A new species of Ptychochromis from northeastern Madagascar (Teleostei: Cichlidae), with an updated phylogeny and revised diagnosis for the genus
Author
Sparks, John S.
Author
Stiassny, Melanie L. J.
text
Zootaxa
2010
2341
33
51
journal article
10.5281/zenodo.193312
becbe9e7-b072-464c-a565-fb0915d012f4
1175-5326
193312
Ptychochromis ernestmagnusi
,
new species
Figures 1– 5
;
Table 1
Holotype
:
AMNH
249490, 146.6 mm
SL, male, Mananara (du nord) River at Antanambaobe Village (16° 16′ 0 1″S, 49°40′ 0 1″E),
Madagascar
,
120 m
a.s.l., MG 10-06, coll. P.V. Loiselle,
4 October 2006
.
Paratypes
:
AMNH
249488, 9 ex., incl. 1 ex. C&S,
75.9–99.5 mm
SL, Mananara (du nord) River at Antanibaolina Village (16° 15′ 0 1″S, 49 ° 40′ 41″E),
Madagascar
,
108 m
a.s.l., MG 09-06, coll. P.V. Loiselle,
4 October 2006
.
AMNH
249489, 8 ex., incl. 1 ex. C&S, 51.5–99.0 mm SL, data as for
holotype
.
MNHN
2009-1674, 2 ex., 79.4–98.0 mm SL, data as for
holotype
.
UMMZ
248823, 2 ex., 83.0–
86.4 mm
SL, data as for
AMNH
249488.
Non-Type Material Examined:
MNHN
1935-0007, 1 ex.,
128.5 mm
SL, Mananara (du nord) River,
Madagascar
. No additional collection locality information available.
FIGURE 1.
Ptychochromis ernestmagnusi
, AMNH 249490, holotype, adult male, 146.6 mm SL (scale bar = 1cm).
Diagnosis.
A
Ptychochromis
exhibiting the eastern-type palatine morphology (
Fig. 2
) and further distinguished from all congeners except
P. grandidieri
,
P. loisellei
,
P. m a k i r a
and
P. curvidens
by an anterior displacement of the first supraneural such that it overlies the dorsoposterior margin of the supraoccipital (
Fig. 3
A– C). Among eastern group
Ptychochromis
,
P. m a k i r a
shares with
P. ernestmagnusi
the possession of supraneurals with a characteristically flattened dorsal profile, which is interpreted here as a synapomorphy uniting these two geographically proximate species. Also shared with
P. m a k i r a
is the presence of strong (paired) lateral barring as a prominent component of pigmentation patterning; however,
P. m a k i r a
can be distinguished by distinctively V-shaped lateral flank bars, whereas in
P. ernestmagnusi
the bars are oriented vertically.
Ptychochromis ernestmagnusi
is further distinguished from
P. m a k i r a
by conspicuous iridescent spangling on the flank and dorsal fin near its base, dusky grayish-green base coloration (vs. whitish), four lachrymal laterosensory foramina (vs. three), and a total of seven (vs. six) infraorbital bones.
Description.
Morphometric and meristic data presented in
Table 1
. External anatomical characteristics and general pigmentation pattern in life and preservation can be observed in
Figures 1
,
5
and
7
B. Moderately deep bodied and laterally compressed. Dorsal body profile convex, becoming significantly more so in larger specimens (ca.
100 mm
SL and larger). Ventral body profile weakly to moderately convex. Lateral snout outline straight in smaller specimens, and becoming weakly curved (convex) in larger individuals. Predorsal profile moderately to strongly convex from mid-orbit to dorsal-fin origin, becoming more pronounced in larger specimens and creating weak “nuchal hump”. Supraoccipital crest prominent in lateral view and conspicuously deep bodied in larger individuals (ca.
100 mm
SL). Caudal peduncle short, deep, and laterally compressed. Origin of dorsal fin located well anterior to vertical through pectoral-fin insertion. Origin of pelvic fin located considerably posterior to vertical through pectoral-fin insertion.
Total vertebral count 27 or 28, with formulae of 13 + 14 (mode), 13 + 15, and 14 + 14 precaudal and caudal vertebrae, respectively.
Oral jaws isognathous and small. Oral dentition bilaterally symmetrical and bicuspid, with moderately to well-developed distally expanded and slightly recurved cusps (
Fig. 2
). Outer row teeth of both premaxilla and dentary enlarged relative to teeth of inner rows and graded in size (i.e., becoming smaller) posterolaterally.
Outer row teeth procumbently implanted in rostral portion of lower jaw, and oriented vertically elsewhere. Outer row teeth in upper jaw more or less vertically oriented. Upper jaw with three or four rows of teeth anteriorly and tapering to single (i.e., outer) row posteriorly. Lower jaw with three rows of teeth rostrally, and tapering to single (i.e., outer) row posteriorly. Although smaller, inner rows of teeth on both premaxilla and dentary of same morphology (bilaterally symmetrical and bicuspid) as those of respective outer row. Dentition covers about anterior 2/3 of dentary and nearly entire surface (>80%) of premaxillary arcade.
TABLE 1.
