.M
Ps e.wdot*nais (P) fc«!•*•% nawfi
P&«u4otan«iis (P) c a l i f o t n i e n s i s
P s e u d o t a n a i s (P) f n t e i p a t u s
Pseudotanais ( P ) j o n e s i
P s e u d o t a n a i s (P) colonws
P s e u d o t a n a i s (P) l i l l j e b o r g i
P s e u d o t a n a i s (P) n o d e n s k i o l d i i
Para p s e u d o t a n a i s abyssais's
Para p s e u d o t a n a i s s p . A
P s e u d o t a n a i s (A) l o n g i p e s
Pseudotanais ( A ) s i e g i
P s e u d o t a n a i s (A) m a k i o M i r i x
P& wudolanais. (A) m a l i e i i c v i H
P s e u d o t a n a i s (A) <$ t? i Litis i
t* % eu d « l j »j8', (A) g .us>,». a
Pv eudo lanats (A) v i m i lis
Pv ttwilti U n d M (A) ns J Liyesi*. i>.
Pseudotanais (A^seiappi
P s e u d o t a n a i s (A) g u i l l c i
Cryptoeope a b i e v i a t a
C r f t o c o p o i d e s areticus
C rytoeopoides pacificws
C ryptwcpoides s p. A
l u n g e n t i t a n a i s primitvus
P a r a i u n g e n t i t a n a i s Ion gidig Status
IP! tmmm L a l i t a n a i s b e k l e m i s h e v i
iililliiUUl,l,«l«l,S«il»lisliUa,U,Bltt.l»lJ»..KMUSMSAl»
m mmmm C u r t i c h e l i a e x p i e s s JI
Sine lob us S t a n f o r d !
fflu
refill Iota waif pairmf
C « i p o * p $ « writs ptospe, ©tints
Figure 59. Character 33 trace - cheliped, dactylus width vs. fixed finger width
245
Chellped dactyius
proximal seta
• B afcser.
S 3 sep^s
JD
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
(P)
(F)
(P)
(P)
(P)
spatula
s p. O
s p. P
affinis
s eaipeiium
p-i e u d t t l a i i a i s ( P ) iOfty k piliM.S
P •» e u d o i d i t a ^ ( P ) d e n i i c u i jl?s i
F«, e i H l t > ! j i i . i i " , ( P ) •, [ m : j t « ' »
P « » « < l « t d l i . t l 1 r ( P ) WttjiJI/9
!*«. *MM:I*>5*1 llull'i ( P ) n i p i U H l I C H ' ,
Ps e « d « IJS n.ai'»
Pseudotanais
Psewriotarjais
Pseudotanass
Pseswdotartais
P s e u d o t a rials
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Ps e u d o L m a i s
PT. tMHioJjji.«»•;,
(P)
(P)
(P)
(P)
(P)
(P)
(P)
(P)
(P)
(P)
(P)
(P)
(P)
m JI«: to «h*k!*>-'j
sp. C
i o r s g n etc * us
corolialus
oculatus
mexikolpos
borceai
unietts
mediterraneus
irtfiatus
s p. A
vulv elia
l^ictciildtta
M^n t u o c e it1t«i% •» p A
P *, <><ssdo S.,1 is..ii'» ( P ) -! b f , •, t
P5.eudoia1t.a1s ( P ) c a l r f o i m c f r s i s
Ps e u d o t a n < i i 5 ( P ) f o r c i p a t u s
Pseudotanais (P)jortesi
P s e u d o t a n a i s (P) eolonus
P s e u d o t a n a i s (P) iilljeborgi
P s e u d o t a n a i s (P) n o d e n s k i o l d i i
Para p s e u d o t a n a i s abyss ails
Parapseudotanais sp. A
P s e u d o t a n a i s (A) i o n g i p e s
Pseudotanais (A)siegi
P s e u d o t a n a i s (i*) m akroltk r i *
P4 e-odwia i t a i ^ (A) n f i o i t « i i ^ > # n i
P •, e «<{ «> Li »k.,«11, ( A ) g & i ! j c h i
P •, ** «*!«1*1 ii a I •. (A) fl,ni'. i. i
P «,
<MI «i
» S,i it j i', ( A ) •. uci B!I',
P % t r i i d o Li is *ss (A) n u l j ^ n ' i »«,
Ps. *»udota n JUS (A) 5 c - i ^ p j n
P s e u d o l a o a i s (A) g u i l l e i
Cryptocope abiewiaia
Crytocopoides aroticus
Crytocopoides paclficus
C r y p t o c p o i d e s s p. A
iungentitartais primitras
Para inn g e n t i t a n a i s i o n g i d i g i t a t u s
Lafitanais beklemishe¥i
Curtiefoelia e x p r e s s a
S i n e l o b us S t a n f o r d !
Hargeriai Q p j x
T y p h i e l a n * is p annus
Figure 60. Character 34 trace - cheliped, dactyius setation
Marsupium
i
Psetidetartats (P) s p a t u l a
P s e u d o t a n a i s ( P ) s p. O
P s e u d o t a n a i s ( P ) s p. P
P s e u d o t a n a i s ( P ) a Ifin is
P s e u d o t a n a i s (P) ± calpellom
P<£eu*lotaft.aiS ( P ) I w EI g i>»" p iri u £
K'. tt«5<l*> Li Eui'i ( P ) d e i t t i v u l a l u S
P * e u d o IA iijs>> ( P ) k» |U<: ,tMs<»
P i. f-udo L* «^!% ( P ) ¥i!j»i^i
l*», <Msrfo Sj »,«»', {V) «* tjtp osii<:»i«,
P i <»«jdo I* IIJMS ( P ) m £C IO c l i r l # £
P s e u d o t a n a i s ( P ) s.p. C
P s e u d o t a n a i s (PJ l o n g i s e t o s u s
Pseudotanais (P) corollatus
P s e u d o t a n a i s (P) ooulatus
P s e u d o t a n a i s (P) mexikoipos
P s e u d o t a n a i s (P) b o r c e a i
P s e u d o t a n a i s (P) unieus
P s e u d o t a n a i s (P) m e d i t e r r a n e u s
P s e u d o t a n a i s (P) inflates
Pseudotanais ( P ) s p . A
P s e u d o t a n a i s (P) vulsella
P« e u d o t a n a k ( P ) f a l c i e u l a t a
M y , h i o c e nliu>k ^ # i u ! i i i
My4t I t i o c e n l i u s '• p A
P^ e u d o La ••«••% ((*) j i i f . '. i
Ps e u d o l a «**(5 ( P ) h atas i M u t i
Ps«Hdaljnai<i (P) ealrforniensis
P s e u d o t a n a i s (P) forcipatas
Pseudotanais (P)jonesi
P s e u d o t a n a i s (P) colonus
P s e u d o t a n a i s (P) lilljeborjpi
P s e u d o t a n a i s (P) n o d e n s k i o l d i i
Para p s e u d o t a n a i s abyss alis
Para pseudotanais s p . A
P s e u d o t a n a i s (A) l o n g i p e s
Pseudotanais (A)siegi
P s e u d o t a n a i s (A) m a k i o t h i i x
P v « u d o t a n a i $ (A) «ti« «SP»»•,<? !M
P * e u d o t j i n a i * (A) u w i l j e h i
P i t e u d o L i i M i ^ (A) y a u v s i
P s <>ud»y it a i* (A) s u n ill*
P s . e u d o f a n a t s (A) m a l a y e n * «w
P s c u 4 » i i n i j i & (A) s e r a p p i
P s e w d o t a n a i s (A)
fluiIIei
Cryptoeope a b i w i a t a
C lytoeopoides aretious
Crytooopoides pacifious
C r y p t o c p o i d e s s p. A
lungentitanais primitwis
Paraiungentitanais longidigjtatus
L a t i t a n a i s belt: ie m i s h e w
Curtitihelia express a
formation
$f*finsp'-ioitea*?
i ~ ^ 4 fli 115:!«,"»«
liziioM-Jigtf I S J rap a x
o T y p t i l w t n nmis parums
nC* ipo«ps«Md«s pi#sp*«tin«¥
Figure 61. Character 35 trace - marsupium formation
247
Pemopod 1 basis
length vs. width
t
1 < S x longer than broad
GZ3
jjll
• •
PS
S.1 to ?.9 * lofi^w than broad
8 to 9-9 K longer than broad
> 10 x longer than broad
equivocal
l^—^Pseudotanais
sfespmpseudotanals
Pseudotanais
Pseudotawais
i
Ps**ud*» la tiais
-^JI .
(P) s p a t u l a
(P)sp. o
(P) s p. P
(P) affinis
(P) s calpellum
: a P & e u d o t a n a i s (P) losiy is JHI>,H>,
I
j
x j P v e u d o L i i i ^ i * (P) d c t i t i e u l a l u f
—' rTJBBiP*' *ud*> Is lists ( P ) •> |H«:-tlsi'.
r j j p p p a i p ^ eudolan.au. (P) W I I M ^ I
*——-——33P\ • H d o i ^ l l ^ l ' k ( P ) n i p p o n i C U t
C s c w d o U i i i i i s ( P ) in Ji«roclieles
P 5 « u d » l « n « i i * ( P ) s p . C.
a P s e u d o t a n i i i s ( P ) l o n g s etosus
! l u P s e u d o t a n a i s ( P ) ©oroSlatus
i l Z P P s e u d o t a n a i s ( P ) ©euiatys
•xiPseudotanais ( P ) m e x i k o l p o s
: Pseudotanais (P) borceai
I D P s e u d o t a n a i s (P) unieus
raedrterrasieus
i n P s e u d o t a n a i s (P)
P s e u d o t a n a i s (P) i n f l a t u s
Psfeudotanais (P) s p. A
P s e u d o t a i s a i s (P) v u l s e l l a
Ps v u d o t a n a i t (P) laI«iCMlata
M y f f n o c e n t i u s >, e»s.il«",
My* t i i o c e n t i u * s p A
P <. <f>tid<n LJ » a i t | P ) J i>¥«. '. i
P « e u d a t * n « i * ( P ) b<*i?<t I M I I I I
P^eudotiPflJiis ( P ) e a l r f a i m e n s i s
___
Ps-«udoian<aii« ( P ) f o r c i p a t u s
I !l
-nPseudotastais (P) j o n « s i
*"~~™aPseudotanais ( P ) c o l o n u s
XiPseudotanais (P) lilljeborgi
SPseudotarsais ( P ) n o d e n s k i o l d i i
Parapseudotanals abyssaiis
V. : u P a r a p s e « d s f a n a i s s p . A
P s e u d o t a n a i s (A) l o n g i p e s
Pseudotanais (A)i;t«gi
Psttudutantais (A) m^ki4»thrty
P.v e u d o t a n j i v (A) tit o l i e n * e n i
m p * e u d « t a i l 4 i v (A) g e i l a o h i
...rp^CP*. iMnio Id sajs', ( A ) g j u v ^ i
; "T XlP % e u d o Uns%\ ( A ) \ raniliti
ft
J I
HZ
f
-.1
JTZIOP*. i M i d o L m a i * (A) m a l a y e n t K
'-"^Pieudolanai* (A)?«r*ppi
— - a j P s . e u d o t a n a i s (AJ g u i l l e i
ITZriD Crypto cope abiewlata
~
^ C r y t o c o p o i d e s arcticus
JP'W'Cri'tocopoides paeificus
••MCryplmcpoides s p . A
luisgeittiianais primitwus
Paraiuiigentitanais longidigitatus
Latitanais bektemishevi
Curticheiia expiess a
n Sine lob us S t a n f o r d !
EH
13 V. a »p» jips elides pio% p t c t n e j :
Figure 62. Character 37 trace - pereopod 1, basis length vs. width
Pereopod 1 basis
setation
C 3 snt-tfrts* ti3ieoiusi*fK*t
l i l i i l rsre tf i • t'iuti m iy
E 3
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
P*. tf«ti« I J ii-«is
(P)spattiia
(P) s p. O
(P) s p P
(P ) a t f i n i s
(P) s e-aipeSSum
(P) l o n g is pin u*,(P) d e n t i c u l a i M *
('•, t ' B ( l » f j l > - I i ' ,
(P)
rttkiV-H'S
P •„ <Msd« L i ti j r .
