Org. Divers. Evol. 2, Electr. Suppl.

6:1-19 (2002)
© Gesellschaft für Biologische Systematik
http://senckenberg.de/odes/02-06.htm

Hyalella armata (Crustacea, Amphipoda, Hyalellidae) and the description of a

related new species from Lake Titicaca
Exequiel R. González1, Charles O. Coleman2
1
Depto. de Biología Marina, Universidad Católica del Norte, Casilla 117, Coquimbo, Chile, egonzale@ucn.cl
2
Museum für Naturkunde Berlin, Humboldt-Universität, Invalidenstraße 43, D-10115 Berlin, Germany;
e-mail: oliver.coleman@rz.hu-berlin.de

Received 13 March 2002 ⋅ Accepted 12 August 2002

Abstract
Hyalella armata (Faxon, 1876) was considered for a long time to be a monotypic species. The peculiar morphology, very distinct from other
species in Lake Titicaca, was not analysed in detail after the original description. However, the present authors’ study of an extensive col-
lection from The Natural History Museum in London has revealed two morphologically distinct species. We herein redescribe H. armata and
describe the new species H. longispina.

Key words: Crustacea, Amphipoda, Hyalella, Lake Titicaca, taxonomy

Introduction Material and methods

The distinctive fauna of amphipod crustaceans of the
genus Hyalella Smith, 1874 in Lake Titicaca has
been extensively sampled, but not properly studied.
The first work by Faxon (1876), based on the
material collected by A. Agassiz and S. W.
Garthman in 1875, identified seven endemic and one
not endemic species. Later, Chevreux (1904, 1907)
described four species collected by Dr. Neveu-
Lemaire during the Mission de Crequi Montfort et
Senegal de la Grange to Lake Titicaca. Chevreux
also redescribed two of Faxon’s species. Two more
major sampling efforts were made in the Lake. In
1937, G. I. Crawford collected extensively in the
Lake during the Percy Sladen Trust Expedition and
deposited the material of about 20,000 specimens in
the Natural History Museum of London. More than
15,000 specimens were collected by C. Dejoux of
ORSTOM, France (Dejoux 1992). The present study
deals with two of the spiny forms of Hyalella
present in the Lake, H. armata (Faxon, 1876) and
the new species H. longispina.
The following material was studied (see Fig. 14):
Hyalella armata
From the collection of C. Dejoux: Lago Menor
(material illustrated), Gulf of Puno, Gulf of Desa-
guadero, Chuquito, Juli, between Taquili and
Amantane. From the Crawford collection, Natural
History Museum, London (numbers in brackets =
specimens): Coata Bay, GIC 253, 23 m (196); Coata
Bay, GIC 286, 12-19m (15); Coata Bay, GIC 275,
30m (30); Coata Bay, GIC 296, 19m (12); Coata
Bay, GIC 321, 1m (1); Coata Bay, GIC 283, 12-15m
(4); Coata Bay, GIC 304, 23m (19); Coata Bay, GIC
268, 30 m (16); Coata Bay, GIC 291, 19m (201);
Coata Bay, GIC 263, 30 m (204); Coata Bay, GIC
160a (2); Coata Bay, GIC 282 (1); Isla Suana, P. 46,
GIC 993/1a 14.2-14.6m (3); without location, GIC
170a, 6m (1), GIC 160a (2).
Material deposited in the Museum of Compara-
tive Zoology (Harvard University) by A. Agassiz
and S. W. Garthman, and identified by Faxon, was
also examined.