Morphometric and meristic data for
Ptychochromis ernestmagnusi
,
new species
. Proportional measurements (mm) in percent standard length (SL) or percent head length (HL), unless noted otherwise. Values in parentheses indicate number of specimens examined with that count. (H) indicates count corresponding to holotype.
| Character |
P. ernestmagnusi
N Holotype Range
|
Mean |
SD |
| Standard length (mm) |
18 146.6 51.5–146.6 |
83.5 |
| Percentage of SL |
| Head length |
18 35.7 34.2–37.1 |
35.3 |
0.92 |
| Body depth |
18 46.1 41.7–46.1 |
43.9 |
1.40 |
| Predorsal length |
18 42.3 40.4–44.6 |
42.3 |
1.10 |
| Preanal length |
18 71.6 71.6–73.5 |
72.7 |
0.66 |
| Prepelvic length |
18 41.3 39.8–48.7 |
42.1 |
1.97 |
| Head width (max.) |
18 18.6 15.9–18.6 |
17.3 |
0.74 |
| Caudal peduncle length |
18 15.0 12.4–15.9 |
14.4 |
0.82 |
| Caudal peduncle width |
16 6.2 4.9–6.7 |
5.9 |
0.60 |
| Caudal peduncle depth |
18 16.5 13.3–16.5 |
15.2 |
0.85 |
| Pectoral-fin length |
18 31.0 29.8–34.1 |
32.1 |
1.24 |
| Pelvic-fin length |
18 30.9 24.3–31.2 |
28.7 |
1.76 |
| Percentage of HL |
| Snout length |
18 46.3 35.3–46.3 |
39.3 |
2.71 |
| Orbit diameter |
18 29.5 29.5–37.4 |
34.2 |
2.13 |
| Upper-jaw length |
18 37.7 26.3–37.7 |
30.1 |
2.69 |
| Lower-jaw length |
18 39.6 35.6–41.2 |
38.4 |
1.38 |
| Interorbital width |
18 32.1 25.6–32.1 |
28.3 |
1.55 |
| Preorbital depth |
18 28.7 20.0–28.7 |
24.6 |
2.24 |
| Caudal peduncle length/depth |
18 0.9 0.8–1.1 |
1.0 |
0.07 |
| Caudal peduncle length/width |
16 2.4 2.1–3.3 |
2.5 |
0.33 |
Scales in lateral line 16 34 (8), 35 (5, H), 36 (2), 37 (1)
Scales: lateral line to dorsal fin 18 4 (8), 4.5 (2), 5 (7, H), 6 (1)
Scales: pectoral to pelvic bases 18 4 (4), 5 (14, H)
Gill rakers (lower limb 1st arch) 18 11 (15, H), 12 (3)
Vertebrae (pre-caudal + caudal) 18 13+14 = 27 (15), 13+15 = 28 (1), 14+14 = 28 (2, H) Dorsal fin
18 XII 13
(1), XIII 11 (1), XIII 12 (15, H), XIII 13 (1) Anal fin
18 III 7
(2),
III 8
(13, H),
III 9
(3)
FIGURE 2.
Left lateral view of the oral jaws and anterior region of the suspensorium of
P. ernestmagnusi
(AMNH 249489, paratype, 87.5 mm SL, female, C&S) illustrating the “eastern type palatine” (shaded in gray) morphology.
Lower pharyngeal jaw (LPJ = fused 5th ceratobranchial elements) robust with interdigitating suture on posteroventral margin. Dentition on LPJ and upper pharyngeal jaw (UPJ) comprised of numerous, closely set, strongly hooked and bicuspid teeth. Cusps on LPJ teeth better developed posteriorly. Posteromedially on both LPJ and third pharyngobranchial toothplate, dentition becoming robust and molariform (Fig. 4A); other teeth of these elements hooked and bicuspid. Expansive second pharyngobranchial toothplate bearing five or six rows of well-developed, hooked and bicuspid teeth. Two or three rows of hooked and bicuspid teeth present on “free” second epibranchial toothplate. Fourth upper toothplate covered with numerous, closely-set rows of smaller hooked and bicuspid teeth; teeth becoming progressively smaller and much less well developed posteriorly. Strong concavity and associated sickle-like prong present on caudomedial margin of fourth upper toothplate. Dorsal surface of fourth ceratobranchial elements bearing numerous robust, laterally expanded, toothplates. Fourth ceratobranchial toothplates confluent with outer row gill rakers of these elements. Dentition on fourth ceratobranchial toothplates unicuspid and more or less conical to weakly hooked and bicuspid laterally, and becoming progressively more strongly hooked and bicuspid medially (similar to pattern observed for lateral LPJ dentition). Contralateral fourth ceratobranchial elements bearing strong concavity and associated prong (= hook) on medial margin.
FIGURE 3.
Left lateral view of the dorsoposterior margin of the neurocranium illustrating morphology of the supraneural bones (in black) and their relationship to the supraoccipital bone. A)
Ptychochromis ernestmagnusi
, B)
P. makira
, C)
P. c u r v i d e n s
, and D)
P. oligacanthus
. Arrows indicate degree of overlap of supraoccipital crest by anterior supraneural.