HUMMUSO
tt=Z==3=
J:
<t |» 1«S J i l l *
Ps e u d o t j i nait, ( P ) vilja.a
P ( # n d o l j i n j i « <P) n i p p f l i u c i i ^
P.". « * u d o l a n a i s ( P ) i n ^ c f n «l(^l#"t>
P s e u d o t a n a i s ( P ) s p. C
P s e u d o t a n a i s ( P ) I «* n a i * e t o s u s
P s e u d o t a n a i s (P) corollatus
P s e u d o t a n a i s (P) oeulatus
P s e u d o t a n a i s (P) mexikolpos
P s e u d o t a n a i s (P) b s i c e a i
P s e u d o t a n a i s (P) «ulcus
P s e u d o t a n a i s (P) m e d i t e f r a n e u s
Pseudotanais (P) infiatus
P s e u d o t a n a i s ( P ) s |*~ A
P s e u d o t a n a i s (P) vulsella
Pseinliti-injrs (P) iaicivuiata
Wyv h i o c # tii««>, «, »»i«\.iS«',
My*, t i t o c e • t i t u s *. p A
P% «<td«l<i I»-*I«. ( P ) * b y « - > i
P s f n i t « U »ai<k ( P ) h j i f . v n a u t l
Ps ewdolanai*.- (P) calHoiniensfei
P s e u d o t a n a i s (P) forcipatus
Pseudotanais (P) j » n e s i
P s e u d o t a n a i s (P) eoionus
P s e u d o t a n a i s (P) liiljeborgi
P s e u d o t a n a i s (P) n o d e n s k i o l d i i
Parapseudotanais abyssalis
Parapseudotanais sp. A
P s e u d o t a n a i s (A) l o n g i p e s
P s e u d o t a n a i s (A) s i e g i
P s e u d o t a n a i s ( A ) tit a k 10 t h risr
P t e u d o t a n a i v (A) m « i t e n f e n i
P *, 4*m:i« L i »i JII'I ( A ) y « i ! i c Iss
t^^'^sf
oo
(A)
JJII',', i
P'-,<:<•«<!«> 5.,i rs,u>, ( A ) •» isi» d«>,
P% *Hirf« l,nt-ii«» ( A ) n*n !*»¥«*»-. •*
P s « j « r f o l - i i*-t»«, ( A ) s « i a p p > i
Ps-eudotanais (A) g i i i i l e i
Ciyptoeope abreviata
Crftoeopoides areticus
Crytooopoides paclficus
C r y p t n c p o i d e s s p. A
iungentiianats primihflis
Paraiungentitanais iongidigitatus
L a l i t a n a i s b e k le m i s h. mm
Curtichelia express a
Sinelobus stanfordi
llargeria lapaK
Y^pblotniijfis parvus
<; atpojps*«d«s prof p*etn«$
Figure 63. Character 38 trace - pereopod 1, basis setation
Pseudotamais
Pseydotiinais
Pseudotanais
Ps««dotanais
P s « « u d o t a nai-s
Pereopod 1 ischium
setatson
(P'
(P|
(P
(P;
(P
s patula
s p. O
j p . P
a I f i n is
s caI p e d u m
l o n g IS plii l|&
d enlic ulalu*
P i eudotaftiiiS (P
P i *??S4loSiI El»t!<, ( P
P >:, <MHJ » L:l Bl ^i S >, ( P '. | ) I C j l l h
P s iMSdo L i Si^t!'. ( P
P •» * M M ! O i,i ti,is>.
Ps p i i d o t a D A I S
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
loPseudotanais
Pseudotanais
Pseudotanais
PSfcUdOtaiiaiS-
*lS;aXI
( P st i p p o l u e t i c
( P P i a 4:10 c h r l e *
(P
(P;
(P;
(P'
(P
(P
(P
(p
(P
(P
(P
l o n g is e t o s u s
GOTO
ocuiatus
mexifcolpos
borceai
unicus
m e d ite r ra n e us
inflatus
sp.A
*uKella
P ' j 1M5<it>trfBS,IS-i ( P
lalCtCUlalS
M^'iiiioct.'itliti1,
ii J I I I V
•,
My*i tssoc** s>tisi«. *<
Ps c«(i»l-i n a u ( P
P"i < > U < i l l M tlJI<>
A
j b f ,
', I
(P
Pseudotanais (P calrfomiP'nsis
P s e u d o t a n a i s (P fojcipaliMS
Pseudotanais (F Jones i
P s e u d o t a n a i s (P eolonus
Pseudotanais (P lilljeborgi
Pseudotanais |P nodenskioldii
Para p s e u d o t a n a i s afeyssalis
Para pseudotanais s p . A
P s e u d o t a n a i s (A) I o n gines
Pseudotanais (A)siegi
Pseudotamais (A) m a k r o l h r i x
P5 e u d o t a i i a i s (A) i i m i l e f t i t i i i
tiPs r i i d o i a n a i * (A) geilaclti
„...rj^BP<i e u d o l a i i a i s ( A ) g j i i 1 , ' . i
C
J u P * . e u d o t a n a i * {A) <• H I M l i t
~ BP $ t t u d o l * n a i s ( A ) nt a M y » n ^ fe»
""HfcBBPseudotanais (A) s ctappi
P s e n d « b n a i s (A) guillei
Cryptoeope abreviata
Crytocopoides arcticus
Crytocopoides pacifieus
Cryptocpoides s p . A
tungentitanais primitvus
Pa?a i t i n g e n t i t a n a i s I o n g i d i g i t a i u s
Latitanais beklemisheiri
Curtiehelia express a
Siitelobus stanfofdi
I I a f g e is J t a p a y
T y p M o l d nai<> p j i v u v
II a i p o a p % e 11 d «r << p j o". | K M : 8 si # •,
Figure 64. Character 39 trace - pereopod 1, ischium setation
Pereopod 1 merus
• D P s e u d o t a n a i s ( P ) s patusia
Pseudotanais C
( PP)) s p. O
P s e u d o t a n a i s ( P ) s p. P
p P s e u d o t a n a i s (P) affinis
nPseudotamais (P) s calpellum
P ^ e w d o l a n a i - * ( P ) t ^ n ^ t s inn>.i':.
P v e u d t t L i i u i * (P) d * i i t i c u l a h i \
P "^ 4f ?s<J« Li it«t:', ( P ) «, j*s« JSW',
setation
• H I ans sfts
S B
Ps cudi»ljin«iii<t (P) ¥iljj!ja
'!U l,Ula-:J
; • [ * ' , p ! l d » b f U I ' . ( P ) it i p p O I H C M i ,
^ P i eudotanais
E—JnPse«dola n a i s
joPssudotanais
uPseudotanais
'.'.—igPseudotanais
isPseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudatanais
XsPseudutanais
5nPs#*id«tawait
My% t u o c e ntiuv
I
I
I
( P ) macroeh<*!•• *
( P ) s p. C
(P) longis etosus
(P) ©orollatus
(P) oculatus
(P) mexikolpos
(P) b o r c e a i
(P) unicus
(P) m e d r l e r r a n e u s
(P) inflafus
( P ) s p. A
(P) vulsella
(P) lAlciculata
teiulus
•jaP** #u4«il*nMi\
(P)*by*ti
icf*** « n i d o l * n a i * ( P ) bai#*
njuti
Ps£#«d»ta«a»5. ( P ) e a l r f o t n i e n s i s
P s e u d o t a n a i s (P) forcipatus
P s e u d o t a n a i s (P) j o n e s i
P s e u d o t a n a i s (P) oolonus
2BPse«idotanais (P) l i l l j e b o r g i
P s e u d o t a n a i s (P) n o d e n s k i o l d i i
fzzsnP a r a p s e u d o t a n a i s a b y s s a l i s
fl^n P a r a p s e u d o t a n a i s s p . A
P s e u d o t a n a i s (A) l e n g i p e s
Pseudotanais (A)sie§i
P s f e u d o t a n a i * ' (A) m a k r o l h r i x
3D PS ewdotanai-S (A) i n o r t e n s e i i i
V\ e u d o l a nm\ ( A ) g e i l a c h i
irpaaP* «udotan,iii% (ft) g j u * * i
" i a a P \ «»udota I U I S (A) % i m i l i s
• H r « » i i d » l j n 4 i s (A) n u l a y e n s K
T h « B P s « y d o t a i R ^ i i s (A) s a r a p p i
l a P s e u d o t a n a i s (A) j j u i l l e i
13C r f p t o c o p e a b r e v i a t a
wiCrftocopoides arctieus
BJCrytocopoides pacificiis
illC r y p t o c p o i d e s s p. A
lungentitanais primitais
Paraiungentitanais longidigitaius
Latitanais beklemishewi
Curtiehelia express a
_j"w53?>lit«lobus
Sittelahu-s ^itit a n l o r d i
rrr
ZD<* <iipo4|»* wiidiM p i o * p#cl»»«»s
Figure 65. Character 40 trace - pereopod 1, merus setation
Pereopod 1 carpus
setation
t
i fry f>* •$;!$. %$Jg§
cm
r~i
tve setae
f t i i l :.r! j r
mm H1l## S*t#§
HB§
?*,*J t tSil»B
WKKH muf set*
f
;0 **ta$
••
gza fcqucH:*
P s e u d o t a n a i s (P) s p at u I a
P s * « d o f c a n a i s ( P ) s p. O
Ps^wdolanais (P) s p . P
Pse«d<*ta«ais (P) affini&
Pseudnntaskais
P$«udotiift<iis
P*. e w d a L a i t i i i v
P\#udot4ndn
Ps »ndoLin.ai%
P•, *•• ?s<l<*!»i t»«ts•,
P*i ewdo La itjn<d
Ps e w d f l t a i n ^ i s
Pssud^tamais
Pseudotanais
Pseudotanais
Pseydotanais
Pseudotanais
P s e u d a t a nais
Psendotanais
Psfcudotanais
Ps<&ud«*tanais
Pseudotanai*
(P) s eaipeiium
( P ) l o i t y i& p i n us
(P) denticulatus
( P ) * pic . i t t i t
( P ) vil|ju9i
(p) nipponicux
(P) m jaiQchcltts
(P)sp. C
( P ) l o n g is rinsus
(P) eorallatus
(P) oculatus
(P) mexikoEpos
(P) borceai
( P ) lira i c e s
IF) medfterraneus
(P) inflatus
( P ) s (t. A
(P) vulsella
P t ' e u d t t t a n j m
( P ) falciClllatj
M y \ i n o c e iiSu«% l e i u i i i s
My', iti*n:*f i l l J««*. ; ft A
P «, «Mf<f<» la nAiv ( P ) 4 b ^ ' , i
P <, «Mnf « I.a it JI*« ( f )
bjn»'jiM!ili
P s e u d o l a ijass ( P ) c A l i f o m i e n s i £
Pscudotanais (P) loicipatus
P5*sudn>tanais ( P ) j o n e s i
Pssudotanais (P) eolonus
Pssudotanais (P) lilljeborgi
Pset.tdofciBa.is ( P ) n o d e n s k i o l d i i
P a r a p s e u d o t a n a i s abyss alis
Parapseudotanais s p . A
P s e u d o t a n a i s (A} l o n g i p e s
Pseudofanais (A)si#gi
P s e u d o t a n a i s (A) m a i t i o l h i i x
P»> e i i d o Lt itJiiv ( A ) Eft o i l e n i e i l l
P * «M:dotj n*i\ ( A ) g e i l a i c l i i
P % e u d o Li itit:s (ft) $ ,na>, •« e
P «. (M!ti»l,i is-us (A) s imds',
P % »MJ«I<> L i l i ^ i t v ( A ) i n j Ijy<*«»', !«,
P& C U < i « t 4 (SJ«IS- ( f t } S- C l ^ p j M
P s e u d e t a t i a i s (A) g u i l i e i
C r v p t o c o p e abrewiata
C r y t o c o p o i d e s artstieus
Crvtoeopoides pacificus
C r y p t o e p o i d e s s p. A
l u n g e n t i r t a n a i s primirltwus
P a r a i u n g e n t i t a n a i s l o n g id ig Status
L a l i t a n a i s bek i e m i s h «•«
Gurtichelia exjtiessa
$itiel*»btis s t a n f a r d i
H a r g t i i n rapax
T y p M o t x n a i s parvus
C a r p w a p s e u d t i p»#s jp *etni«^"
Figure 66. Character 41 trace - pereopod 1, carpus setation
252
Pereopod 1 propodus
length v*. width
1 #,1 to 8 x longer Hum broad
1 >9n
equivocal
Pseudofanais ( P ) s patuia
P s e u d o t a s i a i s ( P ) s p. O
P s e n d o t a n a i s (P s p. P
P s e u d a t a n a i s ( P a i f in is
P s e u d o t a n a i s (P $ o a i p e i l u m
P ^ e u d o t a A J K ( P t o i t y is p i l l u«f
p>» «?«<iofj ti,uv ( P d enlfi*: !!S/!li» *,
P»i **!!«!« L H U f i ( P *j p i c a t m
P*, tMitlt>IJi>ai>. ( P
P «, <•- ii <i» !.,• ti -a i', ( P It l | » p < > I > I < * < J > ,
P s t^snf «Sii !*4ir, ( T m ^ c ! « c ! l « l l > V
P s e u d o b n s i s (P s p C
P s e u d u t a n a i s (P, I n n g f e e t o s u s
F s e u d o t a n a i s (P* e o r o l l a t u s
P s e u d o t a n a i s (P. o c u l a t u s
T n p P s e u d o l a n a i s (P' m e x i k o i p o s
— i o P s e u d o t a n a i s (P^ b o r c e a i
, n P s e u d o t a n a i s (P y nicus
- ^ - r o P s e u d o t a n a i s ( P ' m e d i t e r ra n e u s
( — i p P s e u d o t a n a t s (P' i n f l a t u s
, l™,„, p £ ft u j 0 ijm n a j s ^ps p. A
P ^ e u d o l a ftai«i ( P v u h e l l a
:nP?
ftuilotanaiv
(P f ^ E v i c u i j t a
r z z n a M y i I n o e e n l i u * i * tt J i l l *
——| t—^fct^., | m m » nlm». %
A
-—•—•^nP * «*nd© I J n J K ( P J b y , >, i
z z z z z x i P i 4»Mdotait»Js»^ ( P b a r o i n a u l i
j - :uP-seud«tan»iis (P c a l i f n r n i e h s i s
n J z z D P s e u d o t a n a i s (P; f o t c i p a t u s
jiPseudotanais
jonesi
111 anPseudotanais
eoionus
^oPseudotanais
iilije borgi
TiPseudotanais
nodenskioldii
.__ r p ^ P a r a p s e u d o t a i i a is a b y s s a l l s
1 ' r>P a r a p s ee uu dd oottaannaai s s p . A
P s e a d o t a r t a i s (A) i o r i g i p e s
Pseudotanais (A)«riegi
P*; e u d o t a n a i s ( A ) m a l r i o l h r i x
P«, e t i i l o L i it.ii'i ( A ) nt o r t e t i f e n i
J PE
i
B
„
!
in
P •, * M H I O L I is j i ' i
(A)geHach)
P •, «'«srf« Li »b*«i'» ( A ) (j j(«', «i s
PS ( H l d o L t l t JIS ( A ) ', HSi sSl<.
P'• ( M M ! <> i J n ,I I s ( A ) ra-ib^pii'.