Org. Divers. Evol. 2, Electr. Suppl. 6 (2002)

González & Coleman: Hyalella armata (Faxon) and H. longispina n. sp.
Hyalella longispina n. sp.
Holotype male, 10 mm; PERU, Lake Titicaca,
Choccocoyo Bay, St. P 6, GIC 481, 11-23 m, leg.
Crawford, 25.VI.1937. Paratypes (collecting data as
holotype): allotype female, 9 mm; 106 males, 6-
10mm; 93 females, 5-10 mm. All deposited at The
Natural History Museum, London (holotype:
2002.311; allotype: 2002.312; other paratypes:
2002.313-322). Additional material (in brackets =
number of specimens) from the Crawford collection,
Natural History Museum, London: Taman, GIC
1327, 36-38 m (7); Taman, GIC 1323, St. P67, 56-
61 m (29); Taman, GIC 256/3, 15-30m (3); Taman,
GIC 1325/1 St. P67, 56-61 m (3); Patón, GIC 850,
St. P. 19, 25-34m (5); Patón, GIC 843 P. 17, 25-34m
(5); Siripata Bay, GIC 1005/1 P52, 12.3-16.2m (21);
Molinopampa, GIC 881, P. 24,19-27m (1);
Molinopampa, GIC 884, P. 28, 25m (8);
Molinopampa, GIC 890, P. 28, 25m (4);
Ancoraimes, P. 20, GIC 854/1, 12-24m (3);
Ancoraimes, P20, GIC 855/2, 12-24m (1);
Capachica, GIC 333, 20m (3), Choccocoyo, GIG
468 (478?) (5).
The animals were transferred into glycerol and
drawn with a camera lucida on a Leica Wild M8
dissecting microscope. Specimens were dissected,
appendages and mouthparts were transferred onto
slides in glycerol and drawn under a Leica DMLB
light microscope using a camera lucida. Morphome-
tric measurements were taken with a dissecting
microscope and a camera lucida calibrated with a
stage micrometer. The results were calculated and
plotted with Microsoft Excel using the linear
regression mode.
Systematics
Hyalella armata (Faxon, 1876) (Figs 1-5)
Allorchestes armatus Faxon, 1876: 364–367, figs 1-
18.
Hyalella armata – Stebbing (1888: 455); Della Valle
(1893: 514-515, tav. 58, figs 2-3); Stebbing (1906:
577); Barnard & Barnard (1983: 708).
Hyalella (Austrohyalella) armata – Bousfield (1996:
188).
Type material. Museum of Comparative Zoology,
Harvard University, Cambridge, Mass.
Type locality. Achacache, Lake Titicaca, BOLIVIA.
2
Description of male. Size 9.5 mm. Body with a
posterior dorsal transverse ridge on every segment
(Fig. 1a). Epimeral plate 1 round, 2 straight, 3
acuminate (Fig. 1b).
Head smaller than first two thoracic segments,
typically gammaridean, rostrum absent. Eyes pig-
mented, medium, round, located behind insertion of
antenna 1.
Antenna 1 (Fig. 4d) shorter than antenna 2,
slightly longer than peduncle of antenna 2; peduncle
longer than head, articles 1 and 2 subequal in length,
article 3 shorter than article 1, and same length as
article 2; flagellum of 12 articles, same as peduncle,
basal article not elongated.
Antenna 2 (Fig. 4e) less than half body length;
peduncle slender, longer than head, article 4 shorter
than article 5, setal groups on article 4 and 5
abundant; flagellum of 13 articles, longer than
article 5, basal article elongated. Upper lip (labrum)
wider than long, entire, ventral margin slightly
rounded to truncate.
Basic amphipodan mandible (in the sense of
Watling 1993); incisor toothed (Fig. 4g); left lacinia
mobilis with five teeth; setae row on left mandible
with three main setae without accessory setae, right
mandible with two main setae without accessory
setae; molar large, cylindrical and triturative,
accessory seta absent. Lower lip (Fig. 4c) outer
lobes rounded without notches or excavations,
mandibular projection of outer lobes round.
Maxilla 1 (Fig. 2a, c) palp uniarticulate, longer
than wide, reaching more than half the distance
between base of palp and tip of setae on outer plate,
distal setae strong; inner plate slender, smaller than
outer plate, with two strong and pappose apical
setae; outer plate with nine stout and serrate setae.
Maxilla 2 (Fig. 3b) inner plate subequal in length
and width to outer plate, one strong pappose seta on
inner margin, outer and inner plates with abundant
setules.
Maxilliped (Fig. 3a) inner plates apically trunca-
ted, with three conate setae, pappose setae apically
and medially; outer plates larger than inner plates,
apically truncated, apical, medial and facial setae
simple; palp longer than outer plate, four articles;