Eleven or 12 relatively elongate gill rakers arrayed along lower limb of first arch, excluding raker in angle of arch (Fig. 4B). Lower limb rakers of first gill arch denticulate dorsomedially, bearing numerous conical to weakly hooked and bicuspid teeth. Nine weakly developed and triangular epibranchial gill rakers. Remaining gill arches bearing short, robust, and strongly laterally expanded (particularly, distally near crown) rakers. These rakers strongly denticulate dorsally, bearing numerous elongate and conical (becoming bulbous apically) to weakly hooked and bicuspid teeth.
FIGURE 4.
Ptychochromis ernestmagnusi
. (A) Dorsal view of the lower pharyngeal jaw (fused 5th ceratobranchial elements) illustrating molariform caudomedial tooth morphology. Left lateral view of the: (B) lower limb of the first gill arch (ceratobranchial 1 and hypobranchial 1) with gill rakers shaded in gray, and (C) lachrymal (pores shaded in gray) and additional bones of the infraorbital series.
Flank squamation comprised of large, regularly imbricate, weakly ctenoid scales. Scale margins becoming progressively more ctenoid posteriorly on flank. Ctenoid scales extend from about dorsal-fin origin (i.e., ranging from slightly anterior to somewhat posterior of fin insertion) dorsal to upper branch of lateral line and somewhat posterior to pectoral-fin base below upper lateral line, to proximal portion of caudal fin. Scales on anterior portion of nape and throughout head region cycloid. Scales on opercle and subopercle cycloid. Cheek scales cycloid and comprising four rows; fourth (ventral) row frequently poorly developed and comprising few scales. Snout, lachrymal, and anterior portion of interorbital region to about level of mid-orbit asquamate. Anterior chest scales somewhat reduced in size and embedded. Scales extending onto caudal fin reduced in size and ctenoid anteriorly, markedly smaller and cycloid posteriorly. Pored scales of lower branch of lateral line frequently extending onto caudal fin (i.e., beyond hypural flexure) for one or two rows. Lateral-line scales with well-developed canals, and numbering 34 to 37 (mode 34). Four or five (mode) scale rows between bases of pectoral and pelvic fins. Four (mode) to six scales in diagonal from upper branch of lateral line to dorsal-fin origin. No scale rows extending onto dorsal- and anal-fin membranes proximal to base of fins.
Dorsal fin with XII or XIII spines and 11 to 13 soft rays. Anal fin with III spines and seven to nine soft rays. First anal-fin spine conspicuously short, whereas second and third spines elongate and more or less similar in length. Distal margins of soft dorsal and anal fins becoming produced and tapered in larger specimens; posteriorly margins reaching to about caudal-fin origin in smaller individuals (<
80 mm
SL) and extending well beyond origin in larger specimens (
Fig. 1
). Pectoral fin elongate, deep bodied and paddle-like; becoming tapered distally (i.e., dorsal rays much longer than ventral). Adpressed pelvic fin terminating well before anal-fin origin in smaller specimens, and extending to about anal-fin origin in larger individuals (>
90 mm
SL). Caudal fin emarginate, trailing margins of upper and lower lobes becoming at most weakly produced in larger individuals (>
80 mm
SL).
Miscellaneous osteology and anatomy.
Exoccipital foramina on posterior of neurocranium poorly developed; simple and lacking complex interior chamber (see
Stiassny, 1991
, and Sparks, 2008, for a discussion of exoccipital foramen development in Malagasy-South Asian cichlids). Paired, anterior gas bladder diverticula well developed, elongate, and tube-like; however, rather feeble in structure (i.e., not rigid and thick walled) and similar to main gas bladder chamber. Diverticula in contact with exoccipital region of neurocranium via connective tissue but not penetrating into exoccipital foramina (Sparks, 2008;
Fig. 3
A). Infraorbital series composed of seven distinct elements (Fig. 4C). Lachrymal (= first infraorbital or IO1) with four neurosensory pores; fourth pore communicating with anterior pore of second infraorbital (IO2). Second infraorbital excluded from orbital margin by IO3. Cephalic laterosensory canals well developed with enlarged pores (e.g., including those on preopercle and dentary;
Figs. 2
& 4C). Uncinate process and anterior arm of first epibranchial element short and robust (Fig. 4B). Well-developed process (= prong/hook) and deep indentation (= excavation) present on medial face of fourth ceratobranchial element. Supraneurals (
Fig. 3
A) rostrally positioned, with first supraneural overlying dorsoposterior margin of supraoccipital crest. Both supraneurals characteristically shaped with flat dorsal margins (
Fig. 3
A). Posterior supraneural often reduced in size or absent.
Coloration in life
(
Figure 5
). Overall uniform greenish base coloration with a dusky gray overlay, not notably darker dorsally than ventrally. Many scales bearing a small, conspicuous iridescent spot along posterior scale margin. Pigmentation pattern composed of five or six (mode,
holotype
) prominent midlateral blotches intersected by six to eight less strongly pigmented vertical bars. Nape dark gray, snout and cheek dusky grey, and gular region black. Fins uniformly dark blackish-gray, with some small iridescent spots proximally in soft dorsal. Pectoral fin hyaline.