..4,
r,
P s e u d o t a n a i s (A)*«i<ftppi
::c;Pseuri<»t3nais ( A ) g u i l l e i
:DCffpt«cope abrewiata
TtCrytocopoides aretictis
Crytoeopoides pacifieus
\1 u C r y p t o c p D i d e s s p . A
lungentitanais primrtwis
P a r a l u n g e miliaria is l o n g i d i g i t a t u s
Lafiianais bekiemisheiw
Cnrtieh#lia express a
_ r — n S i n e l o b u s st»nf«»rdi
L r z i a H a i f l e i i a rap ax
uTyphlobiiiJis parvus
IZZZPU -iipo4p,f t u d « * p f « : f p « d t t « S
Figure 67. Character 42 trace - pereopod 1, propodus length vs. width
253
P&mopod 1 propodus
se tali on
I . f •wireus %mm
C D iSurseta*
mm
nut
•1
•1—I•
titrrr* r.«*i}« i
TtfJ StliS
urtte
-itiii
ftO l*ta»*
&<JSKH3>
3DPseudota»ais
nA P
e «;
uM
d <u
o tl a m a i s
! r s sJ — ^ u P s e u d *o t a n a i s
[ _ _ f TBHIP S « W do<t a r i a l s
|
1 • — ' n P s t S t l do*t a n a i s
P v e u d o L l nai-s
P* etidttL) i w m
k
. ,
uP . •ndoLin<iiir
"-'< • • « • ' * ' » n i d o L m a i i .
I,*—— - ~ - T i P i ' « t w d i » t i i m i i ^
1-pgDlPs u n d o L i n * ! *
! L Z n p 5 e „ jf» U n a is
j
uPseudotanais
-jJ^pPseudatanais
——siPseudotanals
rpnPsetidotanais
_Jf
f P ) s patuia
(P)sp.O
(P) s p . P
(P) affinis
(P) * caip^tiuin
( F ) lofigi;, |»!ii«i
(P) denti«ul4lilf>
(P) * picatu*
(P) vit|*£i
(P)mppomeus
( P ) m ats i n cl»«»l***
(P) s p fl
(P) coroilatus
(P) oculatus
(P) mexikolpos
(P) b s r c e a i
^-^-pPseudotanals (P) mediteirarteus
P s e u d o t a n a i s (P) inflates
P a e n d a t a ft a i s ( P ) s p . A
P s e u d « Li i i j i s ( P ) w u i s e i L i
P f e u d t f l a i l j i V (P) f j l c i c u l j l *
My* l i i o c t ' n i i u t
t o i u t n
Mf>, t t U t C # I t I i « ' : , '. ft A
p P * * m d o Lt i**i% ( P ) *l>|Ki % i
n P i ^ u d o I * i t * ) * ( P ) h atf»% n a u f i
_j
u P * « M d » i * n * i s . (P) calitorni«nsi$
r-^ r r i z z o P s e ; u d o t a n * i s ( P ) f o i c i p a t u s
r—'I ..—-nPseudotanais
(P)ionesi
P s e n d o t a n a i s (P) colonus
zzzzoPseudotanais (P) lilljeborgi
;—7~nPseudotafiais (P) n o d e n s k i o l d i i
f p a n P a fa p s e u d at a n a is a b y s s a l i s
"L. a j P a r a p s e u d o t a n a i s s p . A
j — n o P s e y d o t a n a i s (A) l o n g i p e s
i—:nPseudotanais ( A ) s i * g i
P s <ii»<l<tta l i i i s v ( A ) in 3 k ? < t t h i i K
; D P ^ « M i d o t t n'jt'i% ( A ) m « i i t « n ^ w n i
P •> «?li<S« Li n*i<k ( A ) g e-lSiichi
P « t k u d « U i i d i ^ ( A ) y ,«<!',», e
P •» <Mtd« Li i i ^ i ' , ( A ) >• sm i!i>.
P s ? n d o I * »*•<> ( A ) i n * M y t ^ i n w
P s e u r f n t * n*«j. ( A ) s c i * | > p i
P s e w d o i a n a i s ( A ) j) u i II e i
Cryptocope abreviata
Crytoeopoides arctlcus
Crytocopttides pacificus
C r y p t o c p o i d e s s p. A
lungentitanais primitvus
P ara iun g e n t i t a n ais l o n g i d i g i t a t u s
L a l t a n a i s b e l t ie m i s h e w i
Curtieh^lia express a
Sin^l&bds stanfardi
Harg#ria r^pax
T y p h i ©tana i f parwus
33Caiponps«Mdiis |ir»spt«lnts
Figure 68. Character 43 trace - pereopod 1, propodus setation
254
setae
C. J prss€it
• B abs*r:
-CPseudotanais CP) s palula
(P) s p. O
laPsetidotanais (P) s p. P
_ _ j — l y P s e t i d o t a n a i s (P) aHinis
L~33f*£'ud*tanais (P) s eatpeltum
:nP?eudotanais (P) fouu.i<; piitwv
puPseudoianals
P», tMKJoLi BS
I j8<,
P i e u d P L I 11*11%
P%eudolanai\
P i * u d o IJI n a n
P5Cud«liiii<ii5
Pfeudotanjus
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Ps eudota fijis
jnPseiidoianais
My* Inoce nlius
Mf% l i i o c ^ n i i u i
(P) denticuldlu*
( P ) •» pstj!««•:.
( P ) wsij^/a
( P ) it i p p o m c u i
(P) in<• ciodue!«»•:»
( P ) s p r:
(P) long is etosus
(P) corollatus
(P) oeulatus
£P) mexikolpos
(P) butreeai
(P) unicus
(P) m e d i t e r r a n e u s
(P) inflates
(P) s p. A
(PJ vols ella
(P) t J U i c u l a t a
teiialui
'. p
A
jnPi ^ i i d o L m i t u (!*) j l ) f , ' u
[pP* ^ u d o t a n a i s (P) i»Jin»iitau , K
P s e w d » I * n.»is (P) calrfoi metis is
P s e u d o t a n a i s (P) foteipatus
Pseudotanais (P)jomesi
P s e u d o t a n a i s (P) colonus
P s e u d o t a n a i s (P) l i l l j e b o r g i
30Pseudotanais (P) modenskioldii
Para p s e u d o t a n a i s abyssaiis
Para pseudotanais s p . A
P s e u d o t a n a i s (A) iongipes
Pseudotanais ( A ) s i e y i
P s e u d o t a n a i s (A) maitiothiiX
P s « u d « ! j H J I ^ ( A ) iss i» r t « n ^ e mi
P t eudo1.ii i t , i i ' . ( A ) 9 e-si.^4; It•
P i <ME<io IJUS JJ«, j'A) g J U S s i
P-j l»M(J« I J l i J I * . ( A ) 1 HIM 111
P i r i t d o l a n a t * (A) m a layrfts is
P s <*«rfo t«i » J M (A) -i «rap|>i
P s e u d o t a n a i s (A) quillet
Crfptoeope abr«¥iata
Crytocopoides aroticus
Crytocopoides pacifieus
C ryptnopoides s p . A
lung entitan ass primitwas
P ara ion gentitan ais Ion gidigiiatus
Latitanais bek Semis hew
loCurtiefaelia express a
W S i n e l o b us
stanfotdi
JjTyp h l o t a n a n p a i w u i
ID (- <& i p o 4 i p i v- o d e i p i o i p e c t n e v
Figure 69. Character 44 trace - pereopod 1, dactylus setation
255
Pereopod 1 dactylus
seta! type
• • I '.mglift, irm:l
PseudotaRals
Pseudotamais
Pseudotanais
Psftwsdoiamais
P s ewdotanaite
P^eudtotaftai-%
P*< e u d o i a n a i t
Pn e w d o t a n a i s
(P) s patula
( P ) s p. O
( P ) s p. P
(P) affinis
( P ) -s ealpeEium
( P ) l o n f l i s psftlis
( P ) denti<sulafeu$
(P) s picatu*
P» etldolaiiai'V
( P ) Vlt|«I.Ai
Ps «»udntanai<i
P* * u d t t t « n a i i
Pseudotanais.
Psewdotamass
Pseudotanais
Psewdotaisais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudetanais
Pseudotaitiiis
Psftudsitanais
( P ) it ipp oiti«u$
( P ) ii»ac«»c!»t*ie;f
(P) v p , C
( P ) I O D Q I K et«*s«s
(P) eorollatus
(P) d^ulatiis
(P) mexikolpos
(P) b»reeai
(P) urticus
( F ) mediterraraeitis
(P) inflates
( P ) s p. A
(P) vulsella
P«fcudolaiiaHr
( P )
lalttltiulata
y f . l i m t e i t t i i ! ' , vwiiatus
My % l i i v c e itlMis '• |> A
P,s e u d o l a na«% <(P) a by& * i
Ps e u d o t a n a i * (P_) b a»**"» n a u l i
Ps e w d o t a m a i s ( P ) o a l r i o m i e n s i s
P s e u d o t a n a i s ( P ) fo<r*tpatus
Pseudotanais (P)jonesi
Pseudotanais (P) colonus
Pseudotamais (P) lilljeborgi
Psssudotanais (P) n o d e n s k i o l d i i
Parapseudotanais abyssalis
Parapseudotanais s p . A
Pseiidetanais (A)Iengipes
Ps«wdota**ais (A)sifcgi
P s e u d o t a n a i s (A) t i t a k i u l h i t *
Hv e u d o ta i t a i ^ (A) in o itett*.te>t>
Ps e u d o ta itais (A) g e i l a c l i i
PN
t M i d o i * ! itillS
(A)
Jii!!\SI
Ps (Midola I M I « (A) t int ids
BIB
f* <. *Mido I J n a i s (A) I B J I J ^ P H ! . K
Ps e i j d n l a n a i s (A) & c r a p p i
P s e u d o t a n a i s (A) g u i l l e i
Cryptocope abreviata
C r jt©© o p © i d es a rctiens
Crytocopoides pacificus
C r v p t o c p o i d e s s p. A
Itingentitanais primltvus
Paraiumgentitanais Ittngidigitatus
Latitaitais bekleinishewt
Curtichelia «'y|ift'ssa
Sirt«lubu& s l a i t t o r d i
H a i y e ii^i l a p a x
lyp ItEuLi it Afi p a i v u «
33 ^; *i i p o iips f ud«<w p i o* p t>cln«%
Figure 70. Character 45 trace - pereopod 1, dactylus setal type
256
Pweopodldactylus*
vs. length of propodus
Mill
s Uli flfc< iii '-'I f "»'. 3f! fr |r»
'Pseudotsnais
Pseudotamais
Pseuilbtanais
Pseud© taraais
Ps feudetanai«
P*. «-ud» I a n n i s
Ps e u d o f a n d i v
P'i t'?ni<»E«t BiJSI',
SQtl.'M*
( P ) s patula
CP)sp. 0
( P ) s p. P
(P) a If in is
(P) * £ai|»eliutn
(P) lofty »•» piit u%
(P) 4«nt««u 1 alus>
(P) % pi«atu%
P«, e u d e i a i t ^ m
(P)
Ps f u d t l j !i.ir,
(P) ti ipp wisictis
(P) maaiocli* 1 ')**
(P)sp C
(P) longfc etosus
(P) c o r o l l a t u s
(P) o c u l a t u s
(P) m e x i k o l p o s
(P) b o r c e a i
(P) u n i c u s
(P) tried iter ranees
(P) i n f l a t u s
( P ) s p„ A
(P) v u l s e l l a
(P) talttiftulata
P<i p u d a t a n a i t
Ps <Fu4«tan<ii'c
Pseudatanais
Pssudotanais
Pseudetanais
Pseudotanais
Psjsudotanais
Pseudtttarsais
Psetidofanais
Pseud^ianais
Ps eutltttanais
P*> e u d # b n a i s
Ps eud«tan.di.£
¥tlf J ^ I
My% tii©c«*i*lnis s p A
Ps e u d o l a n a i * (P) b a i e i nasi h
Ps ewdotanais (P) c j l i f o m i e n s is
P s e u d o f a n a i s (P) f o r c i p a t u s
P s e u d o t a n a i s (P) j o n e s i
P s e u d o t a n a i s (PJ c o l o n u s
P s e u d o t a n a i s (P) l i l l j e b o r g i
P s e u d o t a n a i s (P) n o d e n s k i o l d i i
iParapsewdotanais abyssaiis
iParapseudotanais s p . A
iPs e u d o t a n a i s ( A J I o n g i p e s
[Pseudotaiiais ( A ) s i e g i
P'i «?«<!» lit rs»ti', (A) malfrtothrix
P f e u d o t a n a i v (A) m o d e n s e ni
P v <-u «f» Lt I I j i •,
(A)gefl<lchi
Ps •ad«tanaiN (A) g a u s s i
P <.<•!!(!« i j SkJJ«, ( A ) * i l N l l l *
P 5 * u d o t a n a i s (A) malay«»n* »
i P s e u d o t a n a i s (A) s c u p p i
P s e u d o t a n a i s (A) guilSei
Cryptocope abrewiata
C r y t o c o p o i d e s aieticus
Crftocopoides pacificus
C r ^ p t o c p o i d e s s p. A
l u n g e n t i t a i t ais primHwtis
Paraiungentitaitais longidigitatus
Lafitanais bekiemishevi
C u r t i c h e l i a esqpress a
Sineiebus stanfardi
H a i g e t i a rap a v.