article 2 wider than long, medial margin with long
simple setae; article 3 outer distal face with several
long simple setae, inner distal face with long
plumose setae, inner distal margin with long setae,
outer margin with one or two strong and long
plumose setae; dactylus unguiform, shorter than
third article, distal setae simple and shorter than nail,
inner border without setae, distal nail present.
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González & Coleman: Hyalella armata (Faxon) and H. longispina n. sp.
Coxae 1 to 4 increasing in size, apically
acuminate (Fig. 1a). Coxa 4 (Fig. 3e) much deeper
than 1 to 3 and at right angle to sagittal plane of
body, wing-like. Coxa 1 subequal to 2 and 3. Coxa 3
(Fig. 4b) narrower than 4. Coxa 4 deeper than wide,
excavated posteriorly. Coxa 5 (Fig. 3f) posterior
lobe deeper than anterior. Coxa 6 anterior lobe
small.
Gnathopod 1 (Fig. 1d) subchelate; carpus wider
than long, subequal in length and as wide as
propodus, with strong and wide posterior lobe,
produced and forming a scoop-like structure, open to
the inside, inner face with four to seven pappose
setae, border pectinate and with several pappose
setae; propodus (Fig. 4a) length less than two times
maximum width (quadrangular), hatchet-shaped,
with no setae on anterior border, inner face with
more than ten pappose setae, without small
triangular setae, distoposterior and distoanterior
borders without setose scales, palm slope slightly
oblique, margin convex, posterior distal corner with
robust setae; dactylus claw-like, no setae on inner
curvature, congruent with palm.
Gnathopod 2 (Fig. 1c) subchelate; basis hind
margin with group of seven or more setae; merus
with less than seven setae on posterior margin,
postero-distal margin straight, distal corner rounded;
carpus posterior lobe elongate, produced between
merus and propodus, border pectinate with several
pappose setae; propodus triangular, distoposterior
and distoanterior borders without setose scales, palm
longer than posterior margin, slope oblique, margin
straight, several medium-size setae, anterior edge
with a small process, posterior distal corner with
weak setae, and with change of slope; dactylus claw-
like, congruent with palm, no setae on inner
curvature.
Peraeopods 3 to 7 simple. Peraeopods 3 and 4
merus and carpus posterior margins each with four
hind marginal clusters of long setae; propodus
posterior margin with two to four groups of setae;
dactylus half length of propodus. Peraeopods 5 to 7
all similar in structure and slightly longer
successively; dactylus more than half length of
propodus. Peraeopod 5 (Fig. 3f) subequal to
peraeopod 4, basis posterior lobe deeper than wide,
smaller than posterior lobe of peraeopod 7, merus
with two hind marginal setae, proximal and distal
setae subequal. Peraeopod 6 (Fig. 2b) longer than
peraeopod 4, basis posterior lobe deeper than wide,
similar to posterior lobe of peraeopod 5, smaller than
posterior lobe of peraeopod 7. Peraeopod 7 (Fig. 2e)
3
subequal to peraeopod 6, basis posterior lobe wider
than deep.
Pleopods not modified; peduncle slender; longest
ramus longer than peduncle.
Uropod 1 (Fig. 2f) longer than uropod 2; peduncle
longer than rami; rami subequal; inner ramus with
two dorsal and six distal setae, two of them longer,
male without curved setae on inner side of the
ramus; outer ramus with one dorsal seta, and three
distal setae; peduncle setation present.
Uropod 2 (Fig. 3d) rami subequal; inner ramus
with two dorsal and three distal setae; outer ramus
without dorsal and four distal setae; peduncle
setation present.
Uropod 3 (Fig. 2d) same length as urosomite 3,
shorter than peduncle of uropod 1, shorter than
peduncle of uropod 2; peduncle globose, wider than
ramus, with two strong distal and two marginal
setae; inner ramus absent; outer ramus uniarticulate,
shorter than peduncle, basal width more than two
times tip of ramus, with three simple apical slender
setae, and one conate seta.
Telson (Fig. 3c) wider than long, entire, apically
rounded, with more than two short simple setae,
arranged symmetrically on the apical margin.
Coxal gills sac-like, on segments 2 to 6. Ventrola-