Coloration in preservation
(
Figure 1
). Ground coloration reddish-brown, slightly paler ventrally than dorsally. Traces of iridescent spangling present, particularly in larger specimens, and most evident ventrally on flank. Pigmentation pattern consisting of five or six (mode,
holotype
) prominent midlateral blotches with significantly paler intersecting vertical bars retained in preservation. Most posterior blotch, located on caudal peduncle (i.e., sixth blotch in series if present), significantly paler than others. Fins pale reddish-brown, trailing margins of soft anal and dorsal fins blackish. Pectoral fin hyaline. Pelvic fin pale reddish-brown, and becoming charcoal to blackish distally. Anterior interorbital region, snout, and lachrymal dark gray. Lower lip creamy brown. Gular region dark grayish-black.
Distribution and habitat
(
Figure 6
). Currently known only from the
type
localities, which are located in the middle to lower reaches of the Mananara (du nord) River, northeastern
Madagascar
. This region is characterized by humid, lowland rainforest (UNESCO - MAB Biosphere Reserves Directory, 2009), and remains poorly surveyed for freshwater fishes. It is unknown whether the new species is more widely distributed within the region, and also whether it occurs in smaller tributaries of the Mananara (du nord) River or other adjacent drainage basins to the north and south. Substantial forest cover remains in the region and it is likely the new species has a significantly more widespread geographic distribution than current collection data would indicate. The Mananara (du nord) River is a relatively large basin that extends from its headwaters through regions of intact forest and low population density. It is a typical eastern drainage, with a steep overall profile, and a generally rocky to sandy substrate. It is likely that the new species of
Ptychochromis
is restricted to the middle to lower reaches of the river and its tributaries, given that suitable habitat and adequate trophic resources are most likely lacking at higher elevations, as is typical for other eastern basins (
Sparks, 2005a
,
b
).
FIGURE 5.
Ptychochromis ernestmagnusi
, AMNH 249490, holotype, 146.6 mm SL, adult male. Freshly captured specimen illustrating adult coloration in life. Photo by Paul Loiselle.
Conservation status.
Although forested portions of the coastal region to the south of the Mananara (du nord) River are encompassed by the Mananara-Nord Biosphere Reserve (designated in 1990), which extends from 16° 09' to
16° 36'S
and 49° 31' to
49° 53'E
, and covers 140,000 hectares, the Mananara River and its south bank tributaries are not included within the terrestrial component of the protected area (which also includes a smaller marine component) (UNESCO - MAB Biosphere Reserves Directory, 2009). We anticipate that the new species is not only more widespread in the region due to similar available habitats, but that it also receives some degree of protection from habitat degradation, deforestation, and overfishing within the biosphere reserve, which protects some of the last remaining remnants of lowland rainforest in eastern
Madagascar
. In general, species of
Ptychochromis
seem able to tolerate a moderate degree of habitat degradation (in particular
P. grandidieri
, which remains relatively common and widespread along the highly developed and densely populated eastern coast of the island). Nevertheless, members of the genus are rapidly extirpated from areas where habitats have become severely disturbed and water quality is negatively impacted (e.g., severe deforestation and development resulting in highly turbid drainage basins) (
Sparks & Stiassny, 2003
,
2008
).
Etymology.
Named in honor of Mr. Ernest Magnus of Berlin,
Germany
, and New York City, at the request of the family of Dr. Rudolph G. Arndt, whose support of ichthyological exploration and research at the American Museum of Natural History is gratefully acknowledged.
FIGURE 6.
Relative geographic distributions of species of
Ptychochromis
exhibiting the “eastern type palatine” configuration, including the geographically proximate sister species,
P. m a k i r a
and
P. ernestmagnusi
,
new species
. Note: Arrow for
P. grandidieri
indicates that distribution for species extends southward along the eastern coast of Madagascar.
Discussion and comparisons.
Stiassny and Sparks (2006)
corroborated the monophyly of, and resolved intergeneric relationships within, the endemic
Malagasy
cichlid subfamily
Ptychochrominae
based on the analysis of a combination of nucleotide sequence data and anatomical features, but resolution of relationships within
Ptychochromis
was problematic in that study. Based primarily on features of the palatine bone of the suspensorium,
Stiassny and Sparks (2006)
recognized two groups of
Ptychochromis
: a “western clade” characterized by a compact palatine head with an upright orientation and marked elevation of the lateral ethmoid process (i.e., “western
type
palatine”), and an “eastern group” in which the palatine exhibits a markedly horizontal orientation, an elongate palatine prong, and less prominent (weakly elevated) lateral ethmoid process (i.e., “eastern
type
palatine”). On the cladogram presented by
Stiassny and Sparks (2006:
Fig. 2
)
, unambiguous optimization of the “western
type
palatine” configuration (
Sparks & Stiassny, 2006: character 14, node D
) was interpreted as evidence for monophyly of a “western clade”; however, the “eastern
type
palatine” morphology described above was not unambiguously optimized on that phylogenetic reconstruction.
As
a result, the relationships of the four eastern species included in our 2006 analysis (viz.,
Ptychochromis grandidieri
,
P. m a k i r a
,
P. loisellei
, and
P. curvidens
) were represented as a polytomy (
Stiassny & Sparks, 2006:
Fig. 2
).
Here we have identified an additional putatively homologous anatomical feature unique to the “eastern group”, rostral displacement of the supraneural elements such that the anterior supraneural comes to overlie the dorsoposterior margin of the supraoccipital of the neurocranium. This supraneural configuration is lacking in all other members of
Ptychochrominae
, and, therefore, is hypothesized to represent an apomorphic feature diagnostic of an “eastern clade” of
Ptychochromis
(e.g.,
Fig. 3
A–C).