IfJfltl«li»SiJi:-»
pami$
C a r p w a p s « i i d « « prosp«etn#.$
Figure 71. Character 46 trace - pereopod 1, length of dactylus + unguis
Pereopoda 2 & 8 basis
setation
E D bream *is«oniv
fMNt r»nr;plf r.r* i r t r '#
M B
HO I.€"af
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
P» « u d « * t a n a i s
Pseudotanais
(P) s patuia
( P ) s p. O
( P ) s p. P
(P) aftinis
(P) s calpellum
( P ) SongiS p m i i v
P«eudol4nai<!k
( P )
d*lltl«UI4ht<k
P t r u d t t tanai* ( P ) t p i c j l t i t
P N HMidoLan*!* ( P ) *S3JJ,TI
P<i ( M i d e t a n a i s ( P ) i i i p p o m c u s
P* e u d o t a n a r *
(P) HIJUSIOCIH*!**;
P s e u d o t a n a i s ( P ) s p . r.
P s e u d o t a n a i s ( P ) l o n g i ? <*tosus
P s e u d o t a n a i s ( P ) nsorollatus
P s e u d o t a n a i s (P) ©culatus
Pseudotanais (P) mextkolpos
Pseudotanais (P) borceai
Pseudotanais (P) unicus
P s e u d o t a n a i s ( P ) m e d iter r a n e u s
Pseudotanais (P) infiatus
Pseudotanais ( P ) s p . A
p£«udotamaiv (P) * u K e l l a
P v e u d o t a n a i s (P) falciiMilata
My*, inoeeniiwuk t e n a f u N
Myv I l i a c * ntiu% v p A
P<i e u d o t a n a i \ ( P ) a b y i < i i
P* e u d o t a n a i s ((P) h ate*, n a y b
Ps *w«lie»la n a i s { P ) o a l r f o r m e n$ is
Pseudotanais (P) torcipatus
P s e u d o t a n a i s (P) Jones*
Pseudotanais (P) eolonus
Pseudotanais (P) lilljeborgi
Pseudotanais (P) n o d e n s k i o l d i i
Parapseudotanais abyssalis
Para pseudotanais s p . A
P s e u d o t a n a i s (A) i o i t g i p e s
Pseudotanais (A)siegi
P s e u d o t a n a i s (A) in a l i i o t h r i x
Ps * u d o t a liars (A) i n ^ i t t n s i ' i i i
P'i «M!«Jo Sj !»,(!«, ( A ) g e i l j c l t l
P^pudotjiiiii^ ( A ) g j u ^ s i
Ps *ll<lolanai*
(A)VlllllllS
P ^ t i i d o b f t J i n ( A ) m a l a y e n & i%
Ps«ndoianais (A)sciappi
P s e u d o t a n a i s (A) g u i i i e i
Cryptooope abrewiata
Crytocopoides arcticus
npsin C r f t o e o p o i d e s p a e i f i c u s
if u C r y p t o c p o i d e s s p . A
l u n g e n t r t a n a i s prisnltwus
Paraiungentitanais longidigitatus
Lalitanais ~beklemishewi
Curticheiia express a
S i n e lob us s t a n f o i d i
H a r y e n a sapax
I y p h l o l a i i a i s paiWJK
%'. a t p o a p k e w d e i
Figure 72. Character 47 trace - pereopods 2 and 3, basis setation
p i »•. p « c l n # f
" ~ i»Pse«j<l«ttaitais
nPseudotamais
_LjPseiid»iattais
, uPseudotanais
—rsPseudittanais
TmnHveudotanais
•i e u d o t i n a i v
Pereopodii 2 & 3 ischium
setation
Lir-c:
1
J"!
uPs ptidoM nai*
—nPteudetanais
—TDPseudottHiJiis
uPsewdotanais
- -t|Ps e w d o t a n a i s
—nPsewdotanais
Pseudotanais
_!_^uPseudotanais
pnPseudotai»ais
—— P s e u d e t a n a i s
I— [""J . u P s e i u d o t a n a i s
nPs«udotanars
-ol'iewilwtiftjisi'
(P)
(P)
(P)
(P)
(P)
(P)
(P)
(P)
(P)
(P)
(P)
(P)
(P)
in a e r o «;»<»•>'$
s p. C
lining is e t o s u s
eorollatus
oculatus
mexikolpos
bsrceai
unicus
med(terraineus
inflatus
s p. A
vulsella
f jlcsiculata
leiiatuv
SfjMyi t t i o c e i i l i u t * p A
TpP> tMidt»tai»ais ( P ) a b y » « i
3nP$ e u d o t a nai$ ( P ) b ait» s nMU%
.uPs«*«dt>tanais. ( P ) c a l r f o i i i i e n s i s
- n P s e u d o l a n a i s (P) fortupatus
"nPseudotanais (P)|*jnesi
.uPs«sudwtanais (P) (jolnnus
• P s e u d o t a n a i s (P) l i i l j e b o r g i
u P s e u d o t a n a i s (P) nodensScioIdii
as P a r a p s e u d ©tan a is a b y s s a l l s
uParapseudatanais sp. A
i l P s e n d t * t a n a i s (A) i e t t g i p e s
J ^
ni
—nP* etidotanais
UP* e u d o t a n a i f
u P i e t i d t i L i ss.is'i
4* U li <> t i l SI ,t ! S
—-np.
( M i d o L l 16*1!',
—DP-UP« <»«sd*• 1.1 s s j i « .
eudotanai*
D
eildolan.ai's.
U WJIMUHlMMtMmiimMiUiU IMJJJHJi l.l iSUU l O U U l l L U
trad6
spatula
& p. O
s p. P
affinis
s calpellum
l o f t y i£ pmw,
d enlieuS*iin »,
N « « i d o i d it At: ( P ) '. pic<iiii^
s p i i d o ! J ii.dc*. ( P ) wi!|a^c
[jP«i <J-!S<l«Ll !»..tl*, ( P ) St ipjl tilt It! I I *
3QMys i n o . c e siting
1
(P)
(P)
(P)
(P)
(P)
(P)
(P)
(AJsiegi
(A) m a k i o S h s i x
(A) its o it«n4 e it:
( A ) g «Sl4«cli!
( A ) fl ..«?!•, '. I
(A) •, imiit*i
(A) m a l a y e n s i*
(ft)
5
(Mppi
u P s e u d o l a n a i s (A) g u i l l e i
DC r y p t o c o p e a b i e v i a t a
nCnjftoGopoides arcticus
aaCrytocopoides paeificus
C i y p t o c p o i d e s s p. A
l u n g e n t i t a n ais p r i m i t w u s
Paraiusgemtiianais iongidigitatus
L a t i t a n a i s bek lent is h e w
Curtiisrielia e g r e s s a
i n t s t a b u s ss t a n f o r d i
rar S
i»in«i0l»u$
t i M i H - n j e i i J iapax
ICK aipti.&p'* * « d i f > pi«*« p t t y l n t i
Figure 73. Character 48 trace - pereopods 2 and 3, ischium setation
Pseiidotanais P ) s pates a
Psetidotanais P ) s p. 0
Pteraopods 2 & S
merus setation
nurtweii® marginal
C!3 3 svtfNt
mm 2setaa
Pseudotanais
Pseudotanais
Psfcudotassais
P) s p. P
P) a If in is
P) s c j i l p e l l u n t
P v
P*
F ',
P>»
P ) i o l t g l!» |>!Si «'i
e ud«fa
e m i o y
c a d « Id
r n d t l j
it a f*
itiH*.
siiii<»
itjr.
P) d «!>!:£ u l d h i ^
P ) S p i C did 1 !.
P)
VtlfdJ?
P) s» tpp o i t i c n ^
P ' , e t a d o t d s*^i«. P) m ar. to ch»l*<t
P <, H» t« d W t d St d IS P ) s p C
P s e u d o t d n d i s (P) lonprc etosus
P s e u d o f a n a i s P) c o r o l l a t u s
P s e u d o t a n a i s P) o c u l a t w s
P s e u d o t a n a i s (P) m e x i k o l p o s
ma* P s e u d o t a n a i s P ) b e r c e a i
^ t i P s e u d o t a n a i s P) u n i e u s
P s e s i d o t a n a i s P) nsed iter r a n ens
P s e u d o t a n a i s P) i u f l a t a s
P s •-!MJ <V 8 J It ,1 E',
PsfcUdotandlS
P)sp. A
Psettdotaifiail-s
P) v u l s e l l a
P> l a f c u u E d t d
Mys Iii«*<» iifsus-,
fciyn I n a G e t t t n i s
'» «' S! d f l i " .
P S * li ft O i d St d I % P )
P<i # n d « Id ndi%
E
J
by^ % i
P) b Jipi
ndisfi
P ^ e m d o l d r t d i ^ P") c d l i f o i m e n s i s
Ps e u r f o t i i n * i S i P ) f o t c i p d l w s
F s e u d o t a n a i s PJ J o n e s i
Pseudotamais (P) colonus
P s e u d o t a n a i s (P) l i l l j e b o r g i
P s e u d o t a n a i s (P) nodeinskioldii
Parapseudotanais abyssalis
Parapseudotanais sp. A
P s e u d o t a n a i s (A) l o n g i p e s
Pseudafanais (AJsiegi
P s * u d * » t a t t a i £ (A) m a k r o l h i i x
Ps « u t l o t d » d i s (A) m o i l e i i t f a r i i
P •:. e««loLi a^ia'. (A) g e t l d t s t t i
Ps e u d o l d i t d i s (A) gaiis. <• i
f*% f t i d t t t d n d t s <A) 5 u m l i s
Ps « w d « l d itdts (A) i« d Idjf^ns •«.
PseudfftanJus ( / ^ s e t d p p i
Ps « u d o t a n < i i 5 (A) g u i l l e i
Crfptocope abreviata
Crftooopoides arcticus
Crftoeopoides pacificus
Cryptocpoides s p . A
l u n g e n t i t a n a i s primitwus
Paraiuitgentitanais longidigitatus
L a l i t a n a i s besklemisheiri
Curtiehelia express"a
Sin globus Stanford!
f i a t g e i i a t a p AX
Typ h l o l d u a i v pj:«u<>
£ (I d l p O d p S * l l d e s
Figure 74. Character 49 trace - pereopods 2 and 3, merus setation
j>S <>', J M M ; ! 5 M M : ,
260
Pereopods 2 Si 3 mams
spatulate setae
i i present
P s e i t d o t a u a i s (P) s palula
Pseudotamais (P) s p. O
P s e u d o t a n a i s (P) s p. P
P s e u d o t a n a i s (P) affinis
P s e u d o t a n a i s (P) s e a i p e l i u m
P-i eud«*l<l nais (P) lung e> |>5Si (i-t
Ps eudwta »*•!% (P) d enlictiliilu •»
P«< eudo ta aidi'i (P) ^ picdiu*.
P «. r u d o I..I n-ij*, (P) ¥ i ! ; j ^ i
P', «MH1O I,t ii^s'j (P) n i p p a m c ! i , l
P <; «H!*I<>!..* «»„us (Py m ACI*> r.fovU*',
Ps etsdolanajs (P) «r p. C
Ps«Mdot»r(ass (P) lowpts etosMS
P s e u d o t a n a i s (P) corollatus
P s e u d o t a n a i s (P) oculatus
P s e u d o t a n a i s (P) m e x i k o l p o s
P s e u d o t a n a i s (P) b o r e e a i
P s e u d o t a n a i s (P) unicus
P s e u d o t a n a i s (P) m e d i t e r r a n e u s
P s e u d o t a n a i s (P) inflates
P s e u d o t a n a i s (P) s p. A
P* wudtttamiSs (P) v u l s e l l a
P<s eudotanai? (P) f a l c i c u l a t a
My* Inoee n t i u * % e i u i ! t ' ,
My*> IHWC<»IIIIU% \ p A
P*. «nid«l^n«is (P) <ibft i i
P<t * w d o l a n a i \ (P) It ,is«>', it-lie h
Ps«wdot>in<9iv (P) cairfoi nse ns 15
Ps eudotaraass (P) f o i c i p a t u s
Pseudotanais ( P ) j o n e s i
P s e u d o t a n a i s (P) eolonus
P s e u d o t a n a i s (P) l i i l j e b o r g i
P s e u d o t a n a i s (P) n o d e n s k i o l d i i
Para p s e u d o t a n a i s a b y s s a l i s
Paia pseudotanais s p . A
P s e u d o t a n a i s (A) l o n g i p e s
Pseudotanais (AJsiefli
Ps e u d u t a n a i s (A) n* akiotliriix:
V '. eu«lufa nAi*; (A) tit o itwiiT e ni
P^ 4Ml<ioLltl,IJ', (A) J|4MLn:ilB
P >, <MB«ltt Lt ES.tS', ( A ) {|»t»8'«S!
P % *• II <i O \* I i J I « . ( A ) «. l l » l i l * >
P'i c i n i o b i i - j r . (A) sn *« layeii;. »*
r&ewdotanais' ( A ) & c i a p p i
P s e u d o t a n a i s (A) g u i l l e i
Cryptocope abrewiata
Crytoeopoides arrfieus
Crytoeopoides p'acificus
C iryptocpoides s p. A
l u n g e n t i t a n a i s primitwus
Paraiungentitanais lengidigitattis
Latitanais beklemishewi
Curtiehelia express a
Sinelobus stanfordi
HaYge.fia rap ax
Typ St. I ofa ii Ji-» p a i'¥tis
CufpwJipitSid't* prospturtn**
Figure 75. Character 50 trace - pereopods 2 and 3, merus spatulate setae
261
Pseudotanais
Psetidatanais
Pseudetanais
Pseudotanais
Pseudotanais
P* eud<*tanai$
Pv e u d o l a it Jul.
P >i e « d o l j i is jr«,
Persopods 2 and 3 carpus
modified setae
(P) s patula
( P ) s p. O
(P)sp. P
(P) atfinis
(P) s caipellum
(P.) langi*. p i n u ^
( P ) 4 e i t l i c u i Jit: v
( P ) ^ piciifu*)
F <» <Mirf«> Li » J I ' , ( P ) VII.JJ.XI
P'i r u d o IA it Af*
P \ <Mido tanai*.