teral sternal gills tubular, present on segments 3 to 7.
Characters of female that differ from male. Size
8.8 mm. Antenna 1 flagellum with eight articles.
Antenna 2 flagellum with ten articles. Gnathopod 1
(Fig. 5a, b) similar in size and of same shape as
gnathopod 2; similar to male gnathopod 1 in size and
shape. Gnathopod 2 (Fig. 5c, d) smaller than male
gnathopod 2 and different in shape, propodus length
less than two times maximum width, normally
subchelate, palm transverse.
Habitat. Freshwater, epigean, normally more than
10 m depth.
Distribution. Endemic to Lake Titicaca, Peru and
Bolivia.
Hyalella longispina n. sp. (Figs 6-11)
Type material. See above section on material and
methods.
Description of male holotype. Head (Fig. 6) longer
than peraeonite 1, rostrum absent. Eyes circular.
Peraeonites and pleonites 1-3 with transverse roun-
ded elevations near posterior margin. Epimera 1-3
with straight posterior margin, epimeral plate 1
rounded ventrally, plate 2 with rounded posteroven-
Org. Divers. Evol. 2, Electr. Suppl. 6 (2002)
González & Coleman: Hyalella armata (Faxon) and H. longispina n. sp.
tral angle, plate 3 pointed posteroventrally.
Urosomite 1 longest; urosomite 2 rather short, with
urosomite 3 attached to the posterodorsal margin;
urosomite 3 surpassed by the telson.
Antenna 1 (Fig. 6b) peduncular article ratios:
1:1:0.67; flagellum damaged, at least 10 articles.
Antenna 2 (Fig. 8d) peduncular article 1 massive,
deeply incorporated in lateral head shield, articles 3-
5 successively longer; 10 flagellar articles. Labrum
(Fig. 6d) truncate, with hair-like setae close to apical
margin. Mandible with 5-dentate incisor on the right
side (4-dentate on left); lacinia mobilis with 2 strong
teeth and a few smaller denticles on the right
mandible (3 teeth on left); setal row consisting of 2
long setae and a row of shorter setae; pars molaris
ridged with stout seta on the anterior face. Lower lip
(hypopharynx) (Fig. 6f) with wide rounded lobes
and short rounded mandibular processes. Maxilla 1
palp uniarticulate, not reaching distal margin of
outer plate, with 1 terminal seta; outer plate with 9
strongly serrate apical spiniform setae; inner plate
slender with 2 apical pappose setae. Maxilla 2 (Fig.
8b) inner plate slightly wider and shorter than outer,
with one stout long medial seta and rows of apical
setae shorter than those on outer plate. Maxilliped
(Fig. 8c) inner plate with 3 apical robust setae; outer
plate longer than inner plate, relatively slender; palp
article 2 medially produced; article 3 ovoid; article 4
slender, with apical nail. Gnathopod 1 (Fig. 8a, e)
coxa with strongly tapering and pointed process,
directed lateroventrally (Fig. 7); basis slightly con-
cave anteromarginally, convex posteroventrally;
ischium subquadrate with anteromarginal notch;
anterior margin of merus with articulation of carpus;
carpus with setose posteromarginal rounded lobe;
propodus expanded distally, two groups of setae on
the outer face, medial face with stout setae; palm
with setae of different length and one posterior
spine-like seta; dactylus stout and only weakly
curved. Gnathopod 2 (Fig. 9d) coxa directed more
ventrally than those of peraeopods 1 and 3, shape
similar to gnathopod 1 but longer (142 %), wider
proximally; basis with anterior straight margin,
posteriorly slightly convex with some marginal
setae; ischium subquadrate, anterior margin
expanded distally; merus subrectangular, posterodi-
stally angular; carpus shorter than that of gnathopod
1, with projecting, relatively narrow carpal lobe,
guarding more than half length of the posterior
margin of propodus, with some terminal short setae
and a row of setae on the medial face; propodus
massive, expanded distally, palm oblique, with two
short spine-like setae at posterior angle, palm lined
4
with short setae; dactylus weakly curved with
microtrichs on inner curvature. Peraeopod 3 coxa
(Fig. 9b) similar to that of gnathopods, but anteriorly
deeply excavate; basis with anterior straight margin,
posteromarginally sinuous; ischium as for gnathopod
2; merus slightly expanded distally and slightly
drawn out anterodistally; merus to propodus sube-