To test this hypothesis, we reran our prior phylogenetic analysis (
Stiassny & Sparks, 2006:
Fig. 2
) using the same methodological approach, search parameters, and combined morphological and molecular dataset, with the inclusion of the new species and including coding for the unique “eastern group” supraneural configuration described above and the two features (discussed below) hypothesized to unite
P. m a k i r a
and the new species (for a total of 24 morphological transformations and 2053 total characters; characters 1–21 are described in detail in
Sparks & Stiassny, 2006
:
Table 1
and Results). A complete list of the morphological character descriptions is presented in the Appendix and the corresponding data matrix is presented in
Table 2
.
As
the results indicate (
Fig. 7
), we again failed to recover a monophyletic “eastern group” despite the fact that these five species possess both the novel supraneural feature (character 22,
Fig. 3
A–C) and “eastern
type
palatine” configuration (
Sparks & Stiassny, 2006: character 21;
Fig. 2
) discussed above.
TABLE 2.
Morphological character matrix for species of
Ptychochromis
and outgroups included in phylogenetic analysis. Inapplicable character assignments are designated by (-). “A” = character states 0 & 1. Character state descriptions presented in Appendix 1.
Characters
| 1 |
2 |
3 |
4 |
5 |
6 |
7 |
8 |
9 |
10 |
11 |
12 |
13 |
14 |
15 |
16 |
17 |
18 |
19 |
20 |
21 |
22 |
23 |
24 |
|
Paretroplus damii
|
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
|
Retroculus
|
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
1 |
0 |
1 |
1 |
0 |
0 |
0 |
0 |
0 |
|
Heterochromis
|
0 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
1 |
1 |
0 |
0 |
0 |
1 |
0 |
1 |
0 |
- |
0 |
0 |
0 |
0 |
|
Paratilapia polleni
|
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
A |
0 |
0 |
0 |
0 |
0 |
1 |
1 |
A |
- |
0 |
0 |
0 |
0 |
|
Oxylapia polli
|
1 |
A |
1 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
A |
0 |
0 |
0 |
0 |
0 |
|
Ptychochromoides betsileanus
|
1 |
1 |
1 |
1 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
1 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
|
Ptychochromoides itasy
|
1 |
1 |
1 |
1 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
1 |
1 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
|
Ptychochromoides vondrozo
|
1 |
1 |
1 |
1 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
1 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
|
Katria katria
|
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
0 |
0 |
0 |
0 |
0 |
1 |
1 |
0 |
1 |
1 |
1 |
0 |
0 |
0 |
0 |
|
Ptychochromis oligacanthus
|
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
0 |
1 |
1 |
1 |
1 |
1 |
0 |
0 |
0 |
0 |
|
Ptychochromis insolitus
|
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
2 |
1 |
1 |
1 |
1 |
0 |
0 |
0 |
0 |
|
Ptychochromis onilahy
|
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
0 |
1 |
1 |
1 |
1 |
1 |
0 |
0 |
0 |
0 |
|
Ptychochromis inornatus
|
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
0 |
0 |
0 |
0 |
|
Ptychochromis grandidieri
|
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
0 |
0 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
0 |
0 |
|
Ptychochromis makira
|
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
0 |
0 |
0 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
|
Ptychochromis loisellei
|
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
0 |
0 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
0 |
0 |
|
Ptychochromis curvidens
|
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
0 |
0 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
0 |
0 |
|
Ptychochromis ernestmagnusi
|
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
0 |
0 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
FIGURE 7.
Lateral pigmentation pattern in preservation for: A)
Ptychochromis makira
(AMNH 237131, holotype, 151.0 mm SL, adult male), and B)
P. ernestmagnusi
(AMNH 249488, paratype, 99.5 mm SL, adult). Dashed lines indicate relative adult lateral barring pattern and orientation.
Among the species of
Ptychochromis
that exhibit an “eastern
type
palatine” configuration,
P. m a k i r a
shares with
P. ernestmagnusi
the possession of supraneurals with a characteristically flattened dorsal profile (
Fig. 3
A, B, character 23). Also shared with
P. makira
is the presence of a characteristic lateral barring pattern, in which each prominent midlateral blotch is intersected by a pair of narrow and less strongly pigmented vertical or oblique bars (
Fig. 7
; character 24). However, in
P. m a k i r a
these pairs of lateral flank bars are distinctively V-shaped (
Fig. 7
A), whereas in
P. ernestmagnusi
the bars are vertically oriented (
Fig. 7
B). Despite inconclusive results in our earlier phylogenetic analysis (
Sparks & Stiassny, 2006:
Fig. 2
), in which
P. makira
was recovered in a polytomy with other “eastern group” species of
Ptychochromis
, our new analysis recovered a sister group relationship between
P. m a k i r a
and
P. ernestmagnusi
supported by the two unambiguously optimized anatomical features (characters 23 & 24) discussed above.