PJ, <;«s<i«M l*-3s«.
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pssudotartais
Pseudotanais
( P ) 5i ij»j» oisM'.ns
(P) iiMcmciicI*m
(PJsji.t
£P) l o n j j i s e t o s u s
(P) corollatus
(P) ocuiatus
(P) mexikolpns
(P) boreeai
(P) unicus
(P) mediterraneus
(P) inflates
( P ) s p. A
(P) vuiseita
Ps eud«>Lii>*m ( P ) I ^ I C I C U I J I I J
My** t u o c e n l i u ^ % # i u l ( i > ,
My*. I n o c e ntius >. J* A
P ( f « d t L i n j n ( P ) J bf», •, i
P* eHdota nam ( P ) b j s p i i i u u l i
P»> *>«d^3,i !6,n-;. ( P ) c a i t f o t n t ? ns • $
Pseudofanais (P) for«ipatu$
Pseudotanais (P)jjonesi
P s e u d o t a n a i s ( P ) colossus
P s e u d o t a n a i s (P) lilljeborjpi
P s e u d o t a n a i s (P) noderssfcioldii
Parapseudotanais abyssaiis
Parapseudotanais s p . A
r z i D P s e u d o t a n a i s (A) l o n g j p e s
[ i'—xiPseudotanais ( A ) s i e g i
r___;uPseudotan.ais (A) m a f e r o t h r i x
Z Z Z f t P f e u d o t a n a i f (A) m « r l e n * e n i
J
inP*. e u d « L i IIJIIV ( A ) g e i l a c f i i
prrnP«, «Miid« l* n a u ( A) « J«S'. •. i
_JLJP«, e u d o l a n a i i i (A)*«imili*
~ 0 P ' » *«idolanai<k (A) m a l a y e n t K
•nP*.«?udolanjii& ( A ) s c i a p p i
:
: u P s e u d o t a n a i s (A) fluiII«i
• ~ T ) C r y p t o e o p e abreviata
1
XiCrytoeopoides arcticus
np^Crftocopoides pacificus
I
u C r i r p t o c p o i d e s s p. A
r ~ o l u n g e n t i t a n a i s primrfanis
I I .rnParaiungentitanais longidigitatus
:ul_afitamais b e k l e m i s h e w i
ioCurtiehelia express a
Sinelobiis stanfordi
n i r i m Ha f i e r i * rap ax
^ ^ r a j T f p h l o ' t a n a i s parvus
no C a r p * a p u t tides pr«sp#c«t:n«s
E
Figure 76. Character 51 trace - pereopods 2 and 3, carpus blade-like setae
262
Pereopods 2 and 3 carpal
MMe setae length
C 3 half or lessfangthof prepeiiy®
than fiaJf length ot
Pse«d«taitais
Pseudotanais
Pseudotamais
Pseudotanais
Psetidotaitaiis
Pseudofanats
P*. * u d « t a f t « i b
P^ #<id»ta t*«i»*
(P) s patitia
(P) s p. O
(P) s p„ P
(P) mMinis
(P) s ealpellum
(P) 5«H<|i% (tSIMSS
(P) *lenfn;ui«il«<,
(P)vpicatut
P<> < > « < ! « ! j is J I » , ( P ) WJIJJIJS
r',<*?!<!«Lili^j-,
P«. «n«*iota nafi
P-seudotanacs
Pseud**tan,»is
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Psewdotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanaiit
(P) it ipintaicw,
(P) m jictof.h^le-%
(P) s p. C
(P) longis efejsws
(P) esorollatus
(P) ocuiatus
(P) mexikolpo s
(P) b o r c e a i
(P) unicus
(P) m e d t t e r r a n e u s
(P) inflatus
(P) s p. A
(P) v u l s e l l a
P-, *H«<l«lii ii j i ' i ( P ) liiS<:»(:«l^l J
My:, lto*M:e rtltu*:, s e n » i f u > ,
Mf, Utoctt-niiu:
\p
A
P.* • • • d o l l rs.ai<» ( P ) * f h y « . , M
P-> «nudof,ss».ais ( P ) b a t # 5 . n a o t a
P«.e«id«*t<*i*.ai<> (P) e a l r t o m i e n * is
Ps ewdotamais (P) foreipatMS
Psewdotartais ( P ) j o n e s i
P s e u d o t a n a i s (P) colonus
P s e u d o t a n a i s (P) i i l i j e b o r g i
P s e u d o t a n a i s (P) n o d e n s k i o l d i i
Parapseudotamais abyssaiis
Para p s e u d o t a n a i s s p . A
P s e w d o t a n a i s (A) i e n g i p e s
P s e u d o t a n a i s (A) 51*91
P s e u d o t a b e s (A) m a i t i a t h i m
Ps*Hidota«»ai\ <A) 10 o i l * m *r* 1
P v # t i d o l « i t A i t (A) g e i l a c h i
P'-. t*niiitljitjsi\
(A)', iraili',
P* * i i d a f a n , a i * (ft) sit <4 My^rt^ P.
Ps ipudotanai* (A) $ nrappi
P s e u d o t a n a i s (A) g u i l l e i
Cryptocope abreviata
Crytocopoides arctteus
Crytocopoides paeifficus
Cryptoepoides s p . A
l u n g e n t f t a n a i s primit¥us
Paraiungentitanais iongidigitatus
L a t i t a n a i s bektemisheiri
Curtieltuslia express a
S sue lob us s t a n f t t r d i
HMt§mtia
rap ax
Tjflph l a t i n * is parvus
C j n f « a p s t i i d « i pros p « stuns
Figure 77. Character 52 trace - pereopods 2 and 3, carpus blade-like setae length
Ftreopods 2 & 3
dactyius + unguis
E I
G23
•1
Hi
C?1
subequalto
S$3teSJ4®f
about 1^2 of
less than 1/2 of
equivocal
• Pseudotanais
wPseudotanais
• Pseudotanais
^nPseiidctaBais
tTjnP^etidciitaftiSis
(P)
(P)
(P)
(P)
(p)
s patuSa
s p. 0
s p. P
a If in is
s calpellum
a P ^ e u d o l a n J » « > ( P ) lttngi!> pittite
j-payiP^, <MS<{OLI n j i s ( P ) <S eikin:«Lti!i •:,
|>^JQP% eud«L>iijtiH ( P ) s p l e n i t i s
P ' , i» !( «! » 8.1 li Jl 11, <[P) V it J J ,11
2Bf* <. <*»ul<> ILi n j i ' • ( P ) n i p p <> is e c •>! >,
P >, t>lltjo{,l JUS', ( P ) lit J « SO cll4»l<>S
Psewdttlanais
Pseudotanais
Psieudotanais
Pssudotanais
BiPseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
P s e u d e t a nais
Pseudotanais
10 P s e u d a t a n a i t
detanai*
(P) ^ p. C
(P) l o n g k elosws
(P) corollatus
(P) oeulatus
(P) m e x i k o i p o s
(P) b c r c e a i
(P) aaicus
(P) m e d i t e r r a n e u s
(P) inflatus
(P) s p. A
(P) v u l s e l l a
(P) f a l c i c t i l a t a
rpuiMystHa«#iitius i c n a t a *
"tiziDMys liio«f>nliiiH t p A
P * e u d o t a n i m ( P > . t l n r , «, t
P', Purffl.l-Ui.ll>, ( P ) b JH>', ItJUfl
P s e u d o t a b e s (P) « * l i f o m i e n s i $
P s e u d o t a n a i s (P) forcipafus
Pseudotanais (P)joitesi
P s e u d o t a n a i s (P) colonus
P s e u d o t a n a i s (P) ( i l l j e b o r g i
P s e u d o t a n a i s (P) n o d e n s k i o l d i i
_-":rrParapseudotanais a b y s s a l i s
"j uPara p s e u d o t a n a i s s p , A
P s e u d o t a n a i s (A) l o n g i p e s
PseiiilufaiUiS ( A ) s i e n i
P s e u d a t a n a i ? (A) m a k r o t h r i x
P*. «(Hl*»Ll!l jt»i ( A ) ItlOltwIIVelH
P s e u d e f a n a i t (A) g « i l j i « h i
f*% e u d o y n j n - . (A) gjii-t 1 * i
P% f>udoliin<rii% ( A ) * i m i l i %
Pskeuriotanai* (A) malayens te
P s e u d e t a n a i s (A) s «.rapp>
2isP$ewd«*tanais (A) g u i l l e i
C lyp to © o p e a b revia fa
C i y t o c o p o i d e s arcticus
Crytocopsides paeiffieys
Cryptocpoides s p . A
l u n g e n t i t a n a i s priirtitwus
Paraiun gentitamais IoitgidigStatus
Latitanais b e k l e m i s h e v i
Curtieltelia express a
n m Sin g l o b u s s t a n f o r d i
yp hSoi.i ik,ii<, jtiiswM'i
i C j i p d J p 1 . e m i t " , pn>', jHf<:l(M»«
Figure 78. Character 53 trace - pereopods 2 and 3, dactyius + unguis length
264
Pereopod 4 basis
Pseudotanais
Pse«<J*tanais
Pseudtttanais
Pseudotanais
Ps eudtttoriass
length vs* width
i i < 4 x longer than broad
JJj 4 to 5 9 x longer than broad
{_ j >|Kform@rthan broad
P*
feMdtttaiiais
(P) spatula
( P ) s p. O
( P ) s p. P
(P) afjfimis
(P) s ealp*llum
( P ) 5tMi#!i, l>nili>.
i i P ^ eudol< r><iiv (P) denticuldtaitt
L ~ Q p t •ud#Lirt.4i<i ( P ) t p i c i i i i i ,
P* end«l<i n a n ( P ) *il|<a,a
Pv rudotai n JII% ( P ) mppom«ws
P s «nid« !:-i «»/•«•. ( P ) m a c r o « l i « l * s
P s e w d r t t a n a i S ( P ) S p. C
FseudGtanais
Pseudtttanais
Pseudotaitais
i g P s e u d o i a rials
Pseudotanais
•Qpseudttianass
xiPseudotanais
Pseudotanais
P^eudatanais
(P) Songis etosus
(P) corollatus
(P) ocuiatus
(P) mexikolpos
(P) borceai
( P ) tinieus
(P) medrteiraneus
(P) inflatus
( P ) *> p. A
P ^ e u 4 » t i ( i j i i { ( F ) vulsella
P i eudolJinai'S ( P ) i J l c i c u l a t J
Mys !!!«*;<'Bitns<> s e i r j i t u t
My«. ttioct* stlci", i p A
P* r u d « U w a i t ( P ) j i b y < t s i
Ps * u d o U nm^ ( P ) b < I I * ^ n a u t i
P<r « u r i o l i i n * i t ( P ) c a l i r o r n i c n s is
P s e u d o t a u a i s ( P ) l»scipatns
Pseudotanais ( P ) j o n e s i
Psssudotanais ( P ) coSonus
P s e u d o t a n a i s ( P ) Silijeborgi
n P s e u d o t a n a i s (P) nodenskioldii
__j~p3aPara p s e u d o t a n a i s a b y s s a l i s
•
t {MUM* P a r a p s e u d ota it a is s p. A
P s e u d o t a n a i s (A) l o n g i p e s
Ps^udotanais (AJsiegi
Ps euidtttasiais (A) matereihrix
P s e u d o t a n a i s (A) a t n l « » ' , u t i
P i e u d o f a n j i t (A) g e i i a c h i
"pSMSp >, 4ns»{w Li ii ji«. ( A ) g a u * t i
n_i_jjP% «nid«»Iartai<i (A) ^ u n i l i *
|j"rZDP*«Midofart,ai« (A) m a l a y * n i w
"~1' n P s e« do t a n a is (A) s crappt
: u P s e w d » l a « a i s (A) g u i l l e i
C r y p t o c e p e abrewiata
C r y t o c o p o i d e s arcticus
Crytocopoides pacifieus
Crypto op aides s p . A
ffpngtungentitanais
primrhrus
. T l f c i o P a r a i u n gerttitan ais I o n g i d ig Status
' ' - -ajLatitanais beltle m i s h ew
l o C u r t i c h e i i a express a
f—3oSin«!«bus s t a n f o r d i
L i r a a H a r g e i i a rap ax
i^Typhloljmjiis pjivui
CjSfptJijpf f U l d r s p«tt», p « c l n e *
Figure 79. Character 55 trace - pereopod 4, basis length vs. width
Fereopods 4 to 6 basis
setation
{...„. J sifppl* -ar j tvotmr $JS:M
M l
r-imjilr *-;r*iir
• M l no$$ii$
(P) s patwla
(P) s p . O
( P ) s p. P
((P
P )) a f f i n i s
(P) s M i p e i t u m
'BPseudotanais
nPseudotanais
,
:nPseudotanais
~ J j|
f—
- —- Q P s e u d o t a n a i s
J |—~i L ^ p | » s e u d t t t a urns
r
ajP«* e u d o L j i i a u ( P ) d t n l i c u l a t w s
i-rj n_™0 P * * U ** * ' J ™ *" ! * ^P) •'> PIC J i l l *
i iBBBBnP*1 »wdota n^ss ( P ) wifj^^i
5
—
- n P <s «n# d o >.a it .a 81 ( P ) n i p p « m c i i %
j-pn^ P •, «» 5B «i o M •> J i <. ( P )
LrrnPs'*"<•*** •»'"*
xiPseudotan-ats
I |B««F5 «!!<!<* t a n a isL^HjPseudotanais
TiPseudotanais
rux: P s e u d o t a n a i s
if—riPseydotanais
Pseudofanais
Pseudotanais
Pseudetanais
Pseudwtanais
:nP< frMdotasi-tis
j-fir:uMy% t u o c e n l i u i
1 '—f >My^ ttmc<» t t i n i s
m^cmchrlf^
( P ) J p. C
( P ) Songis e t w s u s
(P) corollaiiss
(P) sculatus
(P) mexlkolpos
(P) borceai
(P) unicus
(P) medtterraneus
(P.) i n f l a t e s
( P ) s p. A
(P) vulsella
(P) ( j k i c i i M y