qual in length, successively narrower; dactylus stout
with apical nail. Peraeopod 4 (Fig. 9a) coxa with
extremely elongate pointed process, directed
laterally, wing-like (Fig. 7), span from left to right
process apex longer than total body length (Fig. 12),
additional short subacute process on posterior
margin close to body articulation of coxa; basis to
dactylus as for peraeopod 3, except for slightly
longer carpus and dactylus. Peraeopod 5 (Fig. 10a)
coxa wider than long, bilobed, anterior lobe
ventrally rounded and slightly shorter than narrower
posterior lobe; basis tapering distally, weakly
rounded anteromarginally with groups of setae,
posterior margin slightly sinuous, apical region with
weak depression; ischium longer than wide, with
straight anterior margin and weak excavation
posteriorly; merus somewhat expanded distally,
drawn out posterodistally, with apical group of
setae; carpus 1.12 x merus; propodus 1.28 x merus,
dactylus only weakly curved. Peraeopod 6 (Fig. 10b)
coxa bilobed, anterior lobe very short, angular; basis
similar in shape to that of peraeopod 5, but wider
and longer; ischium as for peraeopod 5; merus to
propodus similar in shape, much longer than those of
peraeopod 5, relative lengths 1.00:1.26:1.32, dac-
tylus slightly longer than that of peraeopod 5.
Peraeopod 7 (Fig. 10d) shorter than peraeopod 6,
coxa smallest, rounded anteroventrally, posterior
margin rather straight; basis much wider than those
of peraeopods 5-6, ovoid, lobate posteroventrally,
groups of setae on anterior margin; shape of ischium
to dactylus as for peraeopod 6. Pleopods (Fig. 10c)
normal. Uropod 1 (Fig. 11f) peduncle longer than
inner ramus, outer ramus slightly shortened;
peduncle and rami sparsely setose, rami with
terminal setae of different length. Uropod 2 (Fig.
11c) peduncle as long as inner ramus, outer ramus
slightly shortened compared to inner, with terminal
setae only. Uropod 3 reduced in length, peduncle
longer than ramus, only one ramus present, with
some terminal setae. Telson wider than long, entire
with some marginal short setae (Fig. 11g).
Characters of female allotype that differ from
male holotype. Size 9 mm. Gnathopod 2 (Fig. 11a,
b) resembles gnathopod 1 (Fig. 11e, d) in the wide
Org. Divers. Evol. 2, Electr. Suppl. 6 (2002)
González & Coleman: Hyalella armata (Faxon) and H. longispina n. sp.
and strongly setose carpal lobe. Propodus of
gnathopod 2 very similar to that of gnathopod 1.
Habitat. Freshwater, epigean, more than 10 m
depth.
Distribution. Endemic to Lake Titicaca, Peru and
Bolivia.
Discussion
The high diversity of species of Hyalella in the lake
is unexpected and not well understood (Crawford et
al., 1993). The amphipod fauna in Lake Titicaca has
been compared with the fauna of Lake Baikal and
the speciation events considered similar. However, it
is important to consider that Lake Titicaca is not as
old as Lake Baikal. According to Lavenu (1992), the
present lake system on the Altiplano is derived from
a more ancient system which was there since the
lower Pleistocene, no more than two millions years
ago. Bousfield (1982) suggested that the endemic
fauna of the lake could not be older than the lake
basin, estimated by him as originating 10 million
years ago.
Similar to Lake Baikal also in Lake Titicaca
several amphipod species have body processes. It is
conceivable that these pointed processes act as a
defense mechanism against predation as there are
more than 28 species of fish in the lake.
When we started to work on H. armata we found
striking differences in the length of the coxal
acumination within the extensive samples collected
by George Crawford. At a closer look and after a
morphometric analysis of the length/width ratio of
the animals (Fig. 13), a discontinuity between these
two populations became apparent. These differences
were used to separate two distinct species. Both
species are very different from all hyalellids known
so far. They bear pointed coxal plates that are
directed laterally, which is unique in the Hyalellidae.
We searched for further characters, such as setal
patterns and length relations of articles of