Although similar in many respects,
P. ernestmagnusi
is readily distinguished from
P. m a k i r a
by conspicuous iridescent spangling on the flank and dorsal fin membrane near its base, dusky grayish-green base coloration (vs. whitish), four lachrymal laterosensory foramina (vs. three), and a total of seven (vs. six) infraorbital bones. Although only two specimens of
P. m a k i r a
have been collected to date and admittedly represent a rather limited size range, the new species is also more shallow bodied (41.7–46.1 vs. 48.1–48.9% SL in
P. m a k i r a
), has a larger eye (29.5–37.4 vs. 25.6–27.6% HL in
P. m a k i r a
), a longer head (34.2–37.1 vs. 32.0–32.2% SL in
P. m a k i r a
), a shorter preanal length (71.6–73.5 vs. 75.1% SL in
P. m a k i r a
), and a greater lateral line scale count (34–37 vs.
33 in
P. makira
) than its sister taxon,
P. m a k i r a
.
FIGURE 8.
Strict consensus cladogram of nine optimal topologies recovered (tree length = 1018, CI = 0.67, RI = 0.61) based on the simultaneous analysis of the nucleotide dataset of Stiassny and Sparks (2006) and 24 morphological transformations. Unambiguously optimized morphological features supporting recovered nodes are designated by solid (unique feature) and open (homoplasious feature) circles. Character numbers (above braches) and states (below branches) correspond to the morphological transformations listed in Table 2 and Appendix 1. Indices in bold at nodes indicate jackknife support. Character state descriptions and statistics are listed in Appendix 1. The morphological character matrix is presented in Table 2.
Comparative materials.
Values following catalog numbers indicate count of specimens examined, and do not necessarily correspond to the total number of individuals in that particular lot:
Katria katria
:
AMNH
217739,
holotype
, eastern
Madagascar
, Tamatave Province, River Nosivolo, below Zule’s Village, large side-pool off mainstream.
AMNH
93701, 20 ex., 10 ex. C&S, eastern
Madagascar
, Tamatave Province, River Nosivolo below Ampasimaniona Village,
26 km
east-northeast of Marolambo.
UMMZ
240358
, 1 ex. C&S, eastern
Madagascar
, Marolambo.
Oxylapia polli
: AMNH 97111, 10 ex., 1 ex. C&S, eastern
Madagascar
, Tamatave Province, Mangoro drainage, village of Marolambo, Nosivolo River. AMNH 97150, 4 ex., 1 ex. C&S, eastern
Madagascar
, Tamatave Province, River Nosivolo by Ambarimasina Village, ca.
16 km
east northeast of Marolambo. UMMZ 235046, 1 ex. C&S, eastern
Madagascar
, Tamatave Province, Nosivolo River, near village of Marolambo, Mangoro drainage.
Paratilapia polleni
: AMNH 216068, 25 ex., eastern
Madagascar
, large baylake, behind dunes
1 km
south of turnoff from Marolambo-Mananjary road, ca. 100 meters from sea. UMMZ 235043, 2 ex. C&S, northeastern
Madagascar
, Lac Anjavibe,
Nosy
be. UMMZ 235045, 2 ex. C&S, southeastern
Madagascar
, Sahapindra River, near Vevembe.
Ptychochromis onilahy
: MNHN 1962-0201,
holotype
, southwestern
Madagascar
, Province of Tulear,
Onilahy
River, A. Kiener. AMNH 237130,
paratype
, 1 ex. (C&S in part), data as for
holotype
. MNHN 2006- 0 780,
paratypes
, 3 ex., data as for
holotype
.
Ptychochromis makira
: AMNH 237131,
holotype
, northeastern
Madagascar
, Antalaha Province, north of Maroansetra, near town of Marovonona, Antainambalana River, purchased from local fishermen by Augustin Sarovy, J. S. Sparks, W. L. Smith, and K. L. Tang. AMNH 237132,
paratype
, 1 ex. (C&S in part), data as for
holotype
.
Ptychochromis loisellei
: AMNH 232462,
holotype
, male, northeastern
Madagascar
, Antalaha Province, north of Sambava, Mahanara River at Antsirabe-Nord, just upstream of bridge over route N-5 (
13° 58.49′S
;
49° 57.81′E
), PVL-01-29, P.V. Loiselle and local fishermen. AMNH
231249
,
paratype
, 1 ex., northeastern
Madagascar
, Antalaha Province, main channel of the Mahanara River at Antsirabe-Nord, at bridge on Route N-5 (
13° 38.49′S
49° 57.81′E
), PVL-00-07, P.V. Loiselle. AMNH
231258
,
paratypes
, 3 ex., 1 ex. C&S, northeastern
Madagascar
, Antalaha Province, main channel of the Mahanara River at Antsirsabe-Nord, at bridge on Route N-5 (
13° 58.49′S
49° 57.81′E
), PVL-00-12, P.V. Loiselle. MNHN 2006-0781,
paratype
, 1 ex., data as for AMNH
231258
. AMNH 232458,
paratype
, 1 ex., northeastern
Madagascar
, Antalaha Province, Mahanara River, ca.
4 km
northwest of Antsirabe-Nord (
13° 57.30′S
49° 56.20′E
), PVL-01-27, P.V. Loiselle and local fishermen. MHNG 2676.095,
paratype
, 1 ex., data as for AMNH 232458. AMNH 237135,
paratypes
, 5 ex., data as for
holotype
.