^eiidluv
*p A
P', «»)i<l« !,i it JM", ( P ) j h y t i ' , (
P i e u d o t a nai-t ( P ) b a i i P M i a u t i
_j
- P
P^^eewudd«oi »
l t*;»i£ ( P ) c a l r f a i m e n s i s
d»1t a n A is ( P ) l o i c s p a J u s
. „ _ . : n P s e « r t » 1t a n a i s ( P ) j o n e s i
j P s e u d o t a n a i s (P) colonus
P s e u d o t a n a i s (P) lilljeborgi
P s e u d o t a n a i s (P) n o d e n s k i o i d i i
P a r a p s e u d o t a n a i s abyss alis
Parapseudotanais sp. A
P s e u d o t i s i a i s (A) l e n g i p e s
Ps«id«{aiiajs (A)siegi
P s e u d o t a n a i s (A) m a k s o t h r i x
P* e u d a l a iiai& (A)
modemeni
P i etidv tan«n* (A) it e r l a c h i
P S (>U<iob »*(••. (A) $..•«!•,', t
P', «Mtrf«S-i is *i s', ( A ^ m i i h t
Ps * u d » '*• na»* (A) m a lay«>i»<* w«
P ^ * u «io La n> A is (A) & cr J p p 1
a P s « M d « t « ! i » a i s (A) g u i l l e i
Cifpto<jwpe abreviata
Crytocopoides arctieus
JpaMCrytocopoides pacificus
i f a f i G r y p t f l e p o i d e s s p. A
l u n g e n t i t a n a i s prirrkihras
P a r a i t i m g c u n t f t a n a i s 10ragid ig Status
Latiianais beklemistteiri
gCurtichelia « ] f f e s s a
.lSinetabus stantordi
iJrrulldigeiia iapax
j 1 yp !ti«M jjJiii
pjiva^
:ot:4ip«jps«ii(i# i k pio% p««rlMS
Figure 80. Character 56 trace - pereopods 4 to 6, basis setation
266
Ptrecpds44 ischiumsetatlwi r |=!!!K::i:S»S UilaT
on* sstta
Pseudotanais
Pseudotaikais
Pveudatanaiv
P\*Midt»lai»iii*,
( P ) affiniis
(P) s ealpellum
( P ) ((Hints. p««su&
(P) deiiticuldlu*
P i « U « I O I « I I J I I S ( P ) s |>ICJIIUS
P t t U d o l J I I J I t ( P ) Vll|J/J
oP^i # w d o t j n j i K ( P ) n i p p o n i c u s
Pn « « r f « b n<ii« ( P ) in * « t o c h * l * *
Pseudotanais: ( P ) s p . C
Pseudotanais
Psesidofanais
Pseudotaitais
Pseudotarsais
Pseudotanals
Pseudotauais
Pseudatanais
Pseudotasiais
P&eadotanais
( P ) Iwngfe e t e s u s
( P ) corollatws
(P) oeulatus
(P) mexikolpos
(P) borceai
(P) unicus
(P) m e d i t s i r a n e u s
(P) inflates
(P)sp„A
Ps#udutaft.ai« (P) * U I « * * I S J
j r P & * u d v t 3 i u i v (P) tJleieuMt*
My% l i i o c e n l m t i « U 4 l « \
Mfiv I n n e r n l i u \ t p fk
P*v e u d n t i i n j io ( P ) b a r < > j n A u i
Pseadotanass- (P> c a l i t o i n i c n s i s
PsewdGtao-ais ( P ) f o t e i p a t a s
Pssudotauais (P)jouesi
Pseudotanais (P) colonus
P s e u d o t i n a i s (P) l i l l j e b o r g i
Pseudotanais (P) nodenskioldii
Parapseudotanais abyssalis
P a r a p s e u d otanass s p „ A
— I D P s e u d o t a i t a i s (A) i o n g i p e s
•mPseudotanais (A)siegi
P s e u d t t t a i i a i s (A) m • i k r o l h i i x
D P s e u d o t i i f t a i * . (A) n i o r t * n * « A i
Ps eudotAiirfiv (A) n«il,c*clti
P% t u d n ta l u i t (A)flj«s>,', s
Ps <*«rfo!,i ss»is', (A)%imih%
P ^ j n i d n l j i M i 1 . (A) nulAyiMio K
P s « « d « l i l 5»<1IS ( A ) & e i a p p i
P s e u d o t a n a i s (A) g u i l l e i
Cr#pto*j©pe a b r e v i a t a
C ryt© € © p o i d as a i c t i ciss
• = x i C r y t o c o p o i d e s pacsficus
^ • i C r y p t D c p o i d e s s p. A
lungentManaK primrtwis
P a r a ion g e n t i t a n a i s l o n g i d i g i t a t u s
Lalitaitais beklemishe¥i
Cortfehelia express a
Sine!«bus stanfttrdi
H a r j i f l i a sap AX
T y p i i l t t f a i M i s parvus
C Jip««ip;f t (jd#S ps «', p «f <:Iistf
Figure 81. Character 57 trace - pereopods 4 to 6, ischium setation
267
i%FfH9p#ds 4 to 6 mmmm
aetation
• •
nnr •;>-* j
t z n ! oqji i o n s
P s e u d o t a n a i s (P) s p a t u l a
P s e u d o t a n a i s ( P ) s p>„ O
P s e u d e t a n a l s ( P ) s p. P
Psteudotaitais (P) atfinis
p5n»«d»tai»,ais ( P ) s c a l p e l l t i m
P s e u d o t a a i M i * ( P ) l o n g is p i f t u s
P* e u d o t a n a i * (P) d e n t i c u l a h i *
P i V U d o t d l l . l l 1 ) ( P ) t pl«,allUV
P% r u d o l a n a i s ( P ) v i l j j j n
P •« «ME<l«f..i n ^ i \ (I*) n ipp « m c »
Ps HMid«ta it a i f ( P ) m «a«io «b«*l«»'i
P s e u d o t a n a i s (P) s p C
Pseud**isn*M% ( P ) I f f n s t s etos«is
Pstsudotanais (P) corollatus
P s e u d o t a n a i s (P) oculatus
s i P s e u d e t a n a i s (P) m e x i k o i p o s
P s e u d o t a n a i s (P)
feoreeal
P s e u d o t a n a i s (P) unieus
p P s e u d o t a n a i s (P) mediterrarseus
P s e u d o t a n a i s (P) inflatus
Pseudotanais ( P ) s p . A
P s e u d o t a n a i s (P) v u l s e l l a
P s e u d o t a n a i s ( P ) l»sN:H:islit!,i
My> t l i o c e n l i u * i e i u l u ^
Mf, t u o a o n l i u t i p A
P i eudolanaiii (P) iiby,-.1
Pn e u d o t a n a i s ( P ) baie<vna«iti
P s « d o i a p a i $ (P) c a l i f o i n i e n s i s
P s e u d o t a n a i s ( P ) forssipnttis
Pseudotanais (P)fonesi
P s e u d o t a n a i s (P) eolonus
P s e u d o t a n a i s (P) iilljeborgi
P s e u d o t a n a i s (P) nodensfcioldii
Para p s e u d o t a n a i s abyss alis
Para pseudotanais s p . A
P s e u d o t a n a i s (A) i o n g i p e s
Ps«ud»tanais (A)siegi
P s e u d o t a n a i s (A) sit j l t i o l f s o x .
P <t « u d *» ia it a it (A) m o l i e n $ e n i
P s e u d o t a n a i s (A) fl<,'si,it:tu
P',*'«!<!« ! J is .iB', (A) fl.ui". •, i
Ps w u d o l a n a i s (A) s m u l i s
P s o u d o l a n a i . s (A) m a l a y ^ n s ts
P s e u d o t a n a i s (A) s e s a p p i
P s e u d o t a n a i s (A) n u i I I e i
Cryptocope abteviata
n C r y l o e o p o i d e s aretlcus
Crftocopoides pacificus
Cryptoepoides s p . A
iungentitanais primihms
Paraiungentitanais loitgidigitafus
L a M a n a i s bek Is m i s hewi
Curtiehelia express J
S i n e lob us s t a n f o r d i .
H a r g e r i a uapMX
Yyphlotanaus paivu&
IP t M »po 4pN » u d r s p i o s p e e t n e s
Figure 82. Character 58 trace - pereopods 4 to 6, merus setation
Peraopods 4 - 6 carpus
modified setae
• •
Mario 'itoes f,»!t3i'i f'i-'pirt
Psetidotanais
Pseudotanais
Psetidotanais
Pseudotanais
Psetidotanais
(P)
(P)
(P)
(P)
(P)
spatula
s p. O
s p. P
a ifin is
s calpellum
P s v-udttiAtmi',
Pi, tM2<i*> i j ftds'i
P^iidoLiRjh
P •» «Mi*f«ti ti.ii 1 ,
P <. *'?irf« Li i u i ' i
( P ) Soitg is |»t» us
( P ) de»{iciii.ahiv
( P ) % i>it: 4it«s
( P ) *«ij*i/i
( P ) it ipp oiticii',
Ps pild« is mm
Pseudotanai*
Psetidotanais
Pseudntanais
Psetidotanais
Pseudotanais
Psetidotanais
Pseudotanais
Pseudotanais
Pseudotanais
Psetidotanais
P s e u tlota it a is
(P) n u t i t c h f l ^
(P) s p T
(P) l o n g i s e k s n s
(P) eoroltatus
(P) oculatus
(P) mexikolpos
(P) b o r e e a i
(P) u n i c u s
(P) meditterraneus
(P) inflatus
(P)sp.A
(P)¥u!s«*i!a
P«, e<8ti«1jl «>,!>', ( P ) f a i c n i u l d i j
M y . l i i u i i i t i i l i i f , », *•• u .»Iil>,
tAy\ Inoc** n t m t '. }* A
P •, tutdo S-i tsji-, ( P ) A b ^ t i
P % e t i d o t a n a i s ( P ) b a i * " i i».i!jh
P«i ftwdojtanass ( P ) C A l r f o m i e n s i s
P s e t i d o t a n a i s (P) f o i c i p a t u s
Psetidotamais ( P ) j # s i e s i
Psetidotanais (P) colonus
P s e u d o t a n a i s (P) l i l i j e b o r g i
Psetidotanais (P) nodenslcioldii
p u P a r a p s e y d o t a n a i s abyssaiis
" E i P a r a psetidotanais s p . A
P s e t i d o t a n a i s (A) I o n (j ip es
Pseudotanais ( A ) s i e g i
zr__ijP-seud»tanai;£ (A) snalfrolhritf
P i r u d o i a n a i & (A) in o fie ft * « n 3
nP•. c « d « SJ ti.jii'i (A) g (Ml^clki
jTZZUPi e i i d o l j it JI% (A)g«iu<>*. i
1_ uP * t*«do 1J »<ii« (A) % iiinh',
P % ?iid<»laitAI% ( A ) in ,t L i f p m , i*.
P i * M I r f o ! j 3ii^is (A) 5 a r a p p i
j o P s e u d o t a n a i s (A) g u i l l e i
iDCrfptocope abreviata
OCrytocopoides arcticus
^ ^ C r f t o c o p o i d e s pacificus
Lr3aCryptocpoid.es s p . A
~33tungeiititanais primitwis
L—nP a r a i n n g e n t i t a n ais Ion gid ig Status
L a t i t a n a i s belt le mis hetri
£iC si siieheSia e x p r e s s J
xt S i n * So fens s t a n f o t d i
L ~ n H a r u e i i a sap ax
_ J J fyp h l o l a i u i *
|*JI*?I',
rrzrot" J i p o j i p s e u d « ^ p i o * p t t r i n t s
Figure 83. Character 59 trace - pereopods 4 to 6, carpus blade-like setae
269
Fereopods 4 - 6 carps
additional modified §iti@
CZ3 &)fkl-tipp$<j
P = I equivocal
sewfJotanais (P) spatula
Pseudotanais
jPseudatariais
Pseudotanais
Pseodotastais
PJ> e u d o t a i t a i * .
P», t M i d w I j SI-IS1,
!'<, c m l i t L i i t J l ' .
(P)
(P)
(P)
(P)
(P)
(P)
(P)
s p. O
s p. P
affinis
s ealpeiium
Eoatgb. psiMis
<i »»iiii«; t i i a h i •.
<i p i c Mu*.
P "i I M I i l o i j IS..II*. ( P ) ¥ l i j « t / J
P<, «mcf o L i JI J I ? .
P s «*wdo1.a nit!*;
P s «tM<ioian«MS
P s «*Md«ta n a i s
Pseudotanais
Pseudotanais
Pseudotinais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Ps ettdofanat*
Ps e d d o l a i u i i ' i
j ' P ) it i j t p »5U<:SH,
(P) n t ^ c t n c h e l ^
(P) j f. C
( P ) Ior»g is e t » s u s
(P) coroltatus
(P) oculatws
(P) mexikolpos
(P) borceai
(P) unicus
( P ) m e d r i e r r a i t ems
(P) inflates
( P ) & p. A
(P) vulsella
P i , tMajiu ! j Hiii<, ( P ) l a l c i t i u l a t *
^ itiocentauv
seuatu*
v 111««: <» I I I H I •» », |t
A
P^ ( M i d o f a (i.n>) ( P ) . a b y t ' t i
P<; * » i i r f * i ! j n a i * ( P ) b an»s n a u l i
P s p i i d o t a n a i t ( P ) c a l i f o n t i e ttsis*
P&eudotiiiJis (P) foieipalus
Pseudotanais (P)jonesi
P s e u d o t a n a i s ( P ) coSoraus
Pseudotanais (P) lilljeborgj
Pseudotanais (P) nodenskioidii
Para pseudotanais abyssalis
Para pseudotanais s p . A
P s e u d o t a n a i s (A) lortgipies
Pseudotanais (A)siegi
P s e u d o t a n a i s (A) m a k r o t t s i i x
P i e i i d o t a i i j i i ' , ( A ) i n o r t e i i v * ssi
P •:• <»1!(1*|L| ! i J ! S ( A ) 1J * I E J C i t !