appendages, in order to find additional morphologi-
cal and morphometrical differences between these
groups. However, we only found a large variability
in these characters. Nevertheless we consider the
non-overlapping character clouds of the length/width
ratios as indicators of genetic separation of these
populations. There is no geographical or obvious
habitat separation of both populations, but they
inhabit various locations in Lake Titicaca (Fig. 14).
In spite of a potential contact of these populations,
5
there is evidently no intermediate character state of
the length/width ratio in these populations.
The low number of morphological characters
separating these two species perhaps reflects their
recent origin. Without success we tried to extract
DNA from the museum samples in order to get
genetic information on these populations. With fresh
material we would like to check in a future study if
there are more characters on the molecular level.
A comparable similarity between hyalellid popu-
lations in Lake Titicaca occurs in the H. echinus
complex we are currently working on.
Acknowledgements
Dr. Ardis Johnston from the Museum of
Comparative Zoology, Harvard University, loaned
Faxon’s material. This work was largely funded by
Universidad Católica del Norte, Chile, and
University of Maine, USA. Thanks also to Dr. C.
Dejoux for providing specimens for this study. Ms.
Jaqueline Meier made the drawings of H. longispina.
References
Barnard, J. L., & Barnard., C. M. (1983): Freshwater
Amphipoda of the World. vol. I. Evolutionary
Patterns, vol. II. Handbook and Bibliography. 830
pp., Hayfield Associates, Mt. Vernon, Virginia.
Bousfield, E. L. (1982): Amphipoda: paleohistory.
Pp. 96-100 in: McGraw-Hill Yearbook of Science
and Technology for 1982-1983.
Bousfield, E. L. (1996): A contribution to the reclas-
sification of Neotropical freshwater hyalellid
amphipods (Crustacea: Gammaridea, Talitroidea).
Boll. Mus. Civ. Storia Nat. Verona 20: 175-224.
Chevreux, E. (1904): Mission de Crequi-Monfort et
Senegal de la Grange. Note preliminaire sur les
amphipodes recueillis par M. le Dr. Neveu-
Lemaire dans le Lac Titicaca. (Julliet, 1903).
Bull. Soc. Zool. Fr. 29: 131-134, 2 figures.
Chevreux, E. (1907): Les amphipodes des lacs des
hauts plateaux de l’Amerique du Sud. Mission
Scientifique G. de Crequi-Monfort et Senegal de
la Grange, Lacs Hauts Plateaux de l’Amerique du
Sud. 22 pp, figs 30-41.
Crawford, G. I., Harrison, K., Lincoln, R. J. &
Boxshall, G. A. (1993): An introduction to the
species flock of amphipods (Crustacea) of Lake
Titicaca. Workshop Speciation in Ancient Lakes.
Evolution, Biodiversity, Conservation. Scientific
Org. Divers. Evol. 2, Electr. Suppl. 6 (2002)
González & Coleman: Hyalella armata (Faxon) and H. longispina n. sp.
Station of the Hautes-Fagnes, Mont-Rigi,
Robertsville, Belgium.
Dejoux, C., (1992): The Amphipoda. Pp. 346-356
in: Dejoux, C. & Iltis, A. (eds) Lake Titicaca. A
Synthesis of Limnological Knowledge. Kluwer
Academic Publishers, Dordrecht, The
Netherlands.
Della Valle, A. (1893): Gammarini del Golfo di
Napoli. Fauna Flora Golf. Neapel angrenz. Mee-
res-Abschn., Monogr. 20: 1-948, 61 pls.
Faxon, W. (1876): Exploration of Lake Titicaca by
Alexander Agassiz and S. W. Garman. IV.
Crustacea. Bull. Mus. Compar. Zool. 3: 361-375,
37 figs.
Fig. 1. Hyalella armata (Faxon, 1876), male 9.5 mm. a: left aspect of habitus; b: epimeral plates 1-3; c: gnathopod 2; d: gnathopod 1. Scale
bars: a, b = 727 µm; c, d = 192 µm.
6
Lavenu, A. (1992): Origins, formation and
geological evolution. Pp. 3-15 in: Dejoux, C. &
Iltis, A. (eds) Lake Titicaca. A Synthesis of
Limnological Knowledge. Kluwer Academic
Publishers, Dordrecht, The Netherlands.
Stebbing, T. R. R. (1888): Report on the Amphipoda
collected by H. M. S. Challenger during the years
1873-1876. Rep. Sci. Res. Voy. H. M. S.
Challenger 1873-1876, Zool. 29: 1-1737, 210 pls.
Stebbing, T. R. R. (1906): Amphipoda I. Gammaridea.
Das Tierreich 21: 1-806.
Watling, L. (1993): Functional morphology of the
amphipod mandible. J. Nat. Hist. 27: 837-849.
Org. Divers. Evol. 2, Electr. Suppl. 6 (2002)
González & Coleman: Hyalella armata (Faxon) and H. longispina n. sp. 7