Ptychochromis curvidens
: MHNG 2623.82,
holotype
, northern
Madagascar
, Antsiranana (Diego Suarez) Province, Andranofanjava, Andranofanjava-Sandriapiana River system, P. de Rham and J.-C. Nourissat. MHNG 2676.096,
paratypes
, 2 ex., data as for
holotype
. AMNH 237133,
paratypes
, 2 ex., 1 ex. C&S, data as for
holotype
. MHNG 2623.84,
paratype
, 1 ex., northern
Madagascar
, Antsiranana (Diego Suarez) Province, Mirosolava, P. de Rham and J.-C. Nourissat.
Ptychochromis insolitus
: UMMZ 237066,
holotype
, juvenile; northeastern
Madagascar
, Antalaha Province, near town of Mandritsara, Sofia drainage basin, Amboaboa (= Ambomboa) River (15° 50′ 1″S; 48° 42′ 51″E), J. S. Sparks and K. J. Riseng.
Ptychochromis inornatus
: UMMZ 237492,
holotype
, adult female, northwestern
Madagascar
, Antalaha Province, northeast of Antsohihy, Ankofia drainage, Anjingo River (14º 50′ 41.0″S, 48º 14′
38.3E
), JSS 94- 19, J. S. Sparks, K. J. Riseng, and local
Malagasy
guides. UMMZ 237063,
paratypes
, 5 ex., 1 ex. C&S, data as for
holotype
. AMNH
230746
,
paratypes
, 2 ex., data as for
holotype
. UMMZ 237064,
paratypes
, 5 ex., 2 ex. C&S, northwestern
Madagascar
, Antalaha Province, northeast of Antsohihy, Ankofia drainage, Bora Special Reserve, Bemahavony River (tributary of Anjingo River) (14º 52′ 20.4″S, 48º 14′ 52.2″E), JSS 94-20. AMNH
230747
,
paratypes
, 2 ex., data as for UMMZ 237064. UMMZ 237065,
paratype
, 1 ex., northwestern
Madagascar
, Antalaha Province, northeast of Antsohihy, Ankofia drainage, Lake Andrapongy (14º 41′ 49.3″S, 48º 0 7′ 54.3″E), JSS 94-21. UMMZ 237067,
paratypes
, 7 ex., northwestern
Madagascar
, Antalaha Province, northeast of Antsohihy, Ankofia drainage, Anjingo River (14º 50′ 39.7″S, 48º 14′ 39.5″E), JSS 94-54.
Ptychochromis grandidieri
:
MNHN
A.4147,
holotype
,
Madagascar
, region of high forests, Humblot and Grandidier. See discussion in Sparks (2003) regarding the locality of the
holotype
. Additional Non-Type Material Examined:
MNHN
A.310, 1 ex., rivers that cross the eastern slope, Lantz.
AMNH
88018, 56 ex., 1 ex., C&S, southeastern
Madagascar
, Mananjary, estuary of Mananjary River,
21º05′S
48º27′E
.
AMNH
88053, 2 ex., southeastern
Madagascar
, Mananjary, estuary of Mananjary River, 21º
0 5′S
,
48º 27′E
.
AMNH
88076, 2 ex., eastern
Madagascar
, Vatomandry,
19º 20′S
, 49º
0 0′E
.
AMNH
88090, 3 ex., eastern
Madagascar
, Mahanoro,
19º 55′S
,
48º 50′E
.
AMNH
88092, 17 ex., eastern
Madagascar
, Mahanoro, Pangalanes canal north of Mangoro River.
AMNH
88102, 36 ex., 14 ex. C&S, eastern
Madagascar
, Baylake behind dunes, ca.
100 m
from sea.
AMNH
88117, 18 ex., eastern
Madagascar
, Tamatave market,
18º 10′S
,
49º 25′E
.
AMNH
88140, 1 ex., eastern
Madagascar
,
25 km
north of Tamatave, Pangalanes canal, 18º
0 0′S
,
49º 25′E
.
AMNH
88153, 11 ex., eastern
Madagascar
, between Fenerive and Tamatave, Pangalanes canal, 18º
0 0′S
,
49º 25′E
.
AMNH
96999, 52 ex., 2 ex. C&S, eastern
Madagascar
, Tamatave Province, Mangoro River near mouth.
AMNH
97008, 185 ex., eastern
Madagascar
, Salehy village,
1 km
south of turnoff from Marolambo-Mananjary road,
19º 55′S
,
48º50′E
.
AMNH
97012, 12 ex., eastern
Madagascar
, Mahanoro market.
AMNH
97028, 7 ex., 3 ex. C&S, eastern
Madagascar
, Tamatave Province, Bay Lake behind first dune ca.
100 m
from sea, east of road by Sahey Village,
1 km
south of turnoff from Marolambo-Manajary Road.
AMNH
97057, 1 ex., eastern
Madagascar
, Ambodisovoka village, Savalany River.
AMNH
228067, 3 ex., southeastern
Madagascar
, Lopary, Mananizo River.
AMNH
228072, 6 ex., southeastern
Madagascar
, Ampataka village, Sahambavy River.
AMNH
228074, 3 ex., southeastern
Madagascar
,
12 km
north of Farafangana, Manampatrona River, 22º 43′ 47″S, 47º 47′ 25″E.