P«, t>(««!*> Li nj«i«, ( A ) g j u ' . v i
P ^ **3s«J« L i it jii% ( A ) s MII i l i t
P s c D i i o U n J i ^ ( A ) t i l . . « I j f t»!ii is.
P s < > i i i l # y n*«5 ( A ) % f . i ^ p f n
P s « u d o t a n a i s (A) g u i l l c i
CrTfljtoeope abrewiata
Crytocopoides arcticus
rijtfoGopoides p a c i f i c u s
Cr$ptocpoides s p . A
iiiisgentrtanais primitwus
Para inn jjentiian ais i o n g i d i g i t a t u s
L a t i t a n a i s b e k l e m i s it ewi
Curtiehelia express a
Sinelobus Stanford!
H a i f « i i a rap ax
Tfptti^tanilijf parvus
l* . l l p o a p ' i CMlde-, p f O S p f s t i t t t t f
Figure 84. Character 60 trace - pereopods 4 to 6, carpus additional specialized setae
1*7 f)
Psewdotastais
j o P s e u d o t a n aai is
s
3Pseyd©tanais
Psetidfltaraais
«d»tai*ais
P®m@p®$® 4to§ prapodu*
broom setae
(P) spatula
P )) :s p. O
((P
(P)sp. P
(P) a ft mis
(P) s e a l p e l l u m
„ D P A e i i d i t L i u a i S ( P ) i o t t y ib pht H^
SHU*'St*
J
y P ^ euduLdn^iV ( P ) d ettliutiialM i
n J ^ p P * * u d o LII»«U':, ( P ) «, |»it jii«s',
I — P
T^
ji|
!
'• r n d o 1-.M I>JI«. ( P ) v i l ; j v 3
p ^ »* ?s tl«l»t n ^ i". ( P ) n i p p o n i c o ^
j-paHgPs gudo l.g II JIS.
>
' ' *iidolanjiis
Pseudotanais
Pseudotanais
Pseudafamais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
P i eudotanais
Pa. «tidotanai>;
nP*. <Mi«lolii ii js-i
^ ™ ™ 1 ^ D '
j--_^t]|y|yS l l l O C e i l l l U *
( P ) m a c i o alt P I e.^
(P) s p C
( P ) long is elosus
(P) cornllatus
(P) oculatus
(P) mexikolpos
(P) bwrceai
( P ) urticus
( P ) m e d iter ranees
(P) inflates
(P) s p. A
(P)vdlsella
(P) 1jicscui.a1.a
^IIJIU',
1 *—.pMy'i l n o c w i i t m v *> p A
P>, tMadoLiii!jn, ( P ) . a b y * M
P ^ 4» <• <i o f^i it J ! ' , ( P - ) b <ai * i ii J I I h
Ps eudotanaisr (P) California n*i£
P s e u d o t a n a i s ( P ) fotcipalws
Pseudotanais ( P ) j o n e s i
P s e u d o t a n a i s (P) eoionus
P s e u d o t a n a i s (P) i i l l j e b o r g i
P s e u d o t a n a i s (P) n o d e n s k i o l d i i
Para p s e u d o t a n a i s abyss alls
Para p s e u d o t a n a i s s p . A
P s e u d o t a n a i s (A) l o n g i p e s
Pseudotanais ( A ) s i e y i
P s e u d o t a n a i s (A) m akrotfo i i x
Pi#Ullutlltjl'i
P», «??2 4l<> i»i it»n»,
P Si i ' l l l i * hi Sfc.iJ',
P«»iid»liiiuH
( A ) Hi O ihfll*. f i l l
( A ) g *-ii.iu:lss
|" A ) (( *IU », ':. J
( A ) « , imtit',
i>
P % **<ado!J u Js•, (A) in a tayenv «
Pseudotanaiv (A)&crappi
P s e u d o t a n a i s (A) g u i l l e i
C r f p t o e o p e abrewiata
• M C r f t o c o p o i d e s arciicus
..r,"~",nCrytwcftpoides p a c i f i c u s
~ L 3 0 C ryptocpoides s p. A
iungentrtanats primitvus
Paraiungeniitanais tongidigitaius
Lafitanais bek Eemishevi
Curtiuhelia e i f r e s s i
Sine lob us S t a n f o r d !
H a i g t r i a rapax
Typhi e l a n s is purwus
3 D C , a i p » i i p s t i i d « « pi»spa«rtn«-s
Figure 85. Character 61 trace - pereopods 4 to 6, propodus broom setae
Pemopods 4 - 6 dae%lus
C Z l fJifrJ X «i L'iobit 15 fOflrr ;sife
mPseudotanais (P)spatult
;
L. —nPseudotamais (P) s p. O
^3jPse«dotai»ais ( P ) s p. "*
-,L_jr — j - P s e u d o t a i s a i s ( P ) affinis
•—n P s e u tl«ta it a is (I*) s c a l p e i l u m
:=rx>P*eudot4n4rv ( P ) l o n g is pin us
!
P * eudotanais (P) denficulatus
^-1
|[-|r
^
J LlfC:
P". r n d s id ii.il', ( f * ) •> ps« .tits',
P «. *M!*lo Li S4 J ! ' , ( P ) «rt|4J9
3 V 5 «* it d a l a •« ii • \ ( P ) n i p p o n i C U f
J — Q P5 p u d o L a n a i *
1'—•nr-s.eiido'fcin.jiK
-T^^^PscMdotainais
j—jQPseudotanais
1 tH^Psewdotanais
——gpPseudotanais
j — i Pseudetanais
"I *~-riPseudotanais
^^jjjPseudotanais
j Ps e t i d o t a n a i s
jjPseudotanais
F
(P) m j c i o c h e l e s
(P) s p. <•
(P) ionjusctosus
( P ) ©»roIlatus
( P ) ocfulatus
(P) mexikolpos
(P) boreeai
(P) unlcus
(P) medtteri-aneus
(P) inflates
(P)sp. A
XJIP* e u d u t a n a i s ( P ) v u l s e l l a
T i P v e u d o t a n a i * (P) f a l e i c u l a t *
I ajM**" l i i o c e n l i M * ^ e i i a t u t
T.—oM*'" l n o c e n l i m s& A
n
in
P t <Hidot,aitait» ( P ) j i b y t ^ i
znV*< ^ M d n t a n a i * ( P ) b j i t M i j « l i
P-sewdralanaix ( P ) c a l r f w r n i « n s i s
_j.
^ x j P s e u d o t a n a i s ( P ) forespatus
-TiPseudetauais ( P ) j o n e s i
n 8 -—••—•••uPsaudotanais ( P ) coioisus
I'—-"
:pPseudotanais (P) lilljeborgi
-—17—7:—ixjPseudotanais ( P ) n o d e n s k i o l d i i
_ nzzaaParapseudotanais
abyssalis
Para p s e u d o t a n a i s s p . A
_ j — 3 ] P s e u d o t a n a i s (A) l o n g i p e s
nJ^pPseudotanais (A)siegi
*— 1. P ^ e u d u t a n a i s (A) makroHiiriy
&
- Q P * «udo t a n a i * (A) in a itens e m
TiPteudolaftMh (A) g « i l 4 « h
.HI-, t i
A ) fl$<*«<**«
j - — J J P V »ndulanai% ((A)
- 1 ' --PsiMidolan
(A) •. imili*.
P t e v d i t t a n a i * (A) m a I* yens, K
J"T1 P s * u d o l a n j i s (A) s c r a p p i
ajPsRMdwtanais (A) g u i l l e i
• Cryptocope abreviata
HiCrytocopoides arcticus
HiC ryfcocopoidsss p aeificus
• Crvptocpcpides s p . A
t u n g e n t i t a n a i s primitvus
P araiun gentiianais lungidigitaius
L a l i t a n a i s belclesmishevi
C urtlchelia e x p t t s s ^
30 S in ie lo b us S t a n f o r d !
Jnu
rc
" " " L E J D H a r g * >sj tap ax
• • • • • C a f p # a i p S t < l d * S p i « ' , p<M:ilMf«,
Figure 86. Character 62 trace - pereopods 4 to 6, dactylus fusion with unguis
272
Pleopod complexity
CZ3 rami well
O rami
terminal * Mural setae
only
Pseudotanais
Pseudotanais
Psesideta rials
Psettdatanais
Pseudotanais
(P) s p a t u l a
(P) s p. O
(P) s p. P
(P) affints
(P) s calpeilum
P \ * u d o t a n J i \ (P) d«htit:ulAtiiv
P«i • udoLm^is (P) <i picatu*
Pt • u d o b n . i i t (P) viljasi
P* (Hidetaiiim (P) m p p o n i c u *
P<« <»ud<»t*fiai* (P) m«»ci«*«:S»«»l*K»
Pi, «Mdot*nai<( ( P ) s p . C
P s e « d ® t j i u i s (P) long is etesns
P s e u d o t a n a i s (P) corollatus
P s e u d o t a n a i s (P) oeulatus
Pseudotanais (P) mescikolpos
Pseudotanais (P) borceai
Pseudotanais (P) unicus
Pseudotanais (P) mediterraneus
Pseudotanais (P) inflatus
P s e u d o t a n a i s (P) s p. A
P s e u d o t a n a i s (P) vulsella
Pseudotanais, (P) laleiculJita
My* t n o e e n t i u t •, e j i ^ l u * .
Myv ln**c<*nliu% '. p A
P v # u d a t a i M i \ (P) J bf> «> i
Ps e u d o t a n j i * (P) b<ii^sn««h
Pseudetianats (P) c ,ilitoin•« n\ is
Pseudotanais (P) torcipatws
Pseudotanais (P)jomesi
P s e u d o t a n a i s (P) colonus
Pseudotanais (P) liltjeborgi
Pseudotanais (P) n o d e n s k i o l d i i
M m Para pseudotanais abyss alis
I I M B Para p s e u d o t a n a i s s p . A
Pseudotanais (A) longipes
Pseudotanais ( A ) s i e g i
Pseudotanais (A) m a k i o t h f i x
P<icuil«tindi<i (A) in (tit e n * r i n
P>, <?«<i»biMt>, ( A ) g e t l a c h i
P i F U 4 O | J n « I I \ (fi) ( . H i ' , - , i
Ps #udobn l ii<. (A) >, HISs!i*>
Pi> <»iid«lanais ( A ) m j i U y # n ^ i i i
Psi!iiii»faniii , » (A) 5 ctappi
Pseudotanais. (A) g u i l l e i
Cryptoeope afeieviata
Crytosopoides areticus
Crytocopoides paoifieus
CCryptocpoides
s p. A
ungentitanais primitvus
J
Paraiungeittitanais iongidigitatus
Latitanais beklemishe¥i
D Curtiehelia express a
r—nSinteloltus s t a n f o r d i
d~D H a i g « t n j l a p a x
U Jyphlotanais p4iwu%
33 C <*ipo,*p* » u d n p»»*. p * c t n « $
Figure 87. Character 63 trace - pleopod complexity
273
Uropod exopcd
number of articles
> Q Pse«d<*tanais
iOPseudotanais
Pseudotanais
^ L _ T f z : u PPss««!:U«ddo»ttaann aa ii ss
r
1 .oPsewdotanais
jr==nPseu»l*»faftaiis
.,1 IJJP v e u d o t a ii a is
> I—["LZZOP^ »*udola n a i s
'-M L^rr~rnoP*. e u d o l a n a i s
rrsoPt p u d o l a n a i t
J-~2QPS ^ u d n t a n a r i i
li—nPsewdaiaitJiJs
Pseudotanais
j , :yPse*id«tanais
lizzoPseudotanais
Pseudotanais
rpnPsetidotanais
(P) s p a t u l a
(P) s p. O
(P ) s p. P
(P
liftis
(P)) aj fflints
(P) s ttatpeltum
(P ) l o n g is pin u«
(P ) 4 entic Hiatus
( P ) s piealw*
( P ) wiljj-n
(P) mppt»m<:u%
(P ) maeto«h*»|it>\
(|»j ^ p_ c
(P) longv> e t a s u s
(P) e o r o l l a t u s
(P) o c u l a t u s
(P) mexikolpos
(P) b o r e e a i
p P s e u d o t a n a i s (P ) m e d r l e r r a n e u s
pzoPs eudotanais (P) inflates
l ^ n l — x i p f eudotanais ( P ) s p . A
I 'f "JbrZZDPsftudotaiiai*, (P) v u l s e l l a
-—'• n . ' ™ . ... inP« e u d o t a n a i s (P ) faicKSulafa
]I
. rpmfctys I n o c e i t f i n s s p i i a l u i
l i z : D M ^ iiiHtccnltiis ^ p A
•!0P^ i M i d o l a n a i * ( P ) j b ^ u
aaP*i p u d o t i i nais (PJ b a n * * iiautJ
P<. ( > i ! ( i o l j n j r » (P) c a l i f o r n i u m s is
P s e u d o t a n a i s (P) f o r e i p a t u s
TiPsewdotanais ( P ) j o n e s i
Ps«swd»tamais (P) colomus
I B P s e u d o t a n a i s (P ) l i l l j e b o r g i
B P s e u d o t a n a i s (P) n o d e n s k i o l d i i
„„rp^nParapseudotanais abyssaiis
l_i_juParapseucltttanais s p . A
P s e u d o t a n a i s (A) l e n g i p e s
^ L b r n P s e u d o t a n a i s (A) s i e g i
M—; • • • • • p i s e u d o t a n a i s (A) m a k i o t h r i x
" I Q P s e u d o t a n a i v (A) m o l t e n s ' e n i
' n p t t > t i ( i i i i j s u f . ((A) a e i t a c i n
P% » u d o l a n a i * ( A ) g a u * t i
P s * u d o f a n a i % (A) % tm t i l t
Ps> e u d o t a n a i s (A) m a lavens K
1.i"~:nps e u d o t a n a i s ( A ) s c » a p p i
^ D P s e u d o t a n a i s (A) g u i l l e i
C i y p t e c o p e abrewiala
^ C r y t o c o p u i d e s arcticus
,.i i—^ n Crftocopoides pacificjus
- U C r j p t o c p n i d e s s p. A
l u n g e n t i t a n a is p r i m r t v u s
[ t — n P a r a i u i t § e n t i t a n a i s I o n gidigstatus
>o |_ an"ta n a is b ak le m is h e n
I
DCurtiehelia express a
Sineiobus stanfordi
H a r g ^ r i a rap ax
Typ biota in a is parvus
ioCarp*apstnd#$ pi**spt«tn«
Figure 88. Character 65 trace - uropod, number of exopod articles
274
Uropod exopod
length vs. endopod
? i i Bh»*ri 1,13 |$n«31
seudotanais
seudotanais
seydotanais
setidotamais
seudotanais
semfoijHjjv
seudolanais
^ eudo i j ii,i»'.