Fig. 2. Hyalella armata (Faxon, 1876), male 9.5 mm. a: maxilla 1; b: peraeopod 6; c: palp of maxilla 1; d: uropod 3; e: peraeopod 7; f: uro-
pod 1. Scale bars: a, c, d = 94 µm; b, e, f = 192 µm.

Org. Divers. Evol. 2, Electr. Suppl. 6 (2002)

González & Coleman: Hyalella armata (Faxon) and H. longispina n. sp. 8

Fig. 3. Hyalella armata (Faxon, 1876), male 9.5 mm. a: left maxilliped. b: maxilla 2; c: telson; d: uropod 2; e: peraeopod 4; f: peraeopod 5.
Scale bars: a, b = 94 µm; c-f = 192 µm.

Org. Divers. Evol. 2, Electr. Suppl. 6 (2002)

González & Coleman: Hyalella armata (Faxon) and H. longispina n. sp. 9

Fig. 4. Hyalella armata (Faxon, 1876), male 9.5 mm. a: gnathopod 1 detail; b: peraeopod 3; c: lower lip (hypopharynx); d: antenna 1, distal
flagellomeres omitted; e: antenna 2, distal flagellomeres omitted; f: upper lip (labrum); g: left mandible. Scale bars: a = 94 µm; b-g = 192
µm.

Org. Divers. Evol. 2, Electr. Suppl. 6 (2002)

González & Coleman: Hyalella armata (Faxon) and H. longispina n. sp. 10

Fig. 5. Hyalella armata (Faxon, 1876), female “f” 8.8 mm. a: gnathopod 1; b: chela of gnathopod 1; c: gnathopod 2; d: chela of gnathopod 2.
Scale bars: a, d = 192 µm; b, c = 94 µm.