AMNH
231347, 1 ex., southeastern
Madagascar
, Ampataka village, Sahambavy River,
23º 21’ 04”S
, 47º28′18″E.
AMNH
231352, 4 ex., southeastern
Madagascar
, Manombo Special Reserve, Takoandra River, 23º 0 1′ 27″S, 47º 43′ 16″E.
MNHN
A.7896, 2 ex., central
Madagascar
to the west of Antananarivo, Lac
Itasy
(?).
MNHN
1901-0020, 1 ex., eastern
Madagascar
, Tamatave.
MNHN
1901-0021, 1 ex., eastern
Madagascar
, Tamatave.
MNHN
1932-0082, 1 ex., eastern
Madagascar
, Manompana.
MNHN
1932-0083, 13 ex., eastern
Madagascar
, Manompana.
UMMZ
233524, 17 ex., 2 ex. C&S,
Madagascar
, southeastern coastal region.
UMMZ
237311, 22 ex., southeastern
Madagascar
, Mananjary.
UMMZ
237312, 3 ex., 1 ex. C&S, southeastern
Madagascar
, Manombo Special Reserve.
UMMZ
237495, 5 ex., southeastern
Madagascar
,
6 km
north of Karianga at Mahavelo, Rienana drainage, Andriambondro River, 22º 21′ 47″S, 47º 22′ 0 5″E.
UMMZ
238453, 2 ex., southeastern
Madagascar
, near Manombo Special Reserve.
UMMZ
238471, 1 ex., southeastern
Madagascar
, Mananjary port.
UMMZ
238472, 11 ex., 2 ex. C&S, southeastern
Madagascar
, Farafangana market.
UMMZ
238476, 7 ex.,
Madagascar
, southeastern coastal region.
Ptychochromis oligacanthus
:
RMNH
3.936,
lectotype
, “
Madagascar
, in flumine Samberano, Nossibé, in lacu Pambilao”, Pollen and van Dam. Additional Non-Type Material Examined:
AMNH
18841, 1 ex.,
Madagascar
(probably mainland, Sambirano region).
AMNH
58491, 9 ex., northwestern
Madagascar
, Lake Amparihibe, at the mouth of the inflowing small stream, Lake Antsidihy,
Nosy
Be.
AMNH
215522, 4 ex., northwestern
Madagascar
, Lake Bemapaza,
Nosy
Be.
AMNH
215523, 15 ex., northwestern
Madagascar
, Lakes Djabala and Ampombilava,
Nosy
Be.
AMNH
230699
, 3 ex., northwestern
Madagascar
, Lake Andjavibe,
Nosy
Be.
AMNH
232399, 2 ex., northwestern
Madagascar
, Lake Ampombilava,
Nosy
Be.
AMNH
232415, 3 ex., northwestern
Madagascar
, Lake Djabala,
Nosy
Be.
MNHN
1962-322, 1 ex., northwestern
Madagascar
, Sambirano River.
UMMZ
236591, 26 ex., 4 ex. C&S, northwestern
Madagascar
, Lake Ampombilava,
Nosy
Be.
UMMZ
237498, 22 ex., 2 ex. C&S, northwestern
Madagascar
, Lake Djabala,
Nosy
Be.
UMMZ
237493, 3 ex., northwestern
Madagascar
, Lac de Deux Soeurs,
Nosy
Be.
UMMZ
237494, 1 ex., northwestern
Madagascar
, Lake Amparihibe,
Nosy
Be.
UMMZ
237496, 6 ex., northwestern
Madagascar
, Lake Bempazava,
Nosy
Be.
UMMZ
237497, 8 ex., northwestern
Madagascar
, Lake Anjavibe, island of
Nosy
Be.
UMMZ
237499, 11 ex., 1 ex. C&S, northwestern
Madagascar
, Mananjeba drainage, Andranomaloto River, northeast of town of Ambanja.
Ptychochromoides betsileanus
:
BMNH
1882.2.25:69,
lectotype
, Betsileo,
Madagascar
.
BMNH
1882.2.25:70,
paralectotype
, Betsileo,
Madagascar
.
AMNH
217753, 1 ex., southwestern
Madagascar
, Ilanana River, near Ranohira,
Onilahy
drainage.
AMNH
217763, 1 ex., south-central
Madagascar
, Manantanana River, headwaters near Iaritsena, Ambalavao Region, Mangoky drainage.
UMMZ
238115, 5 ex., dry skeletons, south-central
Madagascar
Ilanana River, south of Isalo National Park.
Ptychochromoides itasy
: AMNH 2336643,
paratype
, 1 ex.,
Madagascar
. MNHN 1919-11,
paratype
, 1 ex. C&S,
Madagascar
, central highlands, Region of Antananarivo, Lake
Itasy
.
Ptychochromoides vondrozo
: AMNH 228488,
paratypes
, 2 ex., southeastern
Madagascar
, Fianarantsoa Province, Region of
Vondrozo
, near Village of Vevembe, Mananara (du sud) drainage. UMMZ 235294,
paratypes
, 3 ex., 1 ex. C&S, southeastern
Madagascar
, Fianarantsoa Province, Region of
Vondrozo
, near Village of Vevembe, Mananara (du sud) drainage.