i t'H(!«ljIIJI»,
*» fudoS^ n j »
5 eudoian.aiv
seudotanais
Ps«udoti«nais
seudotanais
seudotanais
Pseudotarsais
seudotanais
seudotanais
Pseudotanais
seudotanais
seudotanais
seudotanais
s ««ido1jnai<ik
(P)
(P)
(P)
(P)
(P)
(P)
(P)
(P)
spatula
s p. O
s p. P
affinis
s calpellum
iony is pin u-%
d enticul«ilu£
^ fn<: jittf;
( P )
tfllj^JIl
(I*) mpponicti'i
(P) in .K-.tocbel*'*
(P) •* p C.
(P) i o t i g k et«sus
(P) coroliatus
(P) ©cuSatus
(P) mexikolpos
(P) boroeai
(P) unieus
(P) mediterraneus
(P) inflatus
(P)sp.A
(P) wulsrella
(P) fa I *i« «lata
1 tllOCtt IlliUV
^ f J! jIlB':,
\ l n o c r nliu* *. p A
<it>ud«iljnji<i (P)<nby. * •
<;i»«(j«tjn^is (P) calriornie n* i<^
s-eudotanais- (P) forcipatus
seudotanais (P) j o n e s i
seudotanais (P) eslonus
seudotanais (P) liiljeborjji
Pseudotanais (P) nodenskioidii
ara pseudotanais abyss a lis
arapseudotanais s p . A
seudotanais (A) longipes
seudotanais ( A J s i e g i
seudotanais (A) tnakiotftfix
* eudo !<* (Ui'i (A) in o (tens * eti
«. «*«<{*> Lt sid t«. (A) geiiaehi
s e i i d o t j i u i ^ (A) ^| ,(<i'. <, 5
\ pudo Li it JH\ (A) ^ mi ihs
S * U d o l . * IIJII5 ( A ) l l k . i l l j y * l l « IS
s pudota nais (A) s cMjtpi
Pseudotanais (A) Qwillei
lC i"*/ptoc o p e abfftvia feral
Crftooopoides aretieus
rytocopoides pacifieus
ryptoopoides s p, A
lungeittitanais primihms
Para inn gentitanais Ion gidig Status
Lalitanais beklemish ewi
Ciirtiehelia e g r e s s a
, — Sine lob us * t a n f o i d i
~~L~a3Haiflieiia lapax
Typliloti«ii4i> f»*ii«u«,
IDt'aipojips *udi*s
Figure 89. Character 66 trace - uropod, exopod length vs. endopod length
piatptetnftS
s e u d o t a n a i s (P) s p a t u l a
seutietanais (P) s p. O
Pseudotamais (P) s p. P
s e u d o t a n a i s (P) affinis
setidataftais (P) s ealpellwm
* eudtttaiiais (P) latig is inn u<i
% «ml»laKJlv (P) d«»!i« til^his
Uropod mmp®4
proximal article
r; I about 1M length «#
aia 1f3to1/2NMfigthof
2l3J6ngtttef
«. <ftl<l<> t j H . I I ' ,
* (tudotauar?
% «udotjft4li>
P*4 w u d o t j i M i N
Pi, <»u<iobi it-ii".
mmtwgTMtrawrffCTgwtwmwKnw^
< ( * ) > . JUt j S l d
^, «M«d»tenai'i (P) wrtj^^
% e»i<l«laiii.i«<i (P) it ippotticu<;
% <Mid<*t»«ai* (P) »it .aGiocSt**!^
se««l«t>iit»jpi', (P) j p, ('.
Pseudotanats (P) IwngK elwsws
s e u d o t a n a i s (P) corollatus
s e u d c t a n a i s (P) ocuiatus
s e u d o t a n a i s (P) mextleolpos
P s a u d o t a n a i s (P) b o r c e a i
s e u d e t a n a i s (P) unicus
iPsewdotanais (P) m e d i t e r r a n e u s
seudwtanais (P) inflatas
s«ud«tanai£ (P) s p. A
s*ud«»tattdis (P) v u l s e l l a
stsiidotanais (P) f a l c i c u l a t *
ruLE Mys lti«c*nliuitr w i u l i i < i
LBnMys lnoc«t ntiu* •. p A
«<>u<loLin4iti (P)Aby*<%i
s e u d o t a n a i t (P) k a i t " . n a u l i
I"seiid(tl*i!Mn (PJ aalit<»ini«n£i$
Ps€«d»tar»iiis (P) forcipatus
seudotanais ( P ) j o n e s i
Psetid#tanais (P) colonus
s e u d o t a n a i s (P) !i!ljeborf|i
seudtttanais (P) n o d e n s k i o l d i i
arapseudotanais abyss alas
arapseydotamais s p . A
P s e u d « t a n a i s (A) i o n g i p e s
seudotanais ( A ) s i e g i
Pseudtttanais (A) rai Jkiti111 ssx
C
(A) m o i t e n ^ e i i i
( A ) g«Hl,d<:iti
(A)g«»Wksi
(A) s i m i l i *
5 e m d ^ f c u t ^ i s ( A ) «. c t ^ j t j t i
:P&eiido1jnais ( A ) g u i l l e i
Cryptocwpe abreviata
Crylocopoldes arctieus
rytncopoides pacificus
ryp toe p a i d e s s p . A
ungentitanais primitwus
pp ara i u n ge is t i t a n a i s I o n g i d i g i t a t u s
L a f i t a n a i s belt le m i s h e v i
iCurtietieiia e i f i e s s a
S i n e l o b us S t a n f o r d !
H « r a t r i a tap ax
Typ h 19ta tt a ME pJifWUf
nnCatpiftjps « n d t f pt esp tetntts
Figure 90. Character 67 trace - uropod, exopod proximal article proportion of ramus
Uropod endopod
articles
L '..J mora than 2 articles
E 3 2 articles
t i l l 1 + pseudo-articulation
• H 1 1 article
P s e u d o t a n a i s (P) s p a t u l a
P s e u d o t a n a i s (P) s p. O
P s e u d o t a n a i s ( P ) s p. P
P s e u d o t a n a i s ( P ) a Iff i n is
P s e u d o t a n a i s (P) s calpeiiurfi
P s e u d o t a n a i s (P) longis pntus
P ^ e u d o f a i i a i s ( P ) d e«s t i c it 14 h i s
P-, 1» M *J U Sii 8S *t I'» ( P ) «, |>H: .jfli!',
P •. *•• ?nl«> L« s>.11•, ( P ) v i t g a x i
P ^ r - t i d o i a SSJI«, ( P ) n i p p o n i c i i t
P*;. p u d o t a n a i s ( P ) i n a c t a «!*<»•>»
Ps««dt»tjiBjK
(P)sp.C
P s e u d o t a n a i s (P) i o n a k closes
P s e u d o t a n a i s (P) eorollatus
P s e u d o t a n a i s (P) oculatus
P s e u d o t a n a i s (P) mexikolpos
P s e u d o t a n a i s (P) boreeai
P s e u d o t a n a i s (P) unicus
P s e u d o t a n a i s (P) medrterraneus
P s e u d o t a n a i s (P) inflatus
P s e u d o t a n a i s ( P ) s p. A
P s e t i d « t d i U i 5 ( F ) ¥«SseSii»
P*. e t i d o t a Btaii, ( P ) f a i t i i t t u i a t a
M y , Si«<><:«• n S H I 1. s e n a t u t
Mt/<i fttoctt ului«, t p
A
P% <*?N:IoLi «s Ji% ( P ) ,i by*, •; i
P *, t M U l o l i l ll^I!', ( P ) b i l l ( " . Si al« tl
P s e u d o l i i n a i s ( P ) c a l r f o r m e n s is
P s e u d o t a n a i s (P) foitupatus
P s e u d o t a n a i s (P) j o n e s i
P s e u d o t a n a i s (P) coionus
P s e u d o t a n a i s (P) Hlljeborgi
P s e u d o t a n a i s (P) nodenskiofdis
Para pseudotanais abyssalis
Parapseudotanais sp. A
P s e u d o t a n a i s (A) l o n g i p e s
P s e u d o t a n a i s (A) s i e u i
P s e u d o t a n a i s (A) m a k r o l h i i x
P s e u d o t a n a i s ( A ) in a • t e n * e n i
P*. e M d o t r f u a i * , ( A ) $ 4*i!,n:Sii
P«i « < l d o U It J ! S ( A ) fl »i!l", ». «
Vs ( M i d o f a »«!!', ( A ) % eiftch*
P<» <mdo ta n•*««. ( / ^ n i t i M y o i i M j i
Pseudotanais
(A)sc*appi
P s e u d o t a n a i s (A) n u i l l e i
Crytocopoides arcticus
Crytocopoides pacifious
Cryptocpoides s p . A
l u n g e n t f t a n a is p r i m t t w u s
P a r a i u n g e m t t t a n a i s I o n gidi*jStatus
Latitanais beklemishewi
Curtieheiia e i f r e s s a
S i n e lob us S t a n f o r d )
H a i g e i i a rap ax
I yp b i o t a it aiA p a i v u ^
II a i p o jap* «f u d e s p i «•% p «»ei»ei
Figure 91. Character 68 trace - uropod, number of endopod articles
277
proximal article
L J lMQtoi/4l«ngth«f
• • 1/2 to 2/3 length of
S 3 equivocal
Pssudfttanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
PS*MHi**LiiiJ!V
P» * u j t i l a rt J I V
P*. t n d o l i i i j n
Ps f i i d » U i u n
P% «ntdo La iMi%
P s «MI d o t a r t a r
P s «md«f.a nais
(P)spatula
(P) s p, O
( P ) s p. P
(P) affinis
(P) « eai|»^lium
( P ) lOnglS jHfiMV
(P) dentieul^fu'g
( P ) 4 p i c 41MS
( P ) wiljjjai
(P) m p p anicus
( P ) m.r)ci0ehi>i«$
( P ) s> [» C
Psewdotanats
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotanais
Pseudotamaii!.P-ceudotanais
Pteud«t»ft4i«
(P) Joojpfc etesws
(P) c o i o l l a t u s
(P) o c u l a t u s
(P) m e x i k o l p o s
(P) b o r c e a i
(P) unicus
^ p j m e d Iter ran ens
(P) i n f i a i u s
(P)sp.A
(P) tftilssila
(P) talciewlata
Ufi I l l O C f e l t t l l l S % « l l 4 t U t
M y % t l l 0 e # l t t i u % •, }> A
P«. f M i d o Li «,«>•. ( P ) ,i by*, % i
P% «•<«*!<> t i nai.% ( P )
b4t*<Li»4Mti
P & « ud o t* n 4 •& (P) tMliforitutns is
P s e u d o t a n a i s (P) foreipatus
Pseudotanais ( P ) j o n e s i
P s e u d o t a n a i s (P) c o l o n u s
P s e u d o t a n a i s (P) l i l l j e b o r g i
P s e u d o t a n a i s (P) n o d e n s k i o l d i i
Para p s e u d o t a n a i s a b y s s a l i s
Para p s e u d o t a n a i s s p . A
P s e u d o t a n a i s (A) Eongipes
Pseudotanais (A)si*s0i
P s e u d o t a n a i s (A) m a l r r o t h t i x
P s e u d o l a n a i * (A) m a rte n* # a«
P* • u d o t a n a t * (A) jjwil^eliJ
P%ttudal4it4«%- (^) g<insvi
P% eudata is41% (A)%imili<,
P S * l l d o 1 U n 4 l . * (A) m4layf>lt* I*.
P& *lld<»t*nai% (A)%-CI4p|t|
P s e u d o t a n a i s (A) n u i l l e i
Cryptoeope abreviata
C r f t o e o p o i d e s areticus
Crytocopoides paeificus
C r f p t o c p o i d e s s p. A
lurtgentitanais primitvus
Paraiungentitanais longidigitatus
L a titan a is b e k i e m i s h e w i
Curtieheiia e x p i ' M s i
saSinelobus s i a n f o r d i
II4 rgeria rap ,ix
Typ b l o t * It J i f (Mi'Wis
ffmwtfrffmTmrtgffwtrfffwttitiiwrtwmmrwrrrTwreTiiTOWft'riTf
C l i f p « l i p S t W d « « pr»$p#efl!)«$
Figure 92. Character 69 trace - uropod endopod proximal article proportion of ramus
278
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