Org. Divers. Evol. 2, Electr. Suppl. 6 (2002)

González & Coleman: Hyalella armata (Faxon) and H. longispina n. sp. 11

Fig. 6. Hyalella longispina n. sp., holotype male, 10 mm. a: habitus; b: antenna 1; c: left mandible; d: upper lip (labrum); e: right mandible; f:
lower lip (hypopharynx). Scale bars a = 1 mm; b, c, e = 100 µm; d, f = 200 µm.

Org. Divers. Evol. 2, Electr. Suppl. 6 (2002)

González & Coleman: Hyalella armata (Faxon) and H. longispina n. sp. 12

Fig. 7. Hyalella longispina n. sp., holotype male, 10 mm. Dorsal habitus.

Org. Divers. Evol. 2, Electr. Suppl. 6 (2002)

González & Coleman: Hyalella armata (Faxon) and H. longispina n. sp. 13

Fig. 8. Hyalella longispina n. sp., holotype male, 10 mm. a: gnathopod 1 detail; b: maxilla 2; c: maxilliped, palp of right side omitted; d:
antenna 2; e: gnathopod 1; f: maxilla 1. Scale bars: a-d, f = 100 µm, e = 200 µm.

Org. Divers. Evol. 2, Electr. Suppl. 6 (2002)

González & Coleman: Hyalella armata (Faxon) and H. longispina n. sp. 14

Fig. 9. Hyalella longispina n. sp., holotype male, 10 mm. a: peraeopod 4; b: coxa of peraeopod 3; c: basis to dactylus of peraeopod 3; d:
gnathopod 2, setation of medial face of carpal lobe omitted. Scale bars: a-d = 200 µm.

Org. Divers. Evol. 2, Electr. Suppl. 6 (2002)

González & Coleman: Hyalella armata (Faxon) and H. longispina n. sp. 15

Fig. 10. Hyalella longispina n. sp., holotype male, 10 mm. a: peraeopod 5; b: peraeopod 6; c: pleopod 1; d: peraeopod 7. Scale bars: a-d =
200 µm.

Org. Divers. Evol. 2, Electr. Suppl. 6 (2002)

González & Coleman: Hyalella armata (Faxon) and H. longispina n. sp. 16

Fig. 11. Hyalella longispina n. sp., a, b, e, d; female allotype, 9 mm; c, f, g male holotype, 10 mm. a: chela of peraeopod 2; b: peraeopod 2;
c: uropod 2; d: peraeopod 1; e: chela of peraeopod 1; f: uropod 1; g: uropod 3 and telson. Scale bars: a, e = 100 µm; b-d, f, g = 200 µm.

Org. Divers. Evol. 2, Electr. Suppl. 6 (2002)

González & Coleman: Hyalella armata (Faxon) and H. longispina n. sp. 17

Fig 12. Hyalella armata complex. a. H. armata, dorsal habitus aspect. b. H. armata, left lateral view. c. H. longispina n. sp.

Org. Divers. Evol. 2, Electr. Suppl. 6 (2002)

González & Coleman: Hyalella armata (Faxon) and H. longispina n. sp. 18

11

10
width (coxae 4)

9
armata

8 longispina

5
0 2 4 6 8 10 12 14

animal length

Fig. 13. Body length vs. width for two populations of the Hyalella armata complex, n=40. The two separate clouds of dots show the distinc-
tion between H. longispina n. sp. (black triangles) from Choccocoyo-Bay, GIC 481, and H. armata (grey diamonds) from Coata Bay, GIC
263.

Org. Divers. Evol. 2, Electr. Suppl. 6 (2002)

González & Coleman: Hyalella armata (Faxon) and H. longispina n. sp. 19

Fig. 14. Map of Lake Titicaca with collecting sites of Hyalella armata (circles) and Hyalella longispina (diamonds).

Org. Divers. Evol. 2, Electr. Suppl. 6 (2